A Comparative Life History Study of the Swamp Darter, Etheostoma Fusiforme (Girard)

NORTH VERSUS SOUTH: A COMPARATIVE LIFE HISTORY STUDY OF THE SWAMP DARTER, ETHEOSTOMA FUSIFORME (GIRARD)

ADANIA FLEMMING

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

2018

To my husband Jason Fraser-Nash for his support and assistance, and to my family from Trinidad and Tobago my dad, David Flemming, mom, Audrey Greenidge and sisters Adonia and Adenia Flemming for their support and to my advisor Larry Page for his guidance.

ACKNOWLEDGMENTS

Gratitude must be expressed to Jason Fraser-Nash, William Fraser-Nash, Hannah Owen, and Cathleen Bester for their assistance in collecting specimens, to Danielle Diaz for her assistance in sorting and identifying invertebrates, to Kevin Love for his guidance with learning

R programming and technical support, to Zachary Randall, Jarred Randall, and Gustav Paulay for their assistance with photographing specimens, to Kirsten Work for E. f. barratti reproduction video footage, to Ash Bullard for identification of the digenean parasite, to my advisor Larry Page, and committee members Daryl Parkyn, Max Nickerson and Jimmy Liao for their guidance, to my lab mates John Pfeifer, David Boyd, Randy Singer, Rob Robins and collection mates Lindsay French, Samara Nehemia for their support. Distributional data for E. fusiforme were downloaded from iDigBio, supported by NSF EF 1115210 and DBI 1547229.

TABLE OF CONTENTS

page

ACKNOWLEDGMENTS ...... 4

LIST OF TABLES ...... 7

LIST OF FIGURES ...... 8

ABSTRACT ...... 9

CHAPTER

1 INTRODUCTION ...... 10

2 METHODS ...... 14

Field Site Description ...... 14 Field Collection ...... 14 Laboratory ...... 15 Diet ...... 16 Reproduction ...... 16 Development and Growth ...... 17 Aquarium Observations ...... 17 Statistics ...... 18

3 FIELD SITE DESCRIPTION (HABITAT ECOLOGY) ...... 22

4 RESULTS/DISCUSSION ...... 25

Reproduction ...... 25 Reproductive Cycle of the Male ...... 25 Reproductive Cycle of the Female ...... 26 Spawning ...... 27 Development and Growth ...... 28 Demography ...... 31 Territoriality ...... 31 Composition ...... 31 Diet and Feeding Habits or Diet and Parasitism ...... 31 Interactions with Other Organisms ...... 33 Predation ...... 33 Parasitism ...... 33 Conservation ...... 34 Aquarium Observations ...... 34

5 SUMMARY ...... 42

LIST OF REFERENCES ...... 44

BIOGRAPHICAL SKETCH ...... 46

LIST OF TABLES

Table page

4-1 Mean lengths, weights and ova counts of female Etheostoma. f. barratti collected each month during the reproductive period...... 35

4-2 Lengths and dates of the 20 largest specimens collected at Hatchet Creek...... 36

4-3 Mean Number of Food Organisms Per Stomach...... 37

5-1 Summary of life-history information on the Hatchet Creek population of E. f. barratti compared to a southern New England population of E. f. fusiforme (Schmidt and Whitworth 1979, Schmidt 1983)...... 43

LIST OF FIGURES

Figure page

1-1 Distributional data of the Swamp Darter, Etheostoma fusiforme...... 13

2-1 Standard lengths of all specimens collected over the study period...... 21

3-1 Image of Hatchet Creek, taken on 11th December 2015...... 19

3-2 Field site showing the location of Hatchet Creek...... 20

3-3 Hatchet Creek before 15th February 2017 and after the level of water in the creek rose to the floodplain 23rd July 2017. The red boxes outline the limb of a tree at the top of the floodplain showing the change in the water level...... 24

4-1 Variation in temperature and mean GSI values of females and males...... 38

4-2 Etheostoma f. barratti adult male, 46.0-mm-SL (top) and female, 44.0-mm-SL (bottom). Reproductive color is visible in the black bands in the dorsal fin and more pronounced dark pattern on the body of the male...... 39

4-3 The smallest specimen examined from the study areas, a 13.1-mm-SL juvenile of undetermined sex collected on 14th April 2017...... 39

4-4 Diet by size class of E. f. barratti. Numbers in parentheses indicate number of darters in each size class. Percentages <2% are not shown. Insecta refers to taxa other than those found in taxa identified...... 40

4-5 Seasonal diet of E. f. barratti...... 41

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

NORTH VERSUS SOUTH: A COMPARATIVE LIFE HISTORY STUDY OF THE SWAMP DARTER, ETHEOSTOMA FUSIFORME (GIRARD)

Adania Flemming

May 2018

Chair: Lawrence Page Major: Zoology

The life histories of subspecies of fishes differ from one another as a result of several factors including the differences in the environment associated with their geographic ranges. The life history for the Swamp Darter, Etheostoma fusiforme is well documented for the northern subspecies, but little information exists for the southern subspecies despite the opportunity for differentiation. Over a period of 18 months at Hatchet Creek in the Suwannee River drainage of

Florida, an ecological and behavioral study of the Swamp Darter revealed differences between the northern and southern populations with respect to life history characteristics. In the southern population, the Swamp Darter can live for at least fifteen months, spawns from January to April, and consumes a variety of invertebrates not found in the diet of the northern population. By better characterizing the differences in life history between the northern and southern populations of E. fusiforme, policy makers can be informed about strategies to protect or otherwise manage this species and the aquatic ecosystems it inhabits. Looking at why or when differences in life histories of subspecies arose may also help in our understanding of evolutionary processes.

CHAPTER 1 INTRODUCTION

The Swamp Darter, Etheostoma fusiforme (Girard), is a small percid fish distributed

(Figure 1-1) from southern Maine along the Atlantic Coast and Gulf Coast to about the Texas-

Louisiana border and north in the former Mississippi Embayment to Kentucky (Page 1983). As the name implies they occupy swamp habitats, and as a benthic species they are found in non- flowing or slow-flowing water over mud or detritus (Page 1983).

The Swamp Darter has had a complex taxonomic history, with 2 subspecies now recognized based on morphological data. The northern subspecies, E. f. fusiforme, is found from

Maine to Waccamaw River, North Carolina, while the southern subspecies, E. f. barratti, is found throughout the rest of the range (Page and Burr 2011). Etheostoma. f. barratti can be distinguished from E. f. fusiforme by the following characters: preopercle more often partially serrate (36% of specimens) versus usually entire (90% of specimens); usually 1 + 3 (70%) versus

2 + 3 (80%) infraorbital pores; 1–37 (푥̅: 13.2) versus 0–12 (푥̅ = 2.0) interorbital scales; usually over 25% (푥̅ = 57.5%) versus usually 0–20% (푥̅ = 9.5%) of parietal scaled (Collette 1962).

Etheostoma f. fusiforme was first described by Girard (1854) as Boleosoma fusiforme from Massachusetts, while E. f. barratti was described by Holbrook (1855) as Boleosoma barratti from the states of Florida and Georgia. Hubbs and Cannon (1935) recognized 5 subspecies of E. fusiforme based on clinal variation in morphological characteristics: E. f. fusiforme, E. fusiforme insulae, E. f. metaegadi, E. f. atraque and E. f. erochroum, and described a new species, Etheostoma thermophilus. Bailey (1954) recognized the subspecies, E. barratti appalachia, which according to Collete (1962) was based on an introduced population, which slightly differed from E. barratti in having a higher percentage of specimens with serrate preopercles.

Collette (1962) noted that E. fusiforme was the most variable species of the subgenus

Hololepis and argued that there were two ways to treat the variation. All distinguishable populations could be named, as Hubbs and Cannon (1935) had done, or the variations could be described with only two names. Collette opted for the latter, and placed most of the subspecies of

E. fusiforme and E. thermophilus in the synonymy of E. fusiforme fusiforme, while E. barratti and E. barrati appalachia were recognized as E. fusiforme barratti.

Etheostoma f. fusiforme is distributed entirely below the Fall Line except for its limited distribution in Connecticut above the Fall Line. The unusual distribution above the Fall Line prompted life-history studies of E. f. fusiforme in southern New England by Schmidt and

Whitworth (1979) and Schmidt (1983). These studies were initially done to determine if gaps in the Swamp Darter’s distribution in Connecticut were real or a result of inadequate sampling.

Subsequently, since life history data for the Swamp Darter was missing from the literature, they evaluated the feeding habits and age and growth. As a result, the life history for the northern subspecies, E. f. fusiforme, is well documented, whilst little is known for the southern subspecies,

E. f. barratti.

As noted by Cole (1954), comparative studies on life histories are as meaningful as comparative studies on morphology, psychology or physiology. Looking at differences in life histories of subspecies may help in our understanding of evolutionary and ecological processes.

Therefore, a life history study of the southern subspecies of the Swamp Darter, E. f. barratti was conducted to compare data with that on E. f. fusiforme gathered by Schmidt and Whitworth

(1979) and Schmidt (1983). The primary goal of the study was to document and compare life history traits of both subspecies of the Swamp Darter. Although not studied for the northern subspecies, data and observations on fecundity and spawning behavior were added to the study

to help with efforts to understand the environmental and conservation needs of the Swamp

Darter, E. f. barratti within a Florida population.

Figure 1-1. Distributional data of the Swamp Darter, Etheostoma fusiforme. Map obtained from iDigBio data portal.

CHAPTER 2 METHODS

Field Site Description

Hatchet Creek, in the Suwannee River drainage of Alachua County, Florida, supports a large population of E. f. barratti and was selected as the stream for this study. The creek flows south and east from Austin Cary Forest and into Newnans Lake. The principal study area was

Hatchet Creek at State Road 26 (Figures 2-1 and 2-2). The bank of the creek in this area is bordered with tall trees, which provide shade throughout most of the year. The creek has a sand bottom which is littered with woody debris -- mostly dead leaves and branches, from the hillslope canopy, while the tree roots along the bank of the creek provide habitat for small fishes and invertebrates. There are few to no rocks exposed in the creek.

Field Collection

Dip nets (fingerling dip nets, 15’’ X 18’’ 12 ‘’ deep, and 1/16 mesh size) were used to make monthly collections in Hatchet Creek from November 2015 to May 2017 (11/2 years), to capture individuals throughout the predicted lifespan of a Swamp Darter (≈1 year). In addition to dip nets, seines (5'ft deep, 50"-51" netting depth, 1/32"mesh) were used in July and August

2017 to capture individuals on the floodplain, where they had moved as a result of heavy rainfall.

Sampling continued until 20 specimens were caught, except in January and February 2016 when fewer specimens were captured as a result of high water, despite increased sampling efforts.

Beginning in December 2016, the number of specimens collected was increased to 25–35 individuals in order to increase the support for length frequency plots.

Most of the dip netting was done along the banks of the creek in areas containing root nodules of trees and vegetation, which have been noted as suitable Swamp Darter habitat in previous studies (Schmidt and Whitworth 1979, Schmidt 1983). The middle of the creek, which

had a sandy bottom, was also sampled for darters. Initially, the specific locations in the creek and types of substrate where specimens were captured were recorded. However, efforts to record the precise location of capture was abandoned after a few months when it became evident that it varied with the amount of water in the creek. Captured individuals were anesthetized with MS-

222 until limited mobility was observed, then transferred to 10-precent formalin to cease ongoing metabolic processes and preserve gut contents. Specimens were removed from formalin after 5 days and transferred to 70-percent ethyl alcohol.

The water and air temperatures, maximum water depth (measured monthly at the same site in the middle of creek), presence/absence of aquatic vegetation in locations darters were obtained, pH, and all species of fishes found were recorded during each collecting event.

Additionally, potential predators large enough to consume E. f. barratti were collected and dissected to observe gut contents. All invertebrates captured incidentally from March 2016 to

May 2017 were collected and identified to order or family to develop a database of potential prey items.

Laboratory

In the laboratory, preserved specimens were sexed by examining the genital papilla, dorsal pigmentation or, when necessary, the gonads. Very small specimens were identified as indeterminate if their gender could not be established using these methods. The standard length

(SL) and total length (TL) in mm of each darter were measured using a Vernier caliper prior to dissection. All darters were dissected by making an incision on the left side from the anus to the operculum. The liver was removed exposing the esophagus which was then cut at the anterior- most point. The intestine was then cut at its posterior end, which freed the entire digestive system from the body cavity and exposed the gonads, which also were removed from the body cavity.

All organs were placed in a small vial in 70-percent alcohol.

Diet

A representative sample (~ 25%) of the digestive systems of adult darters (greater than

25mm SL) from each collection was examined to identify food items and parasites. In collections containing juveniles (specimens ≤ 25mm SL), at least 5 juveniles were also dissected to compare the diet of juveniles to adults. Percent occurrence (number of specimens with food items divided by number of all specimens containing food in a given sample), and percent composition (the percent each food category contributed to total number of food items) of food items were calculated according to the methods of Schmidt and Whitworth (1979). Both percentages were used to determine significance of each food category (S), with S calculated as:

S= √ (% occurrence) (% composition). Values for the 2 subspecies were (2-1) compared.

Reproduction

To determine fecundity, 2 estimates were made: 1) gonadosomatic index (GSI) as the weight of the gonads (male and female) as a percentage of the adjusted body weight, and 2) counts of ova in both ovaries of a female. After removing the gonads, the specimen and its gonads were placed in water and blotted dry with a paper towel before they were weighed. The water was used to avoid the fluctuation in weight observed from the evaporation of ethanol, which occurred when specimens were weighed directly from the ethanol. The gonads and adjusted weight (weight of fish – weight of organs removed) of each fish was weighed using a

JT-D Analytical Digital Balance Scale to the nearest 0.001 gram and a Mettler Toledo AJ100

Analytical Digital Balance Scale to the nearest 0.0001 gram for smaller specimens. The gonadosomatic index was calculated according to Page (1971) as follows:

GSI = Weight of gonad (g)/Adjusted body weight (g) x 100, where adjusted body (2-2) weight is the weight of the female minus the ovaries, stomach and intestines.

Ova were counted and their diameters measured in mature females caught just prior to and during the reproductive season (Jan – May 2016; Jan – Apr 2017). Two specimens with the heaviest gonad weights were selected from each month for ova counts. Measurements were made using a dissecting microscope and ocular micrometer. All ova were counted and assigned to 1 of

3 categories: mature, intermediate or immature. Mature ova were partially translucent with an invaginated membrane and a diameter of 1.0 – 1.5mm. Intermediate (developing) ova were also

1.0–1.5mm but still opaque with several oil globules, while immature ova were 0.2–0.75 mm in diameter and lacked oil globules. Pearson’s correlation coefficient (r) was used to test for correlations between ova counts and standard length and adjusted body weight.

Development and Growth

To determine the age of the darters, a length-frequency plot was created (Figure 2-3).

Additionally, a scale was removed just below the lateral line of the anterior right side, in line with the origin of the dorsal fin (Scheider et al. 2000) in an effort to use annuli to age fishes. The scales were observed under the microscope to count annuli.

Aquarium Observations

On 19 February 2016, 5 individuals were placed in an aquarium for preliminary observation as a trial run to prepare the tank for spawning observations. The tank contained the following substrates known to be used for spawning by other darter species: sand, gravel, logs, plants (Microsorum pteropus, Java fern), and rocks/boulders (Page 1985). These individuals were removed from the aquarium after 2 weeks and 6 different individuals (3 males and 3 females all ≥ 40 mm standard length (SL) and reproductively mature) were added to the prepped aquarium tank on 1 March 2016. The aquarium temperature was 23°C.

The individuals in the spawning aquarium were recorded using a go-pro camera. The video footage was used to observe and describe reproductive and other social behaviors in the laboratory. Several eggs were collected from the aquarium immediately after the darters spawned and placed in an aquarium mesh breeder to prevent predation. Videography of E. f. barratti was obtained from a separate study of spring fishes by Kirsten Work (pers. comm.).

Statistics

Two-sample t-tests were used to compare the lengths of males to females and the effects of gender on GSI. The null hypotheses H0 (The difference between the population means (μ1 –

μ2) equals the hypothesized difference (δ0) were assumed and α=.05 (maximum error accepted

5%). Means (M) and standard errors (S.D) were reported as measures of central tendency and dispersion. A correlation hypothesis test was done to determine if there was a relationship between ova counts and standard lengths and ova counts and weight. A Chi-squared test was performed to evaluate the sex ratio of male to female darters.

Figure 2-1. Image of Hatchet Creek, taken on 11 December 2015. Photo taken by Zachary Randall.

Figure 2-2. Field site showing the location of Hatchet Creek. Map obtained from ArcMap version 10.5.

Figure 2-3. Standard lengths of all specimens collected over the study period.

CHAPTER 3 FIELD SITE DESCRIPTION (HABITAT ECOLOGY)

Hatchet Creek, watershed has more natural habitat than any of the watersheds in urban

Gainesville (Alachua County.us 2015). The riparian zone is heavily forested. Habitats are varied, consisting of floodplain swamp (Bald Cypress dominant), pine flatwoods, pond cypress domes, and mesic hammocks. Dominant canopy vegetation includes Oak spp., Maple spp., Sweet Gum,

Cypress spp., and Pine spp. Understory vegetation is predominantly Cherry, Elderberry, Wax

Myrtle, and Palmetto Spp. Aquatic Macrophytes consist of Juncus spp. and St. Johnís Wart

(Cinnimomum spp.) (Alachua County.us 2015).

The level of water in the creek during the study period varied depending on rainfall. The maximum water depth measured was 114.3 centimeters except during summer 2017 (June through August), when the level of water in the creek rose to the floodplain because of excessive rainfall (Figure 3-1). On occasions where the water level was low, the creek occupied about 1/3 or less of the width of the streambed or was severed into isolated pools. As described by

Marshall (1946) the water at Hatchet Creek has a reddish-brown hue due to a high content of organic matter, which produces an acidic reaction. In the present study, the acidity of the creek fluctuated with the amount of rainfall, with the lower pH found in high water conditions. The pH ranged from 6 to 7.8, while the water temperature ranged from 11.3°C in March 2017 to 25.1°C in July 2016 (Figure 3-1). Temperatures recorded during the period of study showed a positive correlation between air and water temperatures, with the latter being a few degrees lower.

The fish diversity at Hatchet Creek included Gambusia holbrooki (Mosquitofish),

Heterandria formosa (Least Killifish), Lepomis macrochirus (Bluegill), Lepomis auritus

(Redbreast Sunfish), Lepomis gulosus (Warmouth), Aphredoderus sayanus (Pirate perch),

Noturus gyrinus (Tadpole Madtom), Esox americanus, (Grass Pickerel), Elassoma evergladei

(Everglades Pygmy Sunfish), Elassoma okefenokee (Okefenokee Pygmy Sunfish), Fundulus chrysotus (Golden Topminnow), Notropis chalybaeus (Ironcolor Shiner). In July 2017

Hoplosternum literole (Brown Hoplo) and Lepisosteus platyrinchus (Florida gar) were found after heavy rains caused extensive flooding and probably entered the creek from Newnans Lake.

There were no other species of darters present.

Figure 3-1. Hatchet Creek before 15 February 2017 and after the level of water in the creek rose to the floodplain 23 July 2017. The red boxes outline the limb of a tree at the top of the floodplain showing the change in the water level. Photo taken by Adania Flemming (left) Cathleen Bester (right).

CHAPTER 4 RESULTS/DISCUSSION

During the study period (Nov 2015 – August 2017), 475 darters were collected, of which

431 (collections through May 2017) were dissected. Collections were continued into summer

2017 after discovering a new digenean parasite within the digestive tract. In July 2017, after the initial study period, seining was included as a part of collection methods to facilitate the capture of E. f. barratti and its potential predators in the flooded conditions of the creek. In average depths conditions, individuals of E. f. barratti were mainly found along the banks of the creek.

While the creek was flooded during summer 2017, darters were located in the ‘new’ banks of the creek on the floodplain (Figure 3-3). The preferential habitat of E. f. barratti is slowly flowing water near stream banks with live and dead vegetation. Young-of-the-year were found among or in close proximity to vegetation, including below floating duckweed, in slower, more stagnant, backwaters in the creek. Occasionally, darters were found in the middle of the creek on the sand bottoms, but in such instances they were in close proximity to woody debris. In the aquarium, darters made use of all substrates, including logs, leaves, rocks and gravel.

Reproduction

Reproductive Cycle of the Male

The individual GSI values for males ranged from 0.50 to 2.80 with the highest values occurring in Jan – March 2016 and Feb – May 2017. Male darters did not show any discernable changes in the size of their genital papilla throughout the year; however, their testes enlarged as the spawning period approached as evidenced by the mean GSI values (Figure 4-1). When GSI values of males are compared to females, male darters appear to maintain a relatively low GSI throughout the year, (Figure 4-1). However, when the GSI values of males were plotted independent of females, males GSI values showed similar increases during the reproductive

season (Figure 4-2). The two sample t-test revealed that the mean GSI was significantly lower in male darters (M=1.0, SD=0.92) than in female darters (M=5.8, SD =4.55); t(436) =20.47, p<.001.

In breeding males, testes were white, large, and spongy in appearance compared to translucent and stringy in non-breeding males. Large males became very dark in early spring, with distinctive dark blotches, absent in females, in the membranes between the first 4 dorsal spines (Figure 4-3). Males of many species of darters have been observed to display a change in coloration prior to spawning (Page 1983), including the closely related E. gracile (Braasch and

Smith 1967).

Breeding tubercules (small bony protuberances) were present on the branched rays and the membranes between the rays of the anal and pelvic fins of mature males during the reproductive season. Tubercules have been noted to develop maximally on darters during the breeding season and function as contact organs during spawning, presumably aiding in the tactile stimulation of females (Page 1983). A microscope was necessary to observe the tubercules of E. f. barratti since, as in E. f. fusiforme, the tubercules are barely visible to the unaided eye

(Schmidt 1983).

Reproductive Cycle of the Female

The individual GSI values for females ranged from 0.02–21.70 with the highest values occurring during the spawning period, Jan – May in 2016 and 2017. For females, a GSI value above 5 indicated spawning potential while values above 10 were more typical for females with mature ovaries (Figure 4-1). The highest GSI for a female in 2017 was 21.7 for a fish (37.5 mm

SL) collected April 2017, and in 2016 was 19.3 for a fish (34.8 mm SL) collected February 2016.

In contrast to males, females did not undergo a notable color change prior to the spawning period. However, there was a steady increase in the size of gonads in the months preceding the spawning season (Figure 4-1) and genital papillae were observed to increase as well. Genital

papillae and gonads were largest during the spawning period, Jan – May 2016 and Jan–April

2017.

The number of mature ova in 18 females ranged from 4–226 (Table 4-1). The wide range in number of mature ova could be a consequence of the type of spawning undertaken by E. f. barratti. As an egg attacher (Page 1996), E. f. barratti deposits 1 to 2 eggs on every successful mating bout. This type of batch spawning results in variable numbers of mature ova left within the gonads of females. As expected, there was no correlation between ova counts and standard length, r = 0.198, p=0.431, or between ova counts and adjusted body weight, r= 0.189, p =

0.454.

The lack of correlation can be seen by the counts in Table 4-1. For example, in March

2017, two individuals from the same collection date, both approximately 38.0 mm SL had varying numbers of ova in each category. One individual had 600 immature ova, 20 intermediate ova, and 18 mature ova, while the other individual had 580 immature, 168 intermediate and 0 mature ova. It is likely that the second individual had recently spawned, reducing the number of mature ova compared to the first individual. It is difficult to quantify the total number of mature ova produced during the spawning season for E. f. barratti using wild-caught specimens.

Observations of ova maturity indicated the standard minimum length for reproductive maturity is

≈ 25 mm SL, 30 mm TL (Schmidt 1983) for Etheostoma f. barratti. Table 4-1 illustrates a 29.2 mm SL individual with mature ova.

Spawning

Etheostoma f. barratti was observed spawning several days after being put in an aquarium. The spawning behavior observed for E. f. barratti was similar to the behavior described by Fletcher (1976) for E. f. fusiforme. A male darter would display to a female with his dorsal fin erect and then pursue her. If the female showed interest, she would be mounted by the

male and escorted to vegetation for deposition of 1 or 2 eggs, while being stimulated by the movement of the male’s tuberculate pelvic fins, which was followed by fertilization of the eggs.

The encounter involved the male darter rapidly beating his pelvic fins, as the pair maneuvered through vegetation, usually perching upside down to attach eggs to the undersides of leaves or on logs. The pair would dart in and out of the vegetation, making several attempts at egg deposition.

The eggs were left unguarded after attachment. Other darters in the tank were observed picking at the leaves where eggs were presumably attached immediately after the spawning pair moved on.

Often the darters returned to the same mass of vegetation during spawning, taking short breaks in between bouts of deposition and fertilization by resting on the gravel substrate or a leaf. The temperature in the tank during spawning ranged from 21 to 23°C. Unlike in E. f. fusiforme (Schmidt and Whitworth 1979, Schmidt 1983), aggressive behaviors were observed during spawning in E. f. barratti. On several occasions, a male engaged in spawning activity would stop, unmount the female and chase away approaching males. The male would then remount the awaiting female and continue spawning. Despite the aggression observed by males, females were promiscuous, mating with different males within the aquaria.

Video footage of E. f. barratti from Hart Spring halfway from the headspring to the confluence within the Suwannee River, Florida, showed similar spawning behavior. The temperature of the spring was 21.8°C. In the video, as in the lab, the darters appeared to select a patch of vegetation ideal for egg attachment, and made several attempts at egg deposition and fertilization. The eggs were left unguarded after attachment.

Development and Growth

The eggs collected from the aquarium immediately after spawning did not survive incubation in the lab. No fry were observed in the tank or in the aquarium breeder after the 8–10-

day period where Fletcher (1976) observed eggs hatching. In the tank the darters presumably fed on the unguarded attached eggs and prevented development. The mesh aquarium breeder did not suffice for eggs to develop into fry.

Additionally, the age of E. f. barratti was not determined using scale annuli as the scales were not well marked – seemingly due to the lack of marked seasonality in Florida. The smallest specimen examined from the study areas was a 13.1-mm-SL juvenile of undetermined sex collected in April 2017. Several additional juveniles between 14.8 and 16.0 mm SL were collected between April and June in 2016 and 2017. Schmidt and Whitworth (1979) observed young-of-the-year early in the spawning season (July), but were unable to catch juveniles as they were too small for the mesh of their hand nets. Although our net size (1/16 in. mesh) was small enough to capture juveniles, it may not have been sufficient to capture smaller fry.

Schmidt and Whitworth (1979) categorized young-of-the-year as individuals <30 mm TL and lacking annuli. Yearlings were categorized as individuals with annuli (30.5–55 mm TL) or large breeding individuals collected in May and June but lacking annuli (32–46 mm TL).

Although age was not obtained using scales in the present study, the length frequency distribution and GSI values obtained indicate similar findings for the size relative to age for E. f. barratti (Figs. 2, 6). Noteworthy Schmidt and Whitworth (1979) found aging based on scale examination agreed well with the length frequency distribution. Figure 4-1 shows female GSI values increasing in January the second spring of their lives, darters are 1 year of age and

≥30mm SL (Figure 2-1).

The 13.1 mm SL juvenile examined had a morphology similar to that of an adult, but was not fully developed (Figure 4-4). This specimen had 7 black lateral blotches and dorsal saddles, both of which consisted of black specks (some only visible under the microscope). The lateral

line and scales were not visible. Observations of other juveniles showed that at 16 mm the lateral line canal had begun to form, and scales were present on the caudal peduncle. At 18 mm, the black specks fused to form blotches, and the 10 dorsal saddles found in adults were present. At

19 mm, the lateral line extended to the 1st spiny dorsal. At 21 mm, the lateral line was completely developed (extending to mid-body). Etheosoma f. barratti portrays full adult morphology at 21 mm SL.

The two sample t-test for the 442 specimens identified to gender (n=220 male, n=222 female) revealed that the mean standard lengths were not different between males (M=35.21,

SD= 4.67) and females (M=34.9, SD =5.41), t(431) =-0.69, p=.489. These findings contrast somewhat with previous literature, which states that species of benthic darters living over mud

(i.e., species of the subgenus Boleichthys of Etheostoma) tend to have larger females than males

(Page 1983, Page and Swofford 1984). When observing the specimens collected from Hatchet

Creek individually, only 3 of the 10 largest specimens collected were male, or 8 of the 20 largest were male (Table 4-2). Therefore, without statistically comparing the lengths of E. f. barratti between gender, females may tend to appear larger than males.

Etheostoma f. barratti reached adult size by late winter, and young-of-the-year were produced in early spring (March and April). The length distribution and GSI values indicated darters in the southern population live up to 1 year (possibly 15 months), reproduce and die shortly after (Figure 2-1). The largest recorded specimen for E. f. fusiforme measured 55 mm in

TL and was thought to be in the second fall of life. Comparably, the largest recorded specimen for E. f. barratti was a 59-mm-TL reproductively mature male, in the first spring of its adult life.

Therefore, although the northern subspecies may live longer (2 years) (Collette 1958), it may reach a smaller maximum size than the southern subspecies.

Demography

Territoriality

Etheostoma f. barratti did not exhibit territorial behavior in the laboratory except during the spawning period. A spawning male would unmount a female if the pair was approached by another male and chase the other male out of the vicinity, usually away from the vegetation where spawning was occurring, and return to the female.

Composition

Of the 475 E. f. barratti collected in Hatchet Creek, 3% were 10–19 mm SL, 13% were

20–29 mm SL, 64% were 30–39 mm SL, and 20% were 40–50 mm SL. The 30–39 mm SL size class is the reproductively mature specimens; larger individuals approach death after spawning while smaller individuals are actively feeding and avoiding predation. The results of the χ2 test showed there was no difference in sex ratio among all size classes of darters combined (α< 0.05).

There was also no difference in sex ratio for all size classes individually assessed except within the 20–29 mm SL class (α > 0.05), in which males were larger than females. Female darters are often found to be smaller than males as females put more emphasis on gamete production while males put emphasis on growth since bigger males have an advantage in reproductive success

(Page 1983).

Diet and Feeding Habits or Diet and Parasitism

Stomach contents of 431 darters (0 empty) indicated the darters ate a diverse array of organisms, with the composition varying by size class (Figure 4-5). However, midge larvae

(Chironomidae) and water fleas (Cladocera) were the most numerous food items for all size classes. Darters in the smallest class, 10–19 mm SL, were observed to consume a greater amount of smaller food items (e.g. insect larvae) compared to the larger size classes’ consumption of bigger food items (e.g. insect pupae and adults). The diet of the larger size classes was the most

variable, as their larger mouths facilitated consumption of a greater variety of food items. Figure

4-5 illustrates the variation in consumed food items by season. No discernable seasonality in the diet was found (Table 4-3, Figure 4-6). Cladocera, Chironomidae, Trichoptera (caddisflies) and

Ephemeroptera (mayflies) were consumed monthly throughout each year. Other food items were consumed periodically throughout the study period.

Schmidt and Whitworth (1979) reported the significance of each food category (S); midge larvae (S=23.9) and Cladocera (S=51.5) were the most numerous food items for E. f. fusiforme, while copepods S= (19.78) and ostracods (S=18.7) were frequently eaten and mayfly and caddisfly larvae (S<1) were eaten occasionally. For E. f. barratti, midge larvae (S= 42.5) and

Cladocera (S=57.5) were the most numerous food items. However, there was much more variation in other frequent and occasional food items (caddisfly larvae (S=18.0), copepods

(S=12.0), mayfly larvae (S=8.58), damselfly larvae (Odonata) (S=5.08), diving beetles

(Coleoptera) (S=3.13), amphipods (Amphipoda) (S=3.02), isopods (Isopoda) (S=1.47), and all other miscellaneous (S<1) (Figure 4-5).

The increased complexity in the diet of E. f. barratti relative to that of E. f. fusiforme may be attributed to latitudinal changes in diet or the invertebrate diversity. Schmidt and Whitworth

(1979) explained that the low significance value for midge larva resulted from low representation in terms of the percent of total food items. They noted Swamp Darters could not contain more than 5 midge larvae because of their size, but cladocerans were as numerous as 140 per stomach.

Although there was a high significance value for cladocerans in this study, there was also a comparably high significance value for midge larvae (S= 42.5). In some specimens of E. f. barratti examined, as many as 15 midge larvae were found in the digestive tract. A more

comprehensive understanding of the differences in diet of subspecies of the Swamp Darter will be an interesting addition to our knowledge of the ecology of darter habitats.

Etheostoma f. barratti was observed to be a visual fish. This was seen during captive feeding as the response time to retrieval of food items varied based on the color and movement of the food items. Darters were more responsive to blood worms with bright red coloration versus blood worms which had lost their coloration. Additionally, darters took more time to obtain brine shrimp, and their reaction time was directly proportional to the movement of the brine shrimp; the more the shrimp moved the greater the chance the darters would grab it. As a group, darters have been known to have keen vision and are assumed to be diurnal feeders (Page

1983), which would account for such observations in the lab. Darters collected early in the morning or late afternoon had fewer food items in their stomachs compared to darters collected late to mid-morning.

Interactions with Other Organisms

Predation

No specimens of E. f. barratti were found within the guts of predators large enough to consume the Swamp Darter. However, E. fusiforme is known to be an important forage fish and has been found in the diets of Grass Pickerel (Esox americanus), Largemouth bass (Micropterus salmoides), Black crappie (Pomoxis nigromaculatus) and Longnose gar (Lepisosteus osseus)

(Everhart 1966 and Collette 1962).

Parasitism

A taxonomically undescribed fluke (digenean trematode) of the family Allocreadiidae was discovered as a parasite of E. f. barratti. A total of 676 of these digeneans was removed from the intestines of 130 darters. Approximately 50% of all darters dissected for gut content analysis contained at least 1 digenean. These parasites were located in the digestive tract between

the pyloric caeca and the intestine. Using histology of mounted specimens, the parasite was identified as belonging to the genus Crepidostomum. Adults of this genus are usually located in the intestine, sometimes in the pyloric caeca or gall bladder of freshwater fishes (Hopkins 1934).

These digeneans grow within the fish and migrate to the intestine where adult flukes lay eggs, which are passed with feces into the environment (Hopkins 1934). Metacercariae (digenean larvae) encyst in aquatic insects, crustaceans or mollusc, which in turn, are eaten by a new host fish. The effect of parasitism on darters is unknown. A nematode was found in the gonads of one darter.

Conservation

Although Etheostoma fusiforme has been described as a hardy fish in this and previous studies (Collette 1962, Everhardt 1966, Schmidt and Whitworth 1979, and Schmidt 1983) it has been described as threatened in some parts of its range, for example in New York (Carlson and

Daniels 2004). In such places loss of swamp habitat was described as the underlying issue.

These conservation needs must be assessed and carefully managed to ensure this fish does not become threatened in the rest of its range.

Aquarium Observations

Four (3 males and 1 female) of the 6 individuals of E. f. barratti placed in the aquarium on 1 March 2016 are still alive (February 2018), 11 months after their first spawning event in

March 2017. Although previous studies (Schmidt and Whitworth 1979, Schmidt 1983, Collette

1962) and the current study suggest that E. f. barratti rarely if ever lives 2 years in nature, it clearly is capable of living 2 years in captivity where lack of predation and a steady food source contribute to long-term survival. Males have again become darker in coloration within the tank and are actively pursuing females.

Table 4-1. Mean lengths, weights and ova counts of female Etheostoma. f. barratti collected each month during the reproductive period. Month-year Adjusted Gonad Standard Immature Intermediate Mature collected body weight length ova ova ova weight*(g) (g) (mm) January-16 0.62 0.096 42.5 920 174 4 January-16 0.39 0.036 37.6 614 72 24 February-16 0.594 0.052 42.1 1100 90 24 February-16 0.449 0.051 39.2 400 180 0 March-16 0.47 0.068 41.0 300 180 128 March-16 0.176 0.013 29.2 150 80 10 April-16 0.466 0.053 42.4 560 200 0 April-16 0.537 0.057 42.0 740 320 0 May-16 0.779 0.055 46.8 480 0 190 May-16 0.499 0.038 40.2 280 300 0 January-17 0.438 0.037 38.7 680 214 0 January-17 0.453 0.051 39.6 720 162 2 February-17 0.377 0.007 37.0 600 272 0 February-17 0.47 0.071 41.0 460 220 140 March-17 0.347 0.035 38.0 600 20 18 March-17 0.443 0.033 38.8 580 168 0 April-17 0.404 0.073 38.5 800 240 198 April-17 0.391 0.085 37.5 540 26 226 *Adjusted body weight is the weight of the female minus the ovaries, stomach and intestines.

Table 4-2. Lengths and dates of the 20 largest specimens collected at Hatchet Creek. Date Gender Standard length (mm) Total length (mm) March-16 M 50.1 59.2 May-16 F 46.8 54.4 March-16 M 46.2 54.3 April-16 F 45.8 53.4 January-16 F 45.8 53.3 April-16 F 45.4 52.9 March-16 M 45.2 52.6 April-16 F 45.1 39.5 February-17 F 45.1 49.6 March-16 F 44.5 51.2 March-16 F 44.4 49.8 December-16 F 44.4 51.7 January-17 M 43.6 51.4 March-16 F 43.4 50.8 March-17 M 43.4 50.0 March-16 F 43.1 50.3 January-16 M 43.0 51.0 May-17 M 42.9 51.5 February-16 F 42.9 50.4 March-16 M 42.7 50.2

Table 4-3. Mean Number of Food Organisms Per Stomach. Nov Dec- Jan Feb Mar Apr May Jun- Jul- Aug Sep Oct Nov Dec Jan- Feb Mar Apr May -15 15 -16 -16 -16 -16 -16 16 16 -16 -16 -16 -16 -16 17 -17 -17 -17 -17 Food items (5) (5) (6) (5) (9) (8) (11) (10) (10) (8) (6) (4) (5) (5) (6) (6) (5) (8) (9) consumed Cladocera 0.3 0.3 0.3 1.6 0.1 0.8 0.4 31.9 -- 6.1 3.3 1.0 1.8 44.2 2.5 6.0 0.2 7.9 4.9 Chironomidae 4.2 6.8 1.7 3.4 3.9 5.3 7.0 1.7 4.1 1.4 2.7 12.5 8.0 7.6 5.0 6.6 8.2 1.4 2.3 Trichoptera 0.8 2.0 0.3 2.2 1.6 4.3 4.2 1.5 0.5 0.8 0.5 2.8 0.8 -- 1.3 0.8 -- -- 0.2 Ephemeroptera 0.4 0.6 -- 0.4 0.1 0.4 0.2 0.2 0.1 1.0 -- 7.3 1.0 0.2 0.5 -- 1.4 0.3 1.2 Odonata ------0.4 1.5 0.5 0.1 0.9 0.5 0.2 0.3 -- 0.2 0.2 0.2 ------Coleoptera ------0.5 -- 0.1 -- 1.4 0.2 0.3 0.4 -- -- 0.2 0.3 0.3 0.1 Crustacea ------0.2 -- 0.5 0.1 0.7 0.1 -- 0.3 -- -- 0.4 ------Copepoda ------0.6 0.1 -- 0.8 0.2 -- 0.3 ------5.2 2.2 0.6 4.8 2.3 3.9 Diptera ------1.3 ------0.1 ------0.2 ------Isopoda -- 0.4 0.2 0.4 0.2 1.0 ------0.2 ------0.1 -- Amphipoda ------0.2 1.0 0.4 2.3 -- 0.5 -- -- 0.2 48.2 32.8 4.6 2.4 18.9 15.6 Insecta -- -- 0.2 ------0.3 -- -- 0.2 ------Nematoda ------0.4 ------0.2 ------Gastropoda -- -- 0.3 ------Annelida -- 0.2 ------Hemiptera ------0.1 0.5 ------Ostracoda ------0.2 -- 0.1 -- Numbers in parentheses () indicate the number of stomach dissected

Figure 4-1. Variation in temperature and mean GSI values of females and males

Figure 4-2. Variation in mean GSI values of males.

Figure 4-3. Etheostoma f. barratti adult male, 46.0-mm-SL (top) and female, 44.0-mm-SL (bottom). Reproductive color is visible in the black bands in the dorsal fin and more pronounced dark pattern on the body of the male. Photo taken by Zachary Randall.

Figure 4-4. The smallest specimen examined from the study areas, a 13.1-mm-SL juvenile of undetermined sex collected on 14th April 2017. Photo taken by Jarred Randall.

Figure 4-5. Diet by size class of E. f. barratti. Numbers in parentheses indicate number of darters in each size class. Percentages <2% are not shown. Insecta refers to taxa other than those found in taxa identified.

Figure 4-6. Seasonal diet of E. f. barratti.

CHAPTER 5 SUMMARY

The life-history information on E. f. barratti collected in Hatchet Creek between

November 2015 and August 2017 is summarized in Table 5-1. The life-history information for the Hatchet Creek population of E. f. barratti was compared to the southern New England population of E. f. fusiforme (Schmidt and Whitworth 1979, Schmidt 1983). Additional life history traits were noted for E. f. barratti which have not yet been documented for E. f. fusiforme. It will be useful to obtain this information in future studies of the Swamp Darter.

Table 5-1. Summary of life-history information on the Hatchet Creek population of E. f. barratti compared to the southern New England population of E. f. fusiforme (Schmidt and Whitworth 1979, Schmidt 1983). Characteristics Life History Data of E. f. Life History Data of E. f. fusiforme barratti Principal habitat Slowly flowing water near Soft muddy substrates with dense of adults stream banks with live and dead vegetation, deep undecomposed leaf vegetation piles or less often a rocky bottom littered with sticks and leaves Principal habitat Slowly flowing water near Not mentioned; assumed same as above of young stream banks with live and dead vegetation Size at reaching Approximately ≈25 mm SL Approximately 30 mm TL sexual maturity Sexual Male, dark body with Male, the membranes between the first dimorphism distinctive dark blotches in the four dorsal fin spines become membranes between the first blackened, frequently appearing as a four dorsal spines, tuberculate dark blotch or bar, the anal and pelvic pelvic and anal fin rays; fins develop darker spots than normal, Female, enlarged genital and the cheeks, belly, and breast areas papillae are also darkened, tuberculate pelvic and anal fin rays; Female, not mentioned assumed same. Breeding Present Present tubercules Number of 4–226 Unknown/ not studied mature ova Description of 1.5 mm Unknown/ not studied egg on leaf Spawning period January to April April and May Spawning habitat Aquatic vegetation, logs Leaves of plants, logs Spawning Both male and female inverted, Both male and female inverted, head to position head to head head Egg guarding No; eggs abandoned after No; eggs abandoned after spawning spawning Sex ratio 1:1 male to female Unknown/ Not studied Longevity 1+ year ≈2 years Maximum size 59 mm TL 55 mm TL Migrations None observed Unknown/ not studied Territoriality Extreme in breeding males None observed Principle diet Aquatic insect immatures and Small invertebrates crustaceans Parasites Crepidostomum sp. None observed

LIST OF REFERENCES

Alachua County US. 2015 Alachua County.us creek reports available at http://www.alachuacounty.us/Depts/epd/WaterResources/CreeksAndLakes/Pages/Reports-and- Maps.aspx. Accessed November 2015

Bailey, R.H. Winn, and C. Smith. 1954. Fishes from the Escambia River, Alabama and Florida, with Ecologic and Taxonomic Notes. Proceedings of the Academy of Natural Sciences of Philadelphia, 106: 109–164.

Braasch, M.E., and P.W. Smith. 1967. The life history of the slough darter, Etheostoma gracile (Pisces, Percidae). Illinois Natural History Survey Biological Notes, 58:1–12.

Carlson, D.M., and R.A. Daniels. 2004. Status of fishes in New York: increases, declines and homogenization of watersheds. The American Midland Naturalist, 152(1):104–139.

Collette, B.B. 1962. The swamp darters of the subgenus Hololepis (Pisces, Percidae). Tulane studies in zoology. 9:115–211. doi:10.5962/bhl.part.25183

Cole, L.C. 1954. The population consequences of life history phenomena. The Quarterly Review of Biology, 29 (2):103–137.

Everhart, W.H. 1966. Fishes of Maine. The Maine Department of Inland Fisheries and Game. l– 77 pp.

Fletcher, A.M. 1976. A rare species of darter spawning. American Currents 4(1):20–22

Holbrook, J.E. 1855. An account of several species of fish observed in Florida, Georgia, etc. Academy of Natural Sciences. Proceedings of the Academy of Natural Sciences of Philadelphia 3:47–47.

Hopkins, S.H. 1934. The Papillose Allocreadiidae: A Study of their Morphology, Life Histories, and Relationships. Illinois Biological Monographs, 13(2): 7–80.

Hubbs, C.L. and M.D. Cannon. 1935. The darters of the genera Hololepis and Villora. Miscellaneous Publications Museum. Zoology University. Michigan (30): 1–93. iDigBio. 2015. iDigBio data portal Available online at http://www.idigbio.org/portal.Accessed on 15 October 2015.

Marshall, N. 1946. Studies on the life history and ecology of Notropis chalybaeus (Cope). Quarterly Journal of the Florida Academy of Sciences, 9 (3/4):163–188.

Page, L.M., 1983. Handbook of darters. T.F.H., Inc., Neptune City, N.J. 271 p.

Page, L.M.1985. Evolution of reproductive behaviors in percid fishes. Illinois Natural History Survey Bulletin, 33(3):275–295.

Page, L.M., and R.L Mayden, 1981. The life history of the Tennessee snubnose darter, Etheostoma simoterum in Brush Creek, Tennessee. Illinois Natural History Survey Biological Notes 117:1–11.

Page, L.M. and P.W. Smith. 1970. The life history of the dusky darter, Percina sciera, in the Embarras River, Illinois. Illinois Natural History Survey Biological Notes 69. 15 p.

Page, L.M., and P.W. Smith. 1971. The life history of the slenderhead darter, Percina phoxocephala, in the Embarras River, Illinois. Illinois Natural History Survey Biological Notes, 74:1–14 p.

Page, L.M., and D.L. Swofford.1984. Morphological correlates of ecological specialization in darters. Environmental Biology of Fishes 11:139–159, and Developments in Environmental Biology of Fishes 4:103–123. doi:10.1007/978-94-009-6548-5_12

Schmidt, B.R.E. (1983). The Swamp Darter. North American Native Fisheries Association. American Currents.

Schmidt, R.E., and W.R. Whitworth. 1979. Distribution and habitat of the Swamp Darter (Etheostoma fusiforme) in southern New England. American Midland Naturalist, 102(2):408– 413.

Schneider, James C., P.W. Laarman, and H. Gowing. 2000. Age and Growth Methods and

State Averages. Chapter 9 in Schneider, James C. (ed.) 2000. Manual of Fisheries Survey Methods II: With Periodic Updates. Michigan Department of Natural Resources, Fisheries. Special Report 25, Ann Arbor.

BIOGRAPHICAL SKETCH

Adania Flemming was born and raised in the twin island nation of Trinidad and Tobago.

She attended Arima Government Secondary, where she obtained her GCE O-level in 2004.

Thereafter, she attended Bishop Anstey High School where she attained her GCE A-level in

2006, studying chemistry, biology and physics. Her love for nature and fascination with water and aquatic life compelled her to pursue a Marine Science degree.

Adania left her home in Trinidad to attend The University of Tampa’s undergraduate program in 2007, where she obtained a Bachelor of Science, double majoring in marine science and biology. During her tenure as an undergraduate Adania participated in several research projects. These included a mark and recapture project of sygnathid species to evaluate ecological habitat changes during environmental perturbations, a taxanomic reorganization of cymothoid isopods of Tampa Bay and a project recording the photosynthetic ability of various species of the genus Caulerpa. She also interned at the Florida Aquarium in Tampa, as an education intern where she conducted daily shows, tours and educational presentations about marine life and ecosystems.

Upon graduation in 2011 Adania accepted a position as an exhibit educator at the Florida

Aquarium, where she worked for a year before leaving to further her studies. Adania accepted a position with the Invertebrate Zoology collection at the Florida Museum of Natural History in

2012. She worked in the collection for 2 years, gaining experience, teaching science camps for

K-6 students and participating in science outreach events. Adania was married to Jason Fraser-

Nash on May 20 2013. Adania accepted a position as a Collection Technician in the Ichthyology

Collection and Project Assistant for iDigBio in 2014. Adania continued to teach science camps for K-6 students as well as participate and organize science outreach and education events within both roles, travelling across the US to encourage use of Natural History collections in K-12

classrooms. Fueled by her passion for science and new found understanding of fish, Adania started her Master of Science in zoology working on a Comparative Life History study of the

Swamp Darter, with Dr Larry Page. Adania taught the Integrated Principles of Biology course in fall 2015 and fall 2016 and the Critical Analysis of Biological Research in fall 2017.

To compliment her degree in Zoology Adania added a Certificate in Museum Studies in

August 2017. One of Adania’s long-term goals is to open a Research Aquarium/Museum in her home island, which she hoped to gain more skills for through the certificate program. She created an Introduction to Natural History Collections course at the University of Florida for undergraduates which she taught in spring 2018. The class was aimed to introduce students to alternative career paths from pre-professional fields, through observation of and immersion into the roles of collection personnel. Adania will obtain her Master of Science in zoology and

Certificate in Museum Studies from the University of Florida May 2018.