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Published in final edited form as: J Therm Biol. 2021 Jun 16;100:103004. doi: 10.1016/j.jtherbio.2021.103004

Flight system morphology and minimum flight temperature in North American cicadas (Insecta: Hemiptera: Cicadidae)

Allen F Sanborn 1,1, Earl A Allick 1,2, Sandee V Apang 1, Izyanna D Castillo 1, Erica M Cruz 1, Theophilus H Davis 1,3, Cory H Duncan 1, Fanny Fierro 1, Marla RM Gebaide 1, Abigail Luke 1, Maria L Pacheco 1, Daniel Paz-Castillo 1, Laura M Perez 1, Ana C Poeck 1, Adrian K Seepersaud 1, Carolina G Valdes 1
PMCID: PMC8433486  NIHMSID: NIHMS1716935  PMID: 34503771

Abstract

Thermal responses in cicadas have been studied for many years. The minimum flight temperature (MFT) does not show the same relationship to habitat and behavior as other thermal responses. We measured live mass, wing length, wingspan, wing area and wing loading in an attempt to correlate these morphological parameters to the MFT. We analyzed both intraspecific (in Magicicada cassinii (Fisher, 1852)) and interspecific relationships of the wing morphology and the ability of the cicadas to fly in a large number of North American cicada taxa (n=119). A total of 109 species and 10 subspecies from 17 genera, six tribes, and three subfamilies including all major North American habitats were studied. Analyses show that wing morphology (wing length, wingspan, wing area and wing loading) scales to body size as predicted by geometric similarity (all P<0.0001) for all species and wing area and wing loading (both P<0.0001) in M. cassinii. Mass (P=0.0105), wing length (P=0.0006), wingspan (P=0.0006), wing area (P=0.0055), and wing loading (P=0.0455) all demonstrate a significant correlation to MFT between species, as would be predicted by aerodynamic theory, but not within species. However, the low correlation coefficients suggest the flight system has minimal influence on the MFT of cicadas. Specific physiological adaptations appear to be responsible for the between species variability in MFT rather than being the result of modifications to the flight system morphology.

Keywords: Thermal Responses, wing morphology, flight

1. Introduction

1.1. Cicada thermal responses

Heath (1964, 1967) began the measurement and analysis of thermal adaptation in cicadas with a study of the thermal responses of the periodical cicada Magicicada cassinii (Fisher, 1852). Since that time, thermal adaptation to various environments from five continents has been investigated (see summary in Sanborn 2002, 2004; Sanborn et al. 2003, 2004a, b, c, 2011a, b, c, d, 2012, 2017; Heath and Sanborn 2007; Sanborn and Heath 2009; Sanborn and Phillips 2010, 2011). These analyses have demonstrated a variety of biological principles including convergent evolutionary patterns in cicadas from Argentina and the United States (Sanborn et al. 2004b), Australia and the United States (Sanborn et al. 2004a), and Mediterranean habitats found on four continents (Sanborn et al. 2011a), support for the hypothesis that tropical ecosystem diversity is in part a result of adaptation to specific thermal niches (Sanborn et al. 2011c), the differentiation of species or subspecies (Sanborn and Phillips 1996, 2001, 2010, 2011; Sanborn et al. 2011d, 2012), the influence of ambient conditions on the evolution of thermal responses (Sanborn et al. 2003, 2004a, b, 2011a, 2017), and adaptation to local conditions like altitude (e.g. Heath et al. 1971; Sanborn et al. 2011b).

Two thermal tolerance measurements show relationships to the habitat used by a species. The heat torpor temperature is the maximum body temperature (Tb) to maintain motor control and is related to the potential maximum temperature a species will encounter in its environment (Sanborn et al. 2017). Thermoregulatory behavior and habitat have been shown to influence the maximum voluntary tolerance temperature or upper thermoregulatory set point of a species (Sanborn et al. 1995a, b, 2017; Sanborn 2000, 2004). However, similar relationships to habitat or behavior are not apparent with respect to the minimum flight temperature (MFT) in cicadas even though flight system morphology exhibits geographic variation and is related to habitat in some insects (e.g., Hassall et al. 2009; Hassall 2015).

Early work with cicadas (Heath et al. 1971) demonstrated a relationship between MFT and altitude. However, an increase in the number of species studied showed this relationship to be an artifact of the small number of species included in the original study and the specific species used in the analysis. This conclusion was confirmed in the study of Sanborn et al. (2011b) where the MFT of 12 species in a limited geographic range showed no relationship to altitude while the maximum voluntary tolerance and heat torpor temperature did. Having begun to see the variability in MFT with habitat not illustrated in the maximum voluntary tolerance or heat torpor temperatures, Heath et al. (1972) suggested that the flight system might influence the MFT.

1.2. Temperature influences on flight

We investigate the potential influence of the flight system on MFT with this work. The Tb permitting flight is specific to the species not only in cicadas but in other insect groups as well (e.g., Nève, 2010). The contraction kinetics of insect flight muscle are temperature dependent (e.g., Josephson 1981) so that wing beat frequency is also dependent on temperature. The relationship between wing size and wing beat frequency (Dudley, 1990) can also be related to the thermal parameter since it takes more force to move a larger wing and warmer muscles generate greater force (e.g., Josephson, 1981). The muscle temperature influences the aerodynamics of the cicadas, particularly the lift needed to remain airborne, in a predictable manner based on aerodynamic principles as we have suggested with preliminary studies (Sanborn and Davis 1992; Sanborn et al. 1996, 2001). Wing beat frequency is influenced by body size and forewing area in cicadas (Ha et al. 2013) as well as scaling with body size in insects in general (Dillon and Dudley 2004; Park and Yoon 2008) but is influenced by temperature through the contraction kinetics of the flight muscles. Thermal influences on wing beat frequency appear to have stimulated the development of larger wings in Drosophila Fallén, 1823 found in habitats with colder ambient temperatures (Gilchrist and Huey 2004). Perhaps more importantly, temperature influences the force of muscle contractions (e.g., Josephson 1981) changing the wing stroke amplitude and has a greater influence on lift than wing beat frequency (Grodnitsky 1999; Dudley 2000).

The aerodynamics of the flight system should influence minimum flight temperature as well since lift is influenced by the flight system morphology. Lift is proportional to air density, velocity squared, the coefficient of lift, and the wing area (Vogel 1994). Since lift is proportional to the wing area and square of the air velocity over the wing, greater lift will occur with greater wing area. The air velocity will be correlated to the wing beat frequency, a temperature dependent process as outlined above. In addition, air velocity is inversely related to wing length in flying animals (Greenwalt 1962) and inversely related to body mass in insects (Dudley 2000). The rigidity of the leading edge in cicada wings also increases the lift coefficient in cicadas (Park and Yoon 2008). Similarly, the Kutta-Joukowski Theorem states that lift for any given wingspan is proportional to air velocity (Vogel 1994) suggesting that greater wing length and wingspan will increase lift production at any given air velocity. Body size also influences lift production (e.g., Samjima and Tsubaki 2010) and temperature has been shown to influence both lifting force (Samejima and Tsubaki 2010) as well as acceleration (Berwaerts and Van Dyck 2004) in insects.

Wing area and lift generation will also be influenced by the mass and gravitational pull the wings must overcome to generate sufficient lift to fly. Wing loading is the weight (mass times gravitational acceleration) divided by the wing area (Vogel 1994). Wing loading is related to both lift generation equations through air velocity because velocity is proportional to the square root of wing loading (Alexander 1985; Dudley 2000). Wing loading influences how much thrust is generated per wing beat (Dudley 2000) and wing loading is proportionate to the minimum flight velocity (Alexander 1985; Ahlborn 2004). This results in lower takeoff speeds for larger wings with lower wing loading.

Although these aerodynamic relationships are simplified aspects of insect flight, the outlined aerodynamic principles and temperature effects on insect muscle suggest that wing length, wingspan, and wing area should correlate to minimum flight temperature since lift is proportional to wing area and wingspan. The larger species should also generate greater lift with their larger wings. Wing loading should correlate to minimum flight temperature since wing loading is related to air velocity which influences lift generation and velocity will be dependent on the muscle contraction kinetics which are also temperature dependent (e.g., Josephson 1981). These relationships suggest that morphological characteristics generating greater lift should lead to a lower minimum flight temperature since lower muscle contraction frequencies are necessary to generate sufficient lift to overcome gravitational pull.

We investigate the morphology of the cicada flight system and any potential influence on minimum flight temperature. While there are undoubtedly physiological adaptations that can influence MFT for specific species (e.g. Sanborn et al. 2011c), we attempt to determine if the morphology of the lift generating systems in cicadas as a group plays a role in permitting flight at a particular Tb.

2. Materials and methods

2.1. Ethics statement

The experiments performed in this study used hexapod invertebrates and are thus exempt from meeting the requirements of the Guide for the Care and Use of Laboratory Animals of the National Institutes of Health and EU Directive 2010/63/EU.

2.2. Animals

The thermal responses of 119 North American cicada taxa summarizing more than 50 years of field research and experimentation was recently published by Sanborn et al. (2017). Live cicadas were collected on annual expeditions between 1963–1973 and 1979–2016 to determine the MFT (Sanborn et al. 2017). Most of these specimens were used for determining the morphological variables included here. Specimens are primarily deposited in the first author’s collection with some specimens deposited in the Maxine and James Heath collection (Buchanan Dam, TX). Cicadas were collected in 25 states including all states except Iowa and Nebraska encircled by a transect from Minnesota to Washington to California to Florida to Tennessee to Illinois (Sanborn et al. 2017).

2.3. Taxonomy

The total of 109 species and 10 subspecies represent 17 genera, six tribes and three subfamilies. The higher taxonomy and species diversity are summarized in Table 1. Higher taxonomy follows Marshall et al. (2018) and differs slightly from Sanborn et al. (2017) specifically in the genera Diceroprocta Stål, 1870 and Quesada Distant, 1905 being transferred to the Fidicinini Distant, 1905.

Table 1.

Taxonomic hierarchy and diversity of the cicadas used in this study.

Family Subfamily Tribe Genus
Cicadidae Latreille, 1802
Cicadinae Latreille, 1802
Cryptotympanini Handlirsch, 1925
Neotibicen Hill & Moulds, 2015 – 11 species, 2 subspecies
Megatibicen Sanborn & Heath, 2016 – 7 species, 1 subspecies
Hadoa Moulds, 2015 – 10 species
Cornuplura Davis, 1944 – 1 species
Cacama Distant, 1904 – 3 species
Fidicinini Distant, 1905
Diceroprocta Stål, 1870 – 20 species, 3 subspecies
Beameria Davis, 1934 – 3 species
Pacarina Distant, 1905 – 2 species
Quesada Distant, 1905 – 1 species
Leptopsaltriini Moulton, 1923
Neocicada Kato, 1392 – 2 species, 1 subspecies
Cicadettinae Buckton, 1890
Lamotialnini Boulard, 1976
Magicicada Davis, 1925 – 6 species
Tibicininae Distant, 1905
Platypediini Kato, 1932
Platypedia Uhler, 1888 – 5 species, 1 subspecies
Neoplatypedia Davis, 1920 – 1 species
Tibicinini Distant, 1905
Clidophleps Van Duzee, 1915 – 2 species
Okanagana Distant, 1905 – 32 species, 1 subspecies
Okanagodes Davis, 1919 – 2 species, 1 subspecies
Tibicinoides Distant, 1914 – 1 species
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2.4. Minimum flight temperature experiments

The MFT and live weight values used here (Supplementary Table 1) are the values reported by Sanborn et al. (2017) with n=1–179 for MFT and n=1–298 for live mass depending on taxon. Live weight was measured on a triple-beam balance accurate to ±5 mg on the evening of capture (see procedures in Sanborn et al. 2017). Included in these data is a summary of the previously published thermal responses for North American cicadas (Heath 1967; Heath and Wilkin 1970; Heath et al. 1971, 1972; Heath 1972; Sanborn et al. 1992, 2002a, b; 2004a, b, 2011a, b, d, 2012; Sanborn and Phillips 1996, 2001, 2010, 2011; Sanborn, 2000, 2004; Sanborn and Maté 2000; Heath and Sanborn 2007; Sanborn and Heath 2009). The value with the largest sample size in Sanborn et al. (2017) was used in the analyses here if a species was studied more than once previously.

MFT values were determined following established procedures for cicadas first outlined by Heath (1967) and Heath and Wilkin (1970). Animals were cooled to a torpid state and tossed 1–2 m into the air to stimulate flight. Animals warmed passively to the point where they were able to make a controlled flight or landing. At this time, they were grabbed by the wingtips in an effort to prevent conductive heat transfer from the scientist to the animal and a thermocouple inserted midway into the dorsal mesothorax to measure deep Tb. Tb was measured with a Physitemp Model BAT-12 digital thermocouple thermomenter (Physitemp Instruments Inc, Clifton, NJ) and a type MT 29/1 29 gauge copper/constantan hypodermic thermocouple accurate to ±0.1 °C with a time constant of 0.15 s or a Telethermometer thermometer with a 26 gauge hypodermic thermister probe. Both probes were calibrated with a National Institute of Standards and Technology thermometer to insure accuracy. It is important to note that these procedures produced consistent results within species with individual populations that were studies over distances of 1000 km and time spans of 34 years (Sanborn and Heath, 2009). It is also important to note that habitat rather than phylogeny influenced thermal tolerances where a relationship was noted previously (Sanborn et al., 2011b, 2011c, 2011d, 2017).

2.5. Morphological experiments

Morphological measurements were collected over the past 30 years on dried specimens and wing tracings and are summarized in Supplementary Table 1. Wing length and wingspan were determined with Vernier calipers accurate to ±0.05 mm. Wing length was determined by measuring the distance from mesothoracic attachment to fore wing tip. Wingspan was determined by measuring from wing tip to wing tip in spread specimens or by adding the wing lengths and the mesonotum width at the base of the wings. Wing area was determined from tracings of the wings in a natural flight position so that the coupled junction between the fore and hind wings was perpendicular to the main body axis. A digitizing tablet (Kurta IS/ONE, Wacom INTUOS Creative Pen & Touch Tablet, or Wacom CT-0405-U Pen Partner) and NIH Image software (Image or ImageJ) were used on a Macintosh computer to determine wing area from the tracings. The tablet and software were calibrated at the beginning of each digitizing session. Each wing tracing was digitized a total of ten times with the average of the ten tracings used as the area value for the individual wing to minimize any deviation obtained in a single tracing. Wing area for the left and right wings were added to determine the total wing area for an individual specimen. All imperfections (i.e. damage) were included in the measurement of the area as the wings as the imperfections were present when the MFT value was determined for that specimen and potentially influenced the MFT value determined. Wing loading was determined by dividing the mean live weight by the mean wing area determined for each species. Aspect ratio was not investigated since it is unclear whether the aspect ratio is a significant design parameter determining optimum performance in insect flight (Harbig et al. 2013).

Mass, wing area and wing loading data were correlated to individual specimens for the 42 specimens of Magicicada cassinii (Fisher, 1852) that were studied during the 1990 MFT experiments (Sanborn and Heath 2009) and used to investigate within species relationships of wing area and wing loading and MFT. Additional specimens sent to the first author were used to determine morphology for a species if a small number of specimens were captured for the thermal studies to provide a better estimation of the mean morphological variables for the species.

Each new researcher added to the project was trained in the use of either the Vernier calipers or digitizing system prior to data collection for the study. All individuals practiced obtaining measurements until such time as there was no statistical difference between the data they collected previously and data collected by previous researchers for the training species. Only after an individual demonstrated consistency when measuring a variable and their data did not differ statistically from data collected by previous individuals were the data saved and included in the analyses.

2.6. Statistics

Statistics are reported as mean ± standard deviation. Prism 4.0b was used to perform the regression analyses and produce the graphs. The species-specific means were used to represent the individual species in the graphs and determining the regressions. Because there were no phylogenetic influences between the previously studied thermal responses (Sanborn et al. 2017) and since a complete phylogeny of the included species has not been produced, phylogenetic comparative analyses were not applied during the correlation analyses.

3. Results

3.1. Summary data

All morphological, MFT and mass data are summarized in Supplementary Table 1. The 119 taxa have a live mass mean ranging from 80–3195 mg (n=1–298), mean wing length ranged from 1.51–5.72 cm (n=1–212), mean wingspan ranged from 3.53–13.29 cm (n=1–110), mean wing area ranged from 1.96–19.91 cm2 (n=1–42), and wing loading ranged from 38.69–187.64 mg/cm2. Mean MFT ranged from 15.5°C-23.2°C (n = 1–179).

3.2. Morphology

The morphological variables sampled scale with body size. The allometric relationship between wing length and body mass (Fig. 1) (F=851, d.f.=1, 117, P<0.0001, r2=0.8791), between wingspan and body mass (Fig. 2) (F=1000, d.f.=1, 117, P<0.0001, r2=0.8953), between wing area and mass (Fig. 3) (F=523.1, d.f.=1, 107, P<0.0001, r2=0.8380), and between wing loading and mass (Fig. 4) (F=186.7, d.f.=1, 107, P<0.0001, r2=0.6501) all show statistically significant regressions. The regressions between wing area and body mass (Fig. 5) (F=70.23, d.f.=1, 38, P<0.0001, r2=0.6489) and wing loading and body mass (Fig. 6) (F=365.6, d.f.=1, 38, P<0.0001, r2=0.9058) also show significant relationships in Magicicada cassinii.

Figure 1.

Figure 1.

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Allometric relationship between wing length and body mass in cicadas. y=−0.3501×0.3501, r2=0.8791, P<0.0001.

Figure 2.

Figure 2.

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Allometric relationship between wingspan and body mass in cicadas. y=−0.1303×0.3488, r2=0.8953, P<0.0001.

Figure 3.

Figure 3.

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Allometric relationship between wing area and body mass in cicadas. y=−0.9161×0.6253, r2=0.8380, P<0.0001.

Figure 4.

Figure 4.

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Allometric relationship between wing loading and body mass in cicadas. y=0.9161×0.3747, r2=0.6501, P<0.0001.

Figure 5.

Figure 5.

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Wing area as a function of body mass in Magicicada cassinii. y=0.003832x + 4.907, r2=0.6489, P<0.0001.

Figure 6.

Figure 6.

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Wing loading as a function of body mass in Magicicada cassinii. y=0.09974x + 22.55, r2=0.9058, P<0.0001.

3.3. MFT and morphology

The MFT also correlates with the flight system morphology. The relationship between MFT and body mass (Fig. 7) (F=6.767, d.f.=1, 117, P=0.0105, r2=0.05468), wing length (Fig. 8) (F=12.56, d.f.=1, 117, P=0.006, r2=0.09694), wingspan (Fig. 9) (F=12.53, d.f.=1, 117, P=0.0006, r2=0.09672), wing area (Fig. 10) (F=7.994, d.f.=1, 117, P=0.0055, r2=0.06395), and wing loading (Fig. 11) (F=4.088, d.f.=1, 117, P=0.0455, r2=0.03376) all show significant regressions. These correlations are lost within a single species as shown in the data for Magicicada cassinii. The relationship between MFT and body mass (Fig. 12) (F=0.811, d.f.=1, 39, P=0.3732), wing area (Fig. 13) (F=0.6705, d.f.=1, 38, P=0.4180), and wing loading (Fig. 14) (F=0.7841, d.f.=1, 38, P=0.3815) all lack statistical significance within the species.

Figure 7.

Figure 7.

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Minimum flight temperature as a function of body mass in cicadas. y=−0.0005695x + 19.80, r2=0.05468, P=0.0105.

Figure 8.

Figure 8.

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Minimum flight temperature as a function of wing length in cicadas. y=−0.5614x + 21.06, r2=0.0964, P=0.0006.

Figure 9.

Figure 9.

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Minimum flight temperature as a function of wingspan in cicadas. y=−0.2419x + 21.08, r2=0.09672, P=0.0006.

Figure 10.

Figure 10.

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Minimum flight temperature as a function of wing area in cicadas. y=−0.1000x + 20.10, r2=0.06395, P=0.0055.

Figure 11.

Figure 11.

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Minimum flight temperature as a function of wing loading in cicadas. y=−0.008624x + 20.18, r2=0.04647, P=0.0455.

Figure 12.

Figure 12.

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Minimum flight temperature as a function of body mass in Magicicada cassinii. y=−0.001468x + 21.57, r2=0.01778, P=0.4059.

Figure 13.

Figure 13.

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Minimum flight temperature as a function of wing area in Magicicada cassinii. y=−0.2946x + 22.91, r2=0.01734, P=0.4180.

Figure 14.

Figure 14.

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Minimum flight temperature as a function of wing loading in Magicicada cassinii. y=−0.01444x + 21.96, r2=0.02022, P=0.3815.

4. Discussion

4.1. Morphology

Analyses of the morphological data show the morphological variables each scale as predicted by geometric similarity. Wing length, wingspan, wing area and wing loading of cicadas as a group (Figs. 1–4) all show a significant correlation (P<0.0001) to body mass as would be predicted by geometric scaling, a relationship also seen within a single species (Figs. 5–6). Wing length and wing area show proportional relationships to mass in flying animals so that the corresponding wingspan and wing loading also show significant relationships (Greenwalt 1962; Dudley 2000; Ahlborn 2004). These data suggest there are no dramatic evolutionary changes in particular species that may influence the lift generated by the flight system of particular species.

4.2. Morphology and MFT

MFT shows a significant correlation (P=0.0006–0.0455) to mass, wing length, wingspan, wing area, and wing loading across but not within species (P>0.05). The slopes of these relationships are predicted by the Kutta-Joukowski theorem and the lift equations as longer wings, greater wing area and lower wing loading all lead to greater lift generation, even though the shape of cicada wings makes them relatively inefficient for generating lift and thrust even with their relatively high wing area (Hassanalian et al. 2017). Greater lift generating capacity would require a lower wing beat frequency to generate the lift needed to overcome gravitational pull. Since muscle contraction kinetics are temperature dependent (Josephson 1981), a cicada with greater lift generating structures would be able to produce the necessary lift to overcome gravity at lower temperatures. However, the low r2 values (0.03376–0.09694) of the regressions suggest that morphology has minimal influence on the MFT for each species.

MFT does not correlate significantly with body mass (P=0.4059), wing area (P=0.4180) or wing loading (P=0.3815) within Magicicada cassinii. The variability in MFT values obtained combined with potential damage to wings (specimens were collected late during the 1990 emergence) may have led to the statistically insignificant relationships obtained within the species. A similar between and within species relationships were found in Lepidoptera as well (Nève and Hall, 2016).\

5. Conclusions

We have shown that cicada flight system morphology scales as predicted based on geometric similarity but the low correlation coefficients suggest a minimal influence on minimum flight temperature. There are undoubtedly physiological adaptations that can influence MFT for specific species. For example, the relatively large wings of the species in the genus Platypedia Uhler, 1888 are used for sound production (Sanborn et al. 2002b) but also generate greater lift (vertical force) due to relatively the larger wing area resulting in lower minimum flight temperatures than species of similar size (Sanborn et al. 2017). Undoubtedly the small cicada Herrera umbraphila Sanborn and Heath, 2014 that inhabits the tropical cloud forest understory in Argentina and is the only known diurnal cicada that is a thermoconforming species (Sanborn et al. 2011c) would also require specific physiological adaptations to permit activity at low ambient and body temperatures. The low body temperature over which H. umbraphila is active probably led to specific biochemical adaptations to permit more efficient movement at lower temperatures. Similar adaptations are no doubt found among cicadas as a group as individual species adapt to specific environments so that the basic design of the flight system will permitting flight in these various habitats.

Supplementary Material

1

Highlights.

  • Minimum flight temperature in cicadas is not related to specific environments

  • Wing morphology determines lift and may influence minimum flight temperature

  • Cicada wing morphology scales as predicted by geometric similarity

  • Minimum flight temperature shows significant correlations to wing morphology

  • Minimum flight temperature appears dependent on specific physiological adaptations

Acknowledgements

AFS thanks the late Sr. John Karen Frei for university support of field expeditions to collect cicadas and equipment used in this study. AFS was supported financially by Barry University Mini-Grants. EAA and FF were supported by NIH-NIGMS MBRS Grant GM 45455 and THD was supported by MARC Grant 299420. The funding sources had no involvement in study design, collection, analysis or interpretation of data, writing of the report or decision to submit the article for publication.

Biography

Vitae

Allen F. Sanborn is a comparative physiologist with a Ph.D. in physiology from the University of Illinois at Urbana-Champaign. His research focuses primarily on cicadas where he has studied thermal biology, biodiversity, biogeography, taxonomy, systematics, ecology, and acoustic biology. He is a Professor of Biology at Barry University.

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Earl A. Allick graduated with a B.S. and M.S. from Barry University. He is currently an Assistant Principal for Miami-Dade County Public Schools.

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Sandee V. Apang graduated with a B.S. from Barry University and M.S. from Florida International University. She is currently a teacher for Miami-Dade County Public Schools.

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Izayana D. Castillo graduated with a B.S. from Barry University. She is a certified pharmacy technician working in Miramar, FL.

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Erica M. Cruz is about to graduate with a B.S. from Barry University. She will be enrolling in the Virginia-Maryland College of Veterinary Medicine training to become a veterinarian.

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Theophilus H. Davis (deceased) graduated with a B.S. from Barry University and earned an M.D. from the Medical University of South Carolina.

Cory H. Duncan graduated with a B.S. from Barry University and an M.D. from Florida State University. He is currently in private practice in Jacksonville, FL.

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Fanny Fierro graduated with a B.S. from Barry University and an M.S. from St. Thomas University. She is currently a teacher for Miami-Dade County Public Schools.

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Marla R.M. Gebaide graduated with a B.S. from Barry University and a D.C. from Palmer College of Chiropractic. She is currently in private practice in Northern Virginia.

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Abigael M. Luke graduated with a B.S. from Barry University and an M.D. from Northumbria University/St. Georges University. She is currently in private practice in St. Cloud, MN specializing in internal medicine, hematology and oncology.

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Maria L. Pacheco graduated with a B.S. and D.P.M. from Barry University. She is currently in private practice in Dalton, GA and surrounding areas.

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Daniel E. Paz-Castillo graduated with a B.S. from Barry University and an M.B.A from the Instituto de Estudios Superiores Administrativos. He the CEO of a health clinic in Caracas, Venezuela.

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Laura M. Perez graduated with a B.S. from Barry University and D.V.M. from Cornell University. She is in private practice in Latham, NY specializing in rehabilitation and acupuncture.

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Ana C. Poeck graduated with a B.S. from Barry University and V.M.D. from the University of Pennsylvania. She is in private practice in Boca Raton, FL.

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Adrian K. Seepersaud graduated with a B.S. and M.B.S. from Barry University and an MEDL from Florida Atlantic University. He is currently an assistant principal for the Palm Beach County Public Schools working towards a Ph.D.

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Carolina G. Valdes graduated with a B.S. from Barry University and an M.D. from the University of Puerto Rico, Ponce. She is currently a neurologist in private practice in Ft. Lauderdale, Florida.

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Footnotes

Conflicts of interest

2The authors declare no conflict of interests.

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