Seasonal Fluctuations in the Leaf-Footed Cactus Bug and Promoting Science Communication in the Undergraduate Classroom

BIG CHANGES AND BUG TALK: SEASONAL FLUCTUATIONS IN THE LEAF-FOOTED CACTUS BUG AND PROMOTING SCIENCE COMMUNICATION IN THE UNDERGRADUATE CLASSROOM

LAUREN ANNE CIRINO

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

2016

To my parents who always told me that you can do anything that you put your mind to

ACKNOWLEDGMENTS

I would like to thank my family and friends for their constant support throughout the entirety of this degree. Without their love and encouragement, this degree would have been much more arduous. I would especially like to thank my best friend, Deanna Colton, for her comments on my work, moral support, and going through graduate school first, so I could learn from her.

I would also like to thank Dr. Christine Miller for taking a chance on a high school teacher, who just wanted to get back into science. If it were not for Christine’s dedication to my progress as a scientist and the remarkable mentorship that she provided, this degree would not have happened.

I would like to thank all of the members of the Miller lab including Paul Joseph, Pablo

Allen, Zachary Emberts, Ummat Somjee, Michael Forthman, Paige Carlson, Jasmine Johnson, and Grayson McWhorter for their help and support with both with the execution of these two studies and the writing process. Lastly, I thank Brian Cobble for helping me locate a new field site where the cactus and cactus bugs were plentiful and Travis Tuten for allowing me to use this site for my projects. I am truly grateful for all of your help.

TABLE OF CONTENTS

page

LIST OF TABLES ...... 7

LIST OF FIGURES ...... 8

ABSTRACT ...... 9

CHAPTER

1 INTRODUCTION ...... 11

Plant-Insect Seasonality ...... 11 Science Communication Education ...... 13

2 SEASONAL EFFECTS ON THE POPULATION, MORPHOLOGY, AND REPRODUCTIVE BEHAVIOR OF NARNIA FEMORATA (HEMIPTERA: COREIDAE) ...... 16

Background ...... 16 Methods ...... 19 Study Species Behavior ...... 19 Study Sites ...... 19 Cactus Patch Sampling Study ...... 20 Insect Sampling Study ...... 20 Female Readiness to Mate ...... 21 Beak Plasticity ...... 22 General Statistical Analyses ...... 23 Results...... 24 How Does O. mesacantha lata Phenology Fluctuate Seasonally and Spatially? ...... 24 How Does the Number of N. femorata Change throughout the Seasons and Over Space? ...... 24 How Does Seasonality and Spatiality Affect Sexual Dimorphism of N. femorata? ...... 25 Does Seasonality and Spatiality Affect Female Readiness to Mate in N. femorata? ...... 25 Does Seasonality and Spatiality Affect Beak Length of N. femorata? ...... 25 Discussion ...... 26

3 BROADENING THE VOICE OF SCIENCE: PROMOTING SCIENTIFIC COMMUNICATION IN THE UNDERGRADUATE CLASSROOM ...... 39

Background ...... 39 Course Design and Student Recruitment ...... 41 The Research Talk for Peers ...... 42 The One-Minute Research Monologue for the Public ...... 42 The Research Poster for the Scientific Community ...... 42 Methods for the Assessment of Learning Gains ...... 43

Benefits of Science Communication Education ...... 44 Benefits to Undergraduates ...... 44 Benefits to Graduate Students ...... 45 Discussion ...... 45

4 CONCLUSION...... 51

Plant-Insect Seasonality ...... 51 Science Communication Education ...... 53

LIST OF REFERENCES ...... 54

BIOGRAPHICAL SKETCH ...... 61

LIST OF TABLES

Table page

3-1 Course Elements for the CURE course ...... 48

LIST OF FIGURES

Figure page

2-1 Aerial and understory photos of both Protected and Agricultural Sites ...... 33

2-2 Fruit abundance from the cactus patch study in the Protected Site throughout one full year and in the Agricultural Site for the last four months of the study ...... 34

2-3 N. femorata abundance from the insect sampling study for both Protected and Agricultural Sites ...... 35

2-4 Population level sexual dimorphism index (SDI) of two traits, pronotum width (PW) and hind femora width (HFW), at both the Protected and Agricultural Sites ...... 36

2-5 Pronotum width (PW) and hind femora width (HFW) averages (±SE) used in the calculations of SDI (Figure 2-4) compared temporally and spatially ...... 37

2-6 Scaling relationship of beak length of adult N. femorata for both A) females and B) males across four months in 2015 ...... 38

3-1 Applying science communication curriculum to an undergraduate classroom ...... 47

3-2 Self-rated post course gains to skills and abilities reported by students...... 49

3-3 Self-rated benefits to development reported by students after the course was completed ...... 50

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

BIG CHANGES AND BUG TALK: SEASONAL FLUCTUATIONS IN THE LEAF-FOOTED CACTUS BUG AND PROMOTING SCIENCE COMMUNICATION IN THE UNDERGRADUATE CLASSROOM

Lauren Anne Cirino

August 2016

Chair: Christine Miller Major: Entomology and Nematology

Many insects are influenced by the phenology of their host plants. My first objective here was to conduct an observational field study to examine seasonal changes in prickly pear cactus

(Opuntia mesacantha lata) and simultaneously track population and phenotypic changes in the leaf-footed cactus bug, Narnia femorata (Hemiptera: Coreidae). In north central Florida, O. mesacantha lata (formerly referred to as O. humifusa) begins producing flower buds in April that turn into green fruit and ripen into red fruit through December. I estimated changes in N. femorata abundance, morphology, and female readiness to mate for two field sites over time. I found that these insects were more abundant in the summer and fall as compared to other seasons. Intriguingly, I found a change in sexual dimorphism over space and time; male weapons

(hind femora) increased in size during the time of year when fruit were numerous. I did not find evidence that female readiness to mate varied across the summer and fall months. The field site with the most fruit had larger insects and insects appeared more numerous; yet, both of my sampled populations followed a similar pattern of seasonal fluctuation in body size and insect abundance.

In my second study, I illustrate one way in which science communication training was delivered through a Classroom Undergraduate Research Experience (CURE) course while students conducted novel research. Effective and accurate communication of scientific findings is essential. Unfortunately, scientists are not always well trained in how to best communicate their scientific findings with other scientists nor the importance of seeking out opportunities to speak with the public. With misinformation about science now flowing freely throughout the internet, it is more imperative than ever that scientists are clear, concise, and engaging in their science communication. Here, I argue that the development of science communication skills can and should be woven into the process of science from the very beginning. I describe my experiences developing, running, and evaluating a course for seven undergraduates involved in laboratory research. Four graduate students and a PI from a single laboratory created this course together. Our course design involved active learning about the nature of science and how science is used and interpreted in our society. Students delivered oral presentations, poster presentations, and research dialogues about the authentic research in which they were involved. We evaluated the effectiveness of our approach using the established CURE survey and a focus group. As expected, undergraduates reported strong benefits to communications skills and confidence while the graduate students involved in teaching the course gained beneficial experience in course creation. I provide guidance for those students and faculty interested in motivating and equipping the next generation of scientists to be excellent science communicators.

CHAPTER 1 INTRODUCTION

Plant-Insect Seasonality

Insect herbivores are tightly linked to the plant resource in which they feed. Many insects are polyphagous, meaning they can feed on many different plant genera to acquire the nutrition they need at different times of the year. However, some are monophagous (i.e., feed on a single plant genus), and when the plant resource changes over the seasons, so too can their abundance and phenotypic traits.

Coreidae (Hemiptera: Heteroptera) is a family categorized as primarily phytophagous insects that feed on angiosperms and prefer dicots over monocots (Mitchell 2000, Schaefer and

Mitchell 1983). Those that are monophagous are affected by the phenology of their plant resource (Schaefer and Mitchell 1983, Miller and Emlen 2010a). Narnia femorata Stål 1962

(Hemiptera: Coreidae) is one such monophagous species (Hunter et al. 1912, Mann 1969). They feed on the fruit and pads of the prickly pear cactus, Opuntia spp. (Hunter et al. 1912, Mann

1969). The phenology of Opuntia changes seasonally (Miller et al. 2006, Miller 2008, Gillespie et al. 2014) and at different times of the year these plants can grow ripe fruit, unripe fruit, or no cactus fruit at all (González-Espinosa and Quintana-Ascencio 1986; Hellgren 1994; Gillespie et al. 2014). Specifically, N. femorata feeds on Opuntia mesacantha lata (Small) Majure 2014

(formerly referred to as O. humifusa) in north central Florida (Baranowski and Slater 1986,

Nageon De Lestang and Miller 2009). The fruit of O. mesacantha lata has been shown to be of higher quality for N. femorata than just the pads alone (Nageon De Lestang and Miller 2009,

Miller et al. 2013, Gillespie et al. 2014, Sasson et al. 2016). For example, more eggs are laid in the presence of ripe cactus fruit (Miller et al. 2013) and nymph survivorship is higher when ripe cactus fruit is available (C. W. Miller, unpublished data). N. femorata pass through five instars,

all of which feed on O. mesacantha lata fruit and pads except for the non-feeding first instar

(Baranowski and Slater 1986, Miller et al. 2006, Nageon De Lestang and Miller 2009, Vessels et al. 2013).

Many traits in N. femorata have been shown to be plastic when diet is altered in the laboratory (Proctor et al. 2012, Addesso et al. 2014, Gillespie et al. 2014, Sasson et al. 2016).

Male hind femora, which are used in male contests, and testes size have been shown to change as resource quality changes (Sasson et al. 2016). The hind legs are sexually dimorphic with the males having more incrassate hind femora than females (Proctor et al. 2012). The size of male hind femora are important because males will fight one another to acquire territory which may attract a female for mating (Procter et al. 2012). Leg kicks, squeezes, and full grappling are behaviors that males exhibit to acquire territory (Proctor et al. 2012). Smaller male hind legs may inhibit the chances of winning a high quality territory to attract mates (Proctor et al. 2012, Sasson et al. 2016).

When males of N. femorata attempt to mate with females, the females can reject these mating attempts by running away, kicking the male, or closing her genital plates (Gillespie et al.

2014, L. A. Cirino pers. observ.). Females are more likely to choose a mate that has been reared on a high quality diet (red fruit) and is also living on a high quality environment (red fruit cactus,

Addesso et al. 2014). N. femorata traits and abundance fluctuate based on diet, which are common throughout Coreidae (Mitchell 1980, Eberhard 1998, Miyatake 1993, Miller and Emlen

2010a). Thus, N. femorata is a perfect model for studying seasonal changes in abundance, morphology, and behavioral traits.

I first describe my observational study that examines the temporal and spatial abundance, morphology, and female readiness to mate in the leaf-footed cactus bug, N. femorata. I

concurrently investigated temporal and spatial changes in the phenology of this species food resource, O. mesacantha lata, in north central Florida. Since N. femorata is monophagous, seasonal changes in their traits and abundance may be dictated by the fluctuating cactus quality throughout the year. This study also begins to explore spatial differences between two populations of N. femorata and the abundance of O. mesacantha lata fruit. By understanding the relationship between N. femorata and their host plant, we can gain further insights into how natural changes in the quality of diet affects the ecology and phenotypes of wild insect populations.

Science Communication Education

Science communication is becoming more important than ever with the advent of easily accessible news and social media through many electronic devices. It is crucial for senior scientists to educate younger generations in effective science communication because of the misinformation that is readily available and easily accessible to the public. Even though some undergraduate students obtain a science degree from a four-year university, they are unable to communicate the importance or relevance of scientific research, and why research funding is essential (Brownell et al. 2013, Mercer-Mapstone and Kuchel 2015).

When faced with many other career demands, principal investigators (PI) often find their time extremely limited. However, accomplishing science communication education with a relatively low time investment is attainable. One way in which we can train young scientists, or undergraduate researchers, is through a Classrooom Undergraduate Research Experience

(CURE). A CURE class is a course that addresses a novel research question in which the answer is unknown to both students and instructors (Auchincloss et al. 2014). The course is designed around this question with specific objectives that address the nature of science and support the student researcher’s development as a scientist (Miller et al. 2013b). CURE courses are 13

embedded into a regular academic school year and combine course work with research for students who are first semester researchers (Miller et al. 2013b). Since scientific projects can take a great deal of time and effort, expanding your research team by implementing a CURE class would achieve both research and science communication education goals. Undergraduates may receive training in common laboratory practices, animal care, laboratory techniques, field work and sampling methods, and perhaps others depending on the type of research conducted.

However, using the CURE method, students will learn how to ask scientific questions. During class, they may discuss science ethics, science and the media, be trained in science communication, and learn how to read and write primary scientific literature. Further, incorporating undergraduate students into a research laboratory increases the number of students being trained and receiving the benefits of this effort.

In my second study, I provide a case study on training undergraduate students in effective science communication. I also guide PIs and any other interested parties in setting up a course designed to engage students in both research and science communication. I do so by instructing them in the contents of the CURE course that we implemented in order to train undergraduate students in science communication. These students simultaneously worked on independent projects with a graduate student mentor. Students prepared three separate science communication assignments for the class that addressed both public and professional audiences. Training in science ethics, science in the media, and the nature of science was also employed. Through the implementation of this type of course, many students are able to receive excellent training in science communication with relatively low amounts of additional work for the PI.

The following studies describe how seasonality affects N. femorata and their host plant,

O. mesacantha lata, as well as explores how undergraduate students can be involved in scientific

research with these insects while simultaneously receiving science communication training. My work provides foundational ecological work for N. femorata and also serves as an example of how to train undergraduate students in effective science communication while performing fundamental biological research.

CHAPTER 2 SEASONAL EFFECTS ON THE POPULATION, MORPHOLOGY, AND REPRODUCTIVE BEHAVIOR OF NARNIA FEMORATA (HEMIPTERA: COREIDAE)

Background

Plants typically change seasonally and so does the nutrition that they provide to herbivores (Natuhara 1983, Wojcik et al. 2008, Ribeiro et al. 2011, Miller et al. 2013a). As plant quality declines, herbivore mortality can increase (Van Schaik et al. 1993) and traits vital for reproduction can also be affected (Numata and Hidaka 1984, McLain et al. 1993, Sasson et al.

2016). Many insects temper such fluctuations in diet quality through migration (Dingle 1972,

Van Schaik et al. 1993), while others transition to another type of food plant (Beck and Reese

1976). However, certain insect populations do not migrate nor change their host plants across seasons. Instead, they cope with the phenological changes of their host.

Foundational work on insects including aphids (Dixon and Wellings 1982), forest caterpillars (Janzen 1988), leafrollers (Hunter and McNeil 1997), and apple maggot flies

(Mopper 1996) have revealed the many intriguing ways that insects have adapted to seasonal changes in host plants (Wellings et al. 1980, Awmack and Leather 2002). Recognizing the dynamic ecological relationships between animals and their environment has provided vast insights into evolutionary processes (Reznick and Endler 1982, Schluter 2001, Grant and Grant

2002). Our aim for this study was to investigate seasonal changes in a Hemipteran, the leaf- footed cactus bug Narnia femorata Stål (1962), while simultaneously tracking phenological changes in the host plant.

N. femorata ranges from the southern United States in California, New Mexico, Arizona,

Texas and Florida to Central America in Mexico and Costa Rica (Mann 1969, Brailovsky 1975,

Baranowski and Slater 1986, Brailovsky et al. 1994, Palomares-Pérez et al. 2012). N. femorata was most likely introduced to Florida on nursery stock in the 1960s (Baranowski and Slater

1986). According to Baranowski and Slater (1986), N. femorata had become established in

Florida in the 1980s by feeding on the prickly pear cactus, Opuntia mesacantha lata (Small)

Majure 2014 (formerly referred to as O. humifusa). N. femorata has since been combined with N. pallidicornis retaining the species name femorata (Gibson and Holdridge 1918).

N. femorata overwinters as adults at the base of O. mesacantha lata (Baranowki and

Slater 1986, Mitchell 2000). Upon the arrival of spring, in March or April, the overwintering adults will oviposit in rows on the spines of cactus or pine needles that have fallen near cacti

(Baranowski and Slater 1986, L.A. Cirino, pers. observ.). N. femorata are bivoltine in the southwest (Baranowski and Slater 1986, Miller et al. 2006, Vessels et al. 2013) and the first generation reaches maturity about two months after oviposition (Baranowski and Slater 1986,

Vessels et al. 2013). These Hemipterans develop in five characteristic instars starting with a non- feeding first instar that lasts 4-7 days (Hunter et al. 1912, Vessels et al. 2013). The following four instars last approximately one to two weeks each (Hunter et al. 1912, Vessels et al. 2013).

However, the duration is highly temperature dependent (Allen et al., unpublished data, L.A.

Cirino, pers. observ.).

The developmental environment has a large effect on the phenotypes of adults in the

Coreidae; nutrition during development, for example, can influence the size, shape, internal anatomy, attractiveness to conspecifics, and mating behaviors of individuals (Miller 2008, Miller and Emlen 2010b, Addesso et al. 2014, Gillespie et al. 2014, Somjee et al. 2015, Sasson et al.

2016). Although N. femorata feeds on red ripe fruit, green unripe fruit, flower blooms, and pads of Opuntia spp. (Hunter et al. 1912, Mann 1969), high quality red ripe fruit can produce larger insects that are more attractive to conspecifics (Addesso et al. 2014, Gillespie et al. 2014, Sasson et al. 2016). These insects feed through a tube-like beak that helps them gain access to the

nutrients deep within their host plants and functions through the use of an osmotic pump

(Baranowski and Slater 1986, Mitchell 2000).

In north central Florida, N. femorata specifically feeds on Opuntia mesacantha lata

(Small) Majure 2014 (formerly referred to as O. humifusa, Baranowski and Slater 1986, Nageon de Lestang and Miller 2009). O. mesacantha lata flowers in the late spring and fruit matures from green to red throughout the summer and into the fall. Fruit begins to decline after its peak in June (Gillespie et al. 2014). One reason for the decline is that N. femorata is not the only cactus fruit consumer. Deer, tortoises, birds, coyote, and rodents all compete for this valuable food (Gonzalez-Espinosa and Quintana-Ascencio 1986, Janzen 1986, Hellgren 1994, L.A. Cirino and C.W. Miller, pers. observ.). Variation in the abundance of these vertebrate consumers over space is expected to lead to differences in the number of cactus fruit available for N. femorata to consume. For this reason, cactus fruit should vary not only seasonally, but also spatially.

Based on existing and ongoing work in this coreid and others, I predicted that seasonal and spatial variation in their food should be associated with a host of phenotypic changes in the wild. I predicted that, as low quality unripe green fruit ripens into high quality red ripe fruit, there would be an increase in body size (Miller and Emlen 2010a, Miller and Emlen 2010b,

Gillespie et al. 2014, Miller et al., in press), an increase in the extent of sexual dimorphism

(Miller et al., in press), and an increase in the readiness of females to mate (Miller 2008,

Gillespie et al. 2014). I also predicted that mouthpart length would decrease over this time period

(Allen et al., in prep). Further, because availability of high quality food should be linked to population growth and size (e.g. Miyatake 1994), I expected to find more individuals during the months when cactus fruit was plentiful.

I established two distinct sites for our study that differed strikingly in land use. At these sites, I noted changes in cactus fruit phenology and fruit abundance over time. I visited these sites monthly, counting and measuring insects, and bringing insects back to the lab to test their readiness to mate. I see this study as a first step towards understanding how the differences in these plants over time and space relate to individual differences in N. femorata and the success of these populations.

Methods

Study Species Behavior

Male N. femorata grapple with one another using their hind legs to acquire territory - a cactus pad with ripe or unripe fruit (Proctor et al. 2012). Females fly among cactus patches to mate, feed, and oviposit. Once females are present, males will frequently mount and make genital contact with females (Gillespie et al. 2014). Females that are ready to mate will open up their genital plates and allow male genitalia to penetrate (Gillespie et al. 2014). Successful males will turn 180 degrees and may remain attached for up to several hours (Gillespie et al. 2014).

Females can refuse to mate in a variety of ways, including striking males with their legs, running away from males, pushing their abdominal tip down, and keeping their genital plates closed

(Gillespie et al. 2014). They can also dislodge males once they have started to mate (L. A.

Cirino, pers. observ.).

Study Sites

All of the survey work was completed in north central Florida where N. femorata have been present since the 1960s (Baranowski and Slater 1986). My primary site included a 41,338 square meter survey area and nearby cactus patches at the University of Florida Ordway-Swisher biological field station in Melrose, Florida. This area is used for teaching, research, and outreach

(Figure 2-1a). Fire management is one of the ecologically important practices carried out here. I

considered this site my “Protected Site.” In the last four months of my study, I found another field site that I included as my “Agricultural Site.” The Agricultural Site included a 14,385 square meter survey area and nearby cactus patches at Future Farmers of America (FFA) agricultural plot in Live Oak, Florida (Figure 2-1b). Although not open to the public, this site is used for educational purposes for high school students in Suwannee County. This area is a pine plantation; longleaf pine trees are grown and harvested at this site every 15-18 years. Prickly pear cactus, O. mesacantha lata, are also unintentionally grown. While both sites have sandy soil, the two sites are strikingly different in plant community composition and structure. The

Protected Site is comprised of a diverse mix of oaks, palmettos, pines, grasses, and herbaceous plants; whereas, the Agricultural Site has reduced diversity, permitting growth of only a few species beyond longleaf pine trees and prickly pear cactus.

Cactus Patch Sampling Study

Our objective for this sampling work was to estimate changes in the number and type of cactus fruit of O. mesacantha lata over time. Twenty cactus patches were marked in each site; these twenty locations were chosen based on the presence of N. femorata at the onset of the study. A cactus patch was defined as a group of cactus pads within 25cm of each other (Schooley and Wiens 2004). I revisited these cactus patches in the third week of each month. O. mesacantha lata fruit type (red or green) and abundance were noted. Green fruit was characterized as both the flowers blooms and the green colored fruit. These cactus patches were located nearby the plots where the insect sampling study was completed.

Insect Sampling Study

Our objective for this field sampling work was to roughly estimate the changes in the abundance and document the morphological changes of N. femorata adults and nymphs over an entire year. My survey area at the Protected Site was 41,338 square meters, and our survey area

at the Agricultural Site was 14,385 square meters. I sampled these larger areas to be able to achieve a more representative estimate of changes in N. femorata abundance than would be possible from counting insects on just the twenty patches used in the patch sampling work.

Together with an assistant, I haphazardly surveyed O. mesacantha lata cactus pads for N. femorata for 3.5 to 4 hours at each site. The overall total number of nymphs and adults found at each site were recorded each month. Adults were sexed and their pronotum width (PW) and hind femoral widths (HFW) were measured using Mitutoyo digital calipers (maximum accuracy 0.01 mm). PW has been shown to be an excellent proxy for body size for N. femorata in laboratory studies (Proctor et al. 2012, Gillespie et al. 2014). This work was completed in a day during the second week of each month. I later calculated the sexual dimorphism indexes (SDI) of the PW and HFW by dividing the average female trait size by the average male trait size per month per site and subtracting 1 (Figure 2-4, Lovich and Gibbons 1992).

Female Readiness to Mate

Female mate choice and male-male competition are affected by resource availability and quality in N. femorata in laboratory settings (Proctor et al. 2012, Addesso et al. 2014, Gillespie et al. 2014). My aim for this part of the study was to determine the extent to which female readiness to mate in our field populations changed over time, not including predicted changes due to age and mating status (Gadenne et al. 2016). To minimize potential effects of age and to control for mating status, I collected fifth instar nymphs and brought them back to the lab to complete development to reproductive maturity. Adults become reproductively mature between 10 and 21 days from adult eclosion (C.W. Miller, unpublished data). The effects of seasonality on behavioral differences should be dampened as a consequence of laboratory rearing in the final stage of development. Thus my results for female readiness to mate are conservative. I also

measured pronotum width to examine whether body size predicted readiness to mate for this group of insects.

Insect collection was completed in the months of July through October, when the insects were most abundant, and to ensure an adequate sample size. I standardized nymph care across all months; I kept nymphs individually in an environment composed of a deli cup, soil, a cactus pad with high quality red, ripe fruit, and they were stored in a greenhouse (14 hours light: 10 hours dark throughout the year). Once individuals became sexually mature adults, I randomly paired adult males and females. Tests of readiness to mate were performed by pairing a single male and a single female in a cup for three hours while a single observer documented behaviors. Single pairs of females and males on a cactus are common at the Protected Site, though not as common at the Agricultural Site where the population density is high (L.A. Cirino, pers. observ.).

These trials were carried out for 3 hours in an 80 degree Fahrenheit room that was fully lit with two sets of florescent lights: ceiling and low hanging. I determined female readiness to mate by whether or not the female allowed copulation upon male mounting. I excluded those pairs that did not attempt to mate from my analyses as my question focused on whether or not females mated upon attempts by males. After completion of the readiness to mate trials, I froze males and females, took pictures using a digital camera (Canon EOS 30D) attached to a dissecting microscope (Leica M165C), and took body measurements using ImageJ software

(v1.42d, Abràmoff et al. 2004). Prior to each set of pictures taken, I photographed a scale bar to ensure accuracy of the measurements.

Beak Plasticity

Unpublished work has suggested that beak (mouthpart) length may be plastic, growing longer or shorter depending upon fruit size and phenology. For this reason I used N. femorata from the readiness to mate trials to estimate the extent to which beak length varied spatially or 22

temporally. It is worth noting that the individuals used for these estimates spent part of their last developmental stage in a laboratory setting and on a standardized diet. For that reason, my estimates should be conservative – any differences over space or time should be muted versions of the true differences within the natural populations. I measured beak length from the frozen N. femorata using ImageJ software (v1.42d, Abràmoff et al. 2004) described previously.

General Statistical Analyses

I examined differences in female readiness to mate over space and time using generalized linear model with a binary response variable (mating yes/no following a mounting by a male) and logit-link function. Explanatory variables for this model were site and month.

I examined beak length differences across time by using an Analysis of Variance

(ANOVA), blocking by site, and separately for each sex. A post-hoc Tukey’s (HSD) test was used to identify months with different beak length values. This analysis examined changes in beak length irrespective of changes in body size. However, I expected beak length to vary positively with body size, as is the case for most morphological traits. Thus, I next examined changes in beak length relative to body size (PW) using a full factorial Analysis of Covariance

(ANCOVA). I included month, PW, and the interaction of month and PW as explanatory factors in my model. A statistically significant interaction of month and PW indicates differences in scaling slopes across months. When the interaction was not statistically significant, I removed it and reran the model. The resulting model tested for differences in scaling intercept, where individuals of the same body size have differences in mouthpart length. Both analyses were completed using IBM® SPSS® ver. 22.

Results

How Does O. mesacantha lata Phenology Fluctuate Seasonally and Spatially?

Anecdotally, I noticed that flower blooms started to form on O. mesacantha lata in the warmer months of April and May and were finished blooming by June (data not shown). Green fruit began to turn red and ripen in July, and gradually red ripe fruit became more numerous than green fruit (Figure 2-2). I found a small number of red fruit throughout the entire year (Figure 2-

2). The difference in fruit availability was striking across the seasons and across the two sites.

The Agricultural Site had more fruit available in July (~ 169%), August (~588%), September

(~1106%), and October (~1235%) than the Protected Site (Figure 2-2).

How Does the Number of N. femorata Change throughout the Seasons and Over Space?

Adults were present throughout the entire year, yet I detected low numbers during the winter months (Figure 2-3a). By May, the numbers began to increase and spike in August at the

Protected Site (Figure 2-3a). The Agricultural Site carried a higher number of adults throughout all months of the summer and into the fall relative to the Protected Site (Figure 2-3b). Unlike the adults, nymphs were absent in the winter months in the Protected Site (Figure 2-3c). Although I did not formally sample the Agricultural Site in the winter months (because this site was incorporated after the winter), I observed nymphs in the next year’s winter in both November and February. Nymphs began to appear in April in the Protected Site (Figure 2-3c&d). In the

Protected Site, I detected a peak in abundance twice: once in April and again in August (Figure

2-3c). Interestingly, I did not detect peaks in abundance, rather nymph numbers appeared to increase through the fall months (Figure 2-3d). The difference in numbers of adult N. femorata at the Agricultural Site versus the Protected Site were higher in July (~671%), September (~210%), and October (~850%), but lower in August (~ -3%). The difference between numbers of nymphs

of all developmental stages in the Agricultural Site versus the Protected Site were higher for July

(~71%), August (~61%), September (~183%), and October (~860%).

How Does Seasonality and Spatiality Affect Sexual Dimorphism of N. femorata?

I used the PW and HFW measurements from the field sampling work to estimate the level of sexual dimorphism across months. I found a consistent trend of female-biased sexual size dimorphism (body size) as well as male-biased sexual dimorphism in hind femora across every month of the year (Figure 2-4). The level of sexual dimorphism differed across my two field sites (Figure 2-4). Differences in both PW and HFW were apparent in the month of August, however, those differences decreased in September. Differences between male and female hind femora widths decreased in the Agricultural Site most drastically between July and August. Yet differences remained relatively unchanged in the subsequent months. The Protected Site had larger differences between male and female HFW in August than in September.

Does Seasonality and Spatiality Affect Female Readiness to Mate in N. femorata?

I did not detect a difference in female readiness to mate across months (binary logistic regression: Wald χ2 = 0.162, p = 0.687) nor across sites (Wald χ 2 = 0.327, p = 0.567). There was a trend of increasing female readiness to mate in the Agricultural Site from July through October

(77.8% - 14/18 in July, 84.6% - 22/26 in August, 90% - 18/20 in September, and 100% - 14/14 in

October); however, I did not find a trend of increasing female readiness to mate over time at the

Protected Site with female readiness to mate at 66.7% (12/18) in July, 100% (10/10) in August,

77.8% (14/18) in September, and 66.7% (8/12) in October.

Does Seasonality and Spatiality Affect Beak Length of N. femorata?

I found that beak length (BL) was sexually dimorphic (Figure 2-6). I did not find spatial differences in beak length between the Agricultural Site and the Protected Site (ANOVA for

♂PW: F1,104 = 0.488, p = 0.486 and for ♀PW: F1,116 = 0.842, p = 0.361; ANOVA for ♂BL: F1,104

= 0.204, p = 0.653 and for ♀BL: F1,116 = 0.696, p = 0.406). Thus, I combined the data for both field sites and analyzed it together, yet kept the sexes separate.

I found that the different monthly cohorts of females expressed different allometric slopes between beak length and body size (ANCOVA month x PW: F3,116 = 24.452, p < 0.0001; Figure

2-6). In particular, females in the month of September revealed more homogenous beak lengths across body sizes (Figure 2-6). However, the scaling slope between beak length and body size is not significantly different for males across months (ANCOVA month x PW: F3,104 = 0.632, p =

0.596). Thus, I removed the interaction and reran the model to test for differences in scaling intercept, and found no differences across months (ANCOVA month: F3,104 = 0.614, p = 0.607;

Figure 2-6). Thus, the allometric relationship between body size and beak length does not appear to change in slope or intercept for males across the months of this study.

Discussion

This study provides insight into the intimate relationship between cactus-feeding insects and their cactus host. The number of cactus fruit available peaked in the spring and decreased over time consistent with previous findings across Opuntia spp. (Lenzi and Orth 2012, Gillespie et al. 2014, Arba et al. 2015). I also found pronounced spatial differences in cactus fruit abundance where the Agricultural Site had over 100% more cactus fruit each month than the

Protected Site. This spatial difference is likely related to herbivore abundance at the two sites where I sampled. Many, but not all, features of N. femorata fluctuated over time and space, consistent with the changing phenology of O. mesacantha lata. Body size differences increased at the Protected Site over time, but remained relatively constant at the Agricultural Site. The extent of sexual dimorphism in hind femora (Figure 2-4) and mouthpart length (Figure 2-6) decreased over time at both field sites. Yet, I did not detect differences in female readiness to mate across time and space. 26

Similar to Gillespie et al. (2014), we saw an increase in fruit numbers from April through

June and then a decline from June through the end of the year. Temporal patterns were the same in both field sites; however, I found a tremendous difference in the amount of fruit between sites

(Figure 2-2). The Agricultural Site not only had more fruit overall, but it also had more red fruit than the Protected Site (Figure 2-2). This difference is likely important to N. femorata because red ripe fruit has been shown to be a higher quality resource over green fruit or cactus pads without fruit for this species (Nageon de Lestang and Miller 2009, Miller et al. 2013, Gillespie et al. 2014, Sasson et al. 2016).

I found that the index of sexual dimorphism changed across time and across field sites

(Miller et al., in press). Body size was always larger for females than males, supporting the reverse of Rensch’s rule as found in the majority of insect groups (Stillwell et al. 2010). Yet the degree of sexual size dimorphism increased between August and September in the Protected Site and remained relatively constant for all four months at the Agricultural Site (Figure 2-4). These patterns may reflect differences in food quality at the two sites, with more red ripe fruit at the

Agricultural Site. Existing work has shown that poor quality food can lead to smaller morphological differences between males and females (Miyatake 1997, Miller and Emlen 2010a,

Sasson et al. 2016, Miller et al., in press), especially with regards to sexually-selected traits

(Miyatake 1997, Miller and Emlen 2010a, Sasson et al. 2016, Miller et al., in press) such as the hind femora in this species (Sasson et al. 2016, Miller et al., in press). By having more red ripe fruit, the Agricultural Site may provide the required nutrition for a larger population to produce greater differences between sexes in hind femora and pronotum width than in the Protected Site.

However, we cannot forget that these populations are wild caught and factors such as density, temperature, predators, and others can also affect these traits. Changes in the degree of sexual

dimorphism may have numerous effects on sexual selection dynamics and evolution in natural populations, yet spatial and temporal changes in sexual dimorphism have been given insufficient attention (Cox and Calsbeek 2010, Stillwell et al. 2010, Puniamoorthy et al. 2012, Manicom et al. 2014, Miller et al., in press).

Based on previous laboratory data, I expected larger male hind femora size and larger female body sizes in N. femorata when they developed in a month where red fruit was more numerous (Gillespie et al. 2014, Sasson et al. 2016, Miller et al., in press). Although female body size met this expectation to a certain extent, male hind femora size did not support this red fruit- dependent hypothesis in this particular study. I found that larger hind femora were present at times of the year when the preceding months had plentiful green fruit and very little red fruit

(Figure 2-4). Nymphs develop in approximately two months from egg to adult (Hunter et al.

1912, Vessels et al. 2013, L. A. Cirino, pers. observ.). Thus, the two months prior to when the adults were surveyed were when these insects were developing and feeding on cactus. This suggests that sexual size dimorphism due to environmental conditions in the wild may be more complex than previously thought (Stillwell et al. 2010, Miller et al., in press) and other factors such as density, temperature, and more may be playing a part in sexual dimorphism.

I did not detect changes in female readiness to mate over the four months of data that I collected from the field. These findings are an intriguing contrast to the findings of Gillespie et al. (2014) where female readiness to mate in the laboratory was shown to be dependent on both the rearing environment and the current environmental conditions. Females are likely to be receptive to males when the males are reared on high quality fruit and retaining a territory of high quality food (Gillespie et al. 2014). Although I did not see a significant temporal change in female readiness to mate in this study, my data does show a trend of increased readiness to mate

from July through October in the Agricultural Site. To understand this discrepancy between sites and to get an idea of true female readiness to mate in the wild, subsequent work should be carried out for all months of the year by mimicking the natural conditions found in the field in the laboratory.

I found that the beak length did not change over space. However, beak length relative to body size changed over time. Most traits positively scale with body size. Thus, larger insects might be predicted to have longer body parts than smaller insects. However, my data reveals that the changing beak length over time is not simply due to a change in body size. The length of beaks relative to body size also change. As discussed above, patterns found in this study are likely to be a conservative and muted representation of what occurs for field populations (see methods section above).

Intriguingly, the changes in beak length relative to body size appear to correspond to changes in O. mesacantha lata phenology, a pattern also found in experimental laboratory studies (P.E. Allen, unpublished data). Since Opuntia spp. fruit does not go through an increase of sugars until the end of their development (Barbera et al. 1992), N. femorata may need longer beaks to reach a more sugar or nutrient-rich portion of fruit during early fruit development seasons. The fruit’s outer layer of tissue also decreases in size upon maturity, which may make it easier for N. femorata to access the nutrients within Opuntia fruit (Barbera et al. 1992). The rapid changes in beak length here suggest that phenotypic plasticity, not evolution, is responsible for the changes seen. Such plasticity could be an adaptation that allows the insects to make the most of their changing food.

Mouthparts appear particularly malleable over evolutionary time, and increasing data suggest that phenotypic plasticity in mouthpart size and shape may be common as well (Relyea

and Auld 2005, Gil et al. 2008, Stoler and Relyea 2013). Phenotypically plastic changes in mouthparts have been documented in other animals in the wild such as wood frogs and spotless starlings (Relyea and Auld 2005, Gil et al. 2008, Stoler and Relyea 2013). Plasticity in beak length in female N. femorata is one of the first documented times in the wild that a temporal shift in a plant resource has been documented to influence changes in insect mouthparts within one year (Thompson 1999). This study may provide evidence for how N. femorata has survived on different Opuntia spp. fruits across their range and over time. Not enough is known about this trait strictly in the wild, so we need to explore this more experimentally and in the field.

I expected an increase in N. femorata abundance during seasons where red ripe fruit is more numerous. According to previous studies, more eggs are laid when red ripe fruit is available (Miller et al. 2013) and offspring develop faster and mature more quickly (Nageon De

Lestang and Miller 2009, Gillespie et al. 2014). Hind femora, relative to body size, that males use in intraspecific competition have also been shown to be larger when red ripe fruit is accessible during development (Sasson et al. 2016). Based on this field study and laboratory work, having red ripe fruit available throughout the season can provide N. femorata with higher quality resources which can be exploited for reproductive and territorial purposes (Gillespie et al.

2014, Sasson et al. 2016, Miller et al., in press).

Spatial differences in O. mesacantha lata abundance between the two sites may have contributed to the differences between the populations of N. femorata. Other studies have shown that N. femorata responds to changes in the reproductive effort (e.g. meristems allocated for reproduction versus growth) in the tree cholla cactus, Cylindropuntia imbricata (Haw.) F. M.

Knuth 1935 (=O. imbricata [Haw.] D. C. 1828), with an increased or decreased population size depending on whether or not more fruit is available (Miller et al. 2006, Miller 2008). The results

I report here show a similar pattern. The amount of adults that were found in both the

Agricultural Site and the Protected Site rose and fell over time. The Protected Site had much smaller numbers of adults overall than the Agricultural Site, likely due to a low number of cactus fruit. When adult numbers declined, the adults were almost absent at the Protected Site (Figure

2-3a). Larger O. mesacantha lata fruit numbers at the Agricultural Site also appeared to support a larger and increasing population of nymphs; whereas, the decreasing number of cactus fruit at the Protected Site may be responsible for declining nymph numbers after August.

The Florida populations of N. femorata adults are present throughout the year as are the

New Mexico populations (Vessels et al. 2013). Yet N. femorata in Florida appears to be a multivoltine species compared to the New Mexico population which is bivoltine (Vessels et al.

2013). This may differ from the New Mexico population due to climate or habitat conditions in which they live (Vessels et al. 2013). A single species with a large geographic range may be under more drastic temporal changes with nymphs developing under more dynamic weather and resource quality conditions.

This study provides foundational information about N. femorata in Florida. N. femorata is proving to be a valuable laboratory study organism for investigating questions in the field of sexual selection, yet few details of its life history and relationship with its seasonally-variable host plant have been provided until now. I hope this work will serve as a catalyst for future studies of this species in the wild, including a more in-depth analysis of the ecological interplay between N. femorata and its host plant. Further, the results reported here suggest that the shape and size of insects may change in complex ways with host plant phenology, patterns that have rarely been investigated. The inclusion of an additional field site illustrates that these seasonally-

variable patterns are not consistent across space. Further studies should investigate how land use and competitive herbivory can affect N. femorata traits and demography.

A B

C D Figure 2-1. Aerial and understory photos of both Protected and Agricultural Sites. A) Boundary lines of the Protected Site C) with its corresponding understory photograph B) and the agricultural field site D) with its corresponding understory photograph where N. femorata and O. mesacantha lata were surveyed. Photo A courtesy of author Google Earth. Ordway-Swisher Biological Field Site aerial view. April 1, 2015. Melrose, FL. Source: https://www.google.com/maps. Photo B courtesy of author Google Earth. Future Farmers of America Agricultural Site aerial view. July 28, 2015. Melrose, FL. Source: https://www.google.com/maps. Photo C courtesy of author Pablo Allen. Ordway-Swisher Biological Field Site understory view. February 20, 2016. Live Oak, FL. Photo D courtesy of author Pablo Allen. Future Farmers of America Agricultural Site understory view. February 20, 2016. Live Oak, FL.

Green - Protected Red - Protected 1000 Green - Agricultural Red - Agricultural

800

600

400 Cactus fruit available fruit Cactus 200

2014 2015 Figure 2-2. Fruit abundance from the cactus patch study in the Protected Site throughout one full year and in the Agricultural Site for the last four months of the study.

A B

C D

Figure 2-3. N. femorata abundance from the insect sampling study for both Protected and Agricultural Sites. A-B) N. femorata abundance for both adults C-D) and nymphs in 2014-2015. A) Adults in the Protected and B) Agricultural sites and C) nymphs in the Protected and D) Agricultural sites with fruit abundance matched to the corresponding month.

PW - Agricultural HFW - Agricultural PW - Protected HFW - Protected 0.15 0.1 0.05 0 -0.05 -0.1 -0.15 -0.2

Sexual dimorphism index (SDI) index dimorphism Sexual -0.25 July (54) Aug (33/34) Sept (62/20) Oct (57)

Figure 2-4. Population level sexual dimorphism index (SDI) of two traits, pronotum width (PW) and hind femora width (HFW), at both the Protected and Agricultural Sites. If the number is below zero, males have the larger trait. If the number is above zero, females have the larger trait. The numbers in the parentheses are the sample size for each month (Agricultural/Protected). If the sample size was below 10, the data was excluded from the graph.

Protected Site Agricultural Site PW (mm) HFW (mm) PW (mm) HFW (mm) Males Females Males Females Males Females Males Females Jul 4.86±0.24 (2) 4.92±0.15 (5) 1.93±0.03 (2) 1.42±0.03 (5) 4.84±0.03 (23) 5.08±0.07 (31) 1.75±0.03 (23) 1.41±0.02 (31) Aug 4.65±0.04 (17) 4.72±0.05 (17) 1.59±0.03 (17) 1.31±0.02 (17) 4.40±0.07 (13) 4.74±0.04 (20) 1.43±0.04 (13) 1.29±0.02 (20) Sep 3.95±0.09 (11) 4.43±0.08 (9) 1.40±0.03 (11) 1.20±0.03 (9) 4.34±0.04 (30) 4.66±0.03 (32) 1.53±0.02 (30) 1.35±0.02 (32) Oct 3.87±0.08 (2) 4.45±0.04 (4) 1.41±0.10 (2) 1.31±0.03 (4) 4.38±0.03 (31) 4.63±0.04 (26) 1.50±0.02 (31) 1.31±0.02 (26)

Figure 2-5. Pronotum width (PW) and hind femora width (HFW) averages (±SE) used in the calculations of SDI (Figure 2-4) compared temporally and spatially. Sample sizes are in parentheses behind the average measurement.

[a] [a] [ab] [a] [b] [a]

[c] [a]

A B

Figure 2-6. Scaling relationship of beak length of adult N. femorata for both A) females and B) males across four months in 2015. Statistical significance from an ANCOVA blocked by site, separately for each sex, and a post-hoc Tukey (HSD) test is indicated by the letters within the brackets next to the graph legend.

CHAPTER 3 BROADENING THE VOICE OF SCIENCE: PROMOTING SCIENTIFIC COMMUNICATION IN THE UNDERGRADUATE CLASSROOM

Background

Antibiotics kill viruses (Collett et al. 1999), vaccines cause autism (Ruiz and Bell 2014), and global climate change is a conspiracy (van der Linden 2015). These, among many others, are huge misconceptions of scientific knowledge that are partially perpetuated by the fast flow of information through the internet. Disseminating inaccurate information has become a normal occurrence as avenues for information on the internet have expanded (Del Vicario et al. 2016).

Now, it is more imperative than ever for scientists of all levels to learn how to effectively communicate their science and to do so frequently through multiple mediums. The way in which scientific information is discussed is important for helping the lay public make decisions about their healthcare, the environment, and the quality of food that they consume. Of course, communication of scientific findings is also essential among scientists themselves to enable science to proceed in new and innovative directions.

In the past, science communication to public audiences was seen as largely the responsibility of the media (e.g., newspaper, television; Pew Research Center for the People & the Press 2008b). Intriguingly, the recent insurgence of online communication sources and social media, while problematic in some ways, has also made it easier for scientists to directly communicate with the public (Nisbet and Scheufele 2009). Simultaneously, federal agencies, such as the National Science Foundation, have developed new guidelines for grant proposals where researchers need to explain, from the beginning, how they will accomplish public outreach

(NSF 2016). Yet, most scientists have not been formally trained in how to effectively communicate their research to the public in a way that the public can easily understand (Besley et al. 2015).

Training future generations of scientists in effective communication using modern communication mediums is imperative. Often no formal training in science communication is available for undergraduates majoring solely in scientific fields. Thus, undergraduate students more commonly gain these skills by happenstance (e.g. they join a laboratory that makes science communication a priority) or by seeking formal science communication training independently.

High quality science communication training should be the norm, not the exception for young scientists. Emphasis should be placed on training young scientists how to effectively communicate research to different audiences, so that accurate dissemination of scientific information is shared with both public and the scientific community.

No matter who the target audience is, the ultimate goal of science communication remains the same: to accurately provide people with the latest information on science and to generate support for it (Pace et al. 2010, Kuehne et al. 2014, Besley et al. 2015). Learning how to navigate a poster presentation or a talk at a conference can be extremely beneficial to a young scientist looking to receive feedback with more senior members of the scientific community.

Further, these means of communication can be powerful in highlighting to other scientists new directions and can speed the process of scientific innovation (Osterhaus and Vanlangendonck

2015).

I propose that an important step to improving science literacy in students and, ultimately, in the public sphere, is through engaging undergraduate students in authentic research while simultaneously training them on how to communicate that science to multiple audiences. I describe my experiences developing, teaching, and evaluating a course focused on elevating science communication skills in advanced undergraduates. Together with a teaching team, we recruited undergraduates already involved in laboratory research with the goal of helping these

students become skillful in communicating science to scientists and the public while simultaneously thinking more deeply and critically about their research projects. This course incorporated elements relevant to the development of science communication skills, including materials and discussions about the processes of science, science ethics, and science in the media. Further, we revisited and reinforced student understanding of ecology and evolutionary biology to help them gain deeper context for their ongoing laboratory research. Activities specifically focused on science communication involved (1) the presentation of primary scientific literature to peers, (2) the development of a research monologue for the public, and (3) the construction and presentation of a technical poster based on their current undergraduate research. The time investment of the PI and four graduate students involved in teaching this course was low relative to the strong learning gains reported by the undergraduates involved. My main goal in this manuscript is to provide an example of how to invest in the development of a new generation of scientists with excellent science communication skills. I provide a description of our experiences, a report of student learning gains, and a practical guide for others to use when building one of these courses.

Course Design and Student Recruitment

The teaching team, a primary investigator and four graduate students, co-designed this course for undergraduate students already involved in authentic research in the fields of ecology, evolution, or behavior. We offered this course at the University of Florida (Gainesville, FL) in the summer of 2015 and recruited undergraduate students conducting research at the time. Unlike a traditional lecture-based science course, our course brought the authentic research results that undergraduate students were working on into the classroom. This real research experience provided the platform linking students’ authentic research to broader theoretical contexts and how science is represented in the public sphere. Further, we used students’ research as the 41

material for their science communication training (Box 1). We enrolled six students from the

PI’s laboratory and one from another evolutionary ecology lab. This class used a flipped classroom approach (see Lage et al. 2000) to address course elements while simultaneously supporting the undergraduates’ authentic research (Table 1).

The teaching team trained students in three forms of science communication: a research talk, a 1-minute research monologue, and a research poster. Throughout, we drew on their personal research experiences and included training in the nature of science (Lederman 2007).

The Research Talk for Peers

Students worked in pairs to generate a professional 8-minute PowerPoint to present the introduction, methods, results, and conclusion from a single manuscript published in the primary scientific literature. After the presentation, the students provided critical thinking questions in ecology, evolutionary biology, or behavior, as relevant to the manuscript being addressed.

The One-Minute Research Monologue for the Public

All of our students were already undergraduate members of research teams. Students were asked to draw on their knowledge of their ongoing research to prepare a 1-minute research monologue targeting a general audience. Students presented their monologues in a small group format where they received feedback from members of the research team and their peers. After revising their monologues, students were filmed giving their presentations. A subset of these videos were posted to the laboratory website (http://www.millerlab.net/) as a form of scientific outreach.

The Research Poster for the Scientific Community

Students worked with the teaching team to design and present a poster on their existing laboratory research projects for delivery to peer scientists. Students presented their posters in a

symposium style, where they rotated between presenting to their peers and serving as an engaging audience.

Methods for the Assessment of Learning Gains

We evaluated our course using quantitative and qualitative methods. Quantitatively, students were asked to complete the Classroom Undergraduate Research Experience (CURE) surveys (Lopatto 2008), providing us with self-reported course learning gains to quantify the course’s effectiveness (Anaya 1999).

All enrolled students anonymously and voluntarily completed the CURE survey at the beginning of the course (pre-course) and another survey after the conclusion of the course (post- course). The purpose of these surveys was to invite students to self-report their learning gains and rate the elements of the course on a five-point Likert scale (1 = smallest gain, 5 = largest gain). This course’s CURE survey results were then compared with data from over nine thousand previous student responses to this same survey. These students came from various different ethnicities, attended public or private institutions across the country, and ranged from high school through graduate students. In addition, those CURE responses that overlapped with the

Survey of Undergraduate Research Experiences (SURE) responses were compared (Lopatto

2004). The students surveyed by SURE represented 41 universities and colleges, made up of equal amounts of males and females with eight different ethnicity groupings, and who ranged from first year to third year students (SURE; Lopatto 2004). The seven students in our CURE survey were all Caucasian, approximately equal in number of males and females, and were in either their second or third year at the University of Florida.

We planned a focus group interview to provide a forum for qualitative course evaluation.

The anonymous focus group was moderated by an outside facilitator and included eight open- ended questions written before the class began. The focus group discussion was then transcribed 43

by a professional transcription service. The transcription was individually assigned open codes

(Holstein and Gubrium 2003) by two of the authors. After independently assigning open codes, the open codes were compared and consolidated. Upon consolidation, the open codes were placed into broader categories, which were used to help shape the discussion of the manuscript.

Benefits of Science Communication Education

Benefits to Undergraduates

Science communication skills. Students reported gains in two main areas: benefits to skills and abilities (Figure 3-1) and benefits to development as a scientist (Figure 3-2). Overall, our students reported high learning gains in the areas of communication in each category. The skills and abilities that were rated higher coincided with those required for science communication such as how to give an effective oral presentation (Figure 3-2). Additional non- target areas, such as reading and understanding primary scientific literature were also rated high

(Figure 3-2).

Confidence in delivering a science message. We found that teaching students science communication improved their confidence in speaking to various audiences. For example one student said “personally, I’ve never presented someone else’s research, let alone my own. So, even that little bit of experience was, I thought was, helpful… And I thought I developed my ability to present.” Another student remarked,

I think one of the best projects we’ve done in this course was the one-minute summary of our research that had to be for a public audience. And I think that getting practice, […] presenting your research without all of the technical terms […] can go a long way towards science being better communicated towards the public.

Students received science communication knowledge and constructive feedback directly from more senior researchers making their science message stronger and more impactful.

Students began to recognize that complicated science messages can be ignored or even devalued

by the public. We challenged students to bolster their critical thinking skills and build their confidence in oral communications (Figure 3-1). Our aim was to provide students with a foundation they can use to achieve excellence in science communication in the future.

Benefits to Graduate Students

Graduate students that co-taught the course also benefitted from teaching a CURE class.

They gained higher-level teaching and organizational skills that should raise their competitiveness for academic jobs. For example, this course provided the opportunity to build a syllabus while thinking critically about class objectives, an experience that was new for most instructors. Graduate students also designed and implemented lessons that catered towards their class objectives. Additionally, graduate students improved their own communication skills through teaching others (for more information Cicirelli 1972, Cortese 2005). Strong professional relationships between graduate students and undergraduates were developed through this course that would not be possible in large lectures. Importantly, the students and research team achieved a valuable end product from the class: research videos that were tailored to the public audience, helping with scientific outreach aims (http://www.millerlab.net/).

Discussion

The results from our interviews and surveys illustrate that we provided college students opportunities to develop valuable and foundational skills in communication: recognizing their audience, communicating science effectively to that audience, and evaluating the effectiveness of that message. Communication skills are among the top skills that employers are looking for in job candidates (Robles 2012). The foundational skills provided through this course should benefit students in many career paths. Future studies should follow students as they move into their careers to assess benefits, if any, of this early training in science communication.

Our goal was to make science communication fun, interesting, and achievable for young undergraduate scientists. We achieved this goal with relatively little effort through an abbreviated summer semester. Our learning evaluations illustrate that directing students to present their individual research projects in different ways to difference audiences encouraged an early recognition of some of the challenges, and opportunities, that scientists face. In addition, the course gave us an opportunity to interact in a new way with undergraduate researchers, providing a strong sense of unity and a team mentality.

Find the instructors and provide them an excellent opportunity for career development:

Identify graduate students, postdoctoral researchers, and even experienced undergraduates with similar commitments to excellence in science education. Designate specific meeting dates and times for collaboration prior to and throughout the class. Once your teaching team is assembled, collaborate to design and teach the course, assigning tasks by interest and skill level. By involving advanced students in this process, you will provide them with an excellent training opportunity in teaching.

Determine the science communication assignments: Identify two to three science communication assignments for the class based on the length of the class, the amount of time each assignment will take and the skill level of your students. Determine how each assignment will be taught and who will teach it. There should be other assignments in this class to support the science communication curriculum such as investigating specific research topics using the primary literature or discussing what it means to be a scientist.

Provide support for student science communication learning: Give students the opportunity to practice their science communication with each other and the teaching team. Provide constructive feedback before their final presentation of each science communication assignment.

Showcase undergraduate research: Have your undergraduate students present their research in a school-wide research forum or conference, video record your students presenting their research to the public, or have your students present their research to high school or middle school life science classes.

Figure 3-1. Applying science communication curriculum to an undergraduate classroom

Table 3-1. Course Elements for the CURE course Course Element Topic Training Location Foundational Topics  What is Science? Classroom and Materials  Evolutionary Ecology & Behavior  Science Ethics

Science  Scientific Literacy & Communication Classroom Communication  The Research Talk for Peers Training  The 1-minute Monologue for the Public  The Research Poster for the Scientific Community

Ongoing Authentic  Data analysis training Classroom and Research  Lab techniques training undergraduate student’s  Authentic research respective lab

5 Our course Other CURE courses 4.5

3.5 Mean Rating Mean

2.5

2 Critique work of Discuss reading Present posters Present results Work in small other students materials in class orally groups

Course Elements

Figure 3-2. Self-rated post course gains to skills and abilities reported by students. A rating of 5 is the highest gain and 1 is the lowest gain. The grey circles represent the students in the present study (Our course, n=7). For comparison, we show the overall mean gains reported by students (Other CURE courses, n>9000) who took the CURE survey in 2014-2015, and who used the same CURE survey (white triangles).

5 Our course Other similar courses Typical summer research course

4.5

4 Mean Rating Mean 3.5

2.5 Ability to analyze Ability to read Becoming part of Skill in how to Skill in data and other and understand a learning give an effective interpretation of information primary literature community oral presentation results

Course Elements

Figure 3-3. Self-rated benefits to development reported by students after the course was completed. A rating of 5 is the highest gain and 1 is the lowest gain. The grey circles represent the students in the present study (Our course, n=7). For comparison, we show both the learning gains reported by other students (Similar courses, white triangles) enrolled in classroom research experience (error bars represent 2 standard errors) and by students who completed a summer laboratory research experience in 2014 and completed the Survey of Undergraduate Research Experiences (Typical summer research course, white squares). Since questions on the learning gains surveys for these courses overlapped for self-rated benefits, we were able to compare them here.

CHAPTER 4 CONCLUSION

Plant-Insect Seasonality

Cactus abundance fluctuates seasonally across both space and time. As this fluctuation occurs, N. femorata abundance, morphology, and sexual size dimorphism also changes.

Although there are differences in body and hind femora sizes in wild populations of N. femorata, these differences do not appear precisely matched with O. mesacantha lata fruit abundance and phenology as predicted. This unanticipated result raises questions about the actual nutritional differences between late green fruit and red fruit: does the difference in color actually depict its nutritional value and if so, is this difference impacting the morphological traits of N. femorata differently?

Furthermore, sexual size dimorphism needs to be examined under the influence of simultaneous factors in a controlled setting. Studies on the sexual dimorphism of N. femorata under varying temperatures and densities are underway (Allen et al., in prep). These studies suggest that adult males reared in higher densities as juveniles had larger hind femora than those reared in lower densities (Allen et al., in prep). Males reared under high quality diet had larger hind femora than females regardless of temperature (Allen et al., in prep). However, the difference between male and female hind femora decreased under poor quality fruit conditions

(Allen et al., in prep). When reared on poor quality fruit and cooler temperatures, females, rather than males, had larger hind femora (Allen et al., in prep). These laboratory results show that many factors in the wild are contributing to the plasticity of sexually selected traits in N. femorata such as hind femora size. These three factors should be studied simultaneously for effects on sexually selected traits of N. femorata. Studying the effects of these abiotic and biotic

factors will help us understand how the complexity of the environment can influence the traits of

N. femorata.

Female readiness to mate in both the Agricultural Site and Protected Site did not significantly change temporally or spatially as expected. Early nymphal rearing conditions may not play a role in female mate choice. Rather, it may be the ultimate instar rearing conditions and context-dependence conditions that play a greater role in female readiness to mate (Gillespie et al. 2014). Increasing the sample size, collecting mating pairs throughout each season, and keeping laboratory conditions, including cactus type, similar to environmental conditions is necessary to understand the extent to which female choice fluctuates in the wild.

Intriguingly, we found evidence that female beak length gets shorter over the seasons as

O. mesacantha lata fruit ripens, but we did not find evidence that male beak length changes over time. Recent laboratory work has shown that beak size increases when reared on low quality fruit regardless of temperature and sex (P. E. Allen, unpublished data). However, beak length needs further examination in the field. A more comprehensive look at how beak length changes over one full year in one species, how it may change over space in a single species, and how it may differ between species is also needed to understand the plasticity of this trait. It would also be beneficial to study the phenotypic changes of the host plant while simultaneously studying beak length. This may provide some insights into why there is phenotypically plastic beak lengths in

N. femorata.

Finally, the abundance of N. femorata fluctuates with the change in fruit abundance. By the addition of the Agricultural Site into the end of the study, I was able to begin examining the effects of site on abundance and morphology of N. femorata. Spatially, there is more fruit at the

Agricultural Site than there is at the Protected Site, and as expected, we see a larger population

of adult and nymph N. femorata. Further research should be done to understand how agricultural practices can influence the population size of N. femorata by increasing the number of agricultural and protected sites sampled. Additionally, estimating the immigration/emigration rate and conducting predator-prey studies of these populations would be beneficial in understanding population fluctuations of N. femorata.

Science Communication Education

With the speed in which people receive news today, accurate and effective science communication is crucial. Undergraduate students benefited from a Classroom Undergraduate

Research Experience (CURE) class that combined independent research with science communication training. The class provided undergraduate students with foundational skills in recognizing different audiences, preparing communication for these audiences, and evaluating the effectiveness of that communication. Because this class utilized team-teaching, the relative work load was reduced, so achieving the goals of effective science communication training were attainable.

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BIOGRAPHICAL SKETCH

Lauren Anne Cirino was born in Alexandria, Virginia in 1984. She grew up and went to grade school in northern Virginia where she first became interested in science. It was her high school biology teacher who introduced her to the study of life including evolutionary biology.

Lauren knew from that point on that she wanted to focus her studies on the evolutionary theory.

Lauren attended Clemson University for her B.S. in biology. She worked on sexual selection of sailfin mollies in an evolutionary laboratory while there. Although still inspired by science,

Lauren decided to begin teaching high school science upon graduating from Clemson. She taught in south Florida for one year and then in northern Virginia for six years. This is where Lauren realized her passion and love of sharing her knowledge of science with others. She continues to share her passion of science as a graduate student where she incorporates and teaches undergraduate students the research process. Other than her passion for research, Lauren loves to play many team sports including soccer, field hockey, and volleyball. She received her Master of

Science degree from the University of Florida in the summer of 2016.