Running head: GREEN IGUANA INVASIVE POTENTIAL

ANALYSIS OF THE GREEN IGUANA’S (​IGUANA IGUANA)​ INVASIVE POTENTIAL

BASED ON ECOLOGICAL FUNCTION AND ECOSYSTEM CHARACTERISTICS

Bailey Daniels & Camila Guzmán

Nova Southeastern University
21 April 2020
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Abstract

Invasive species threaten native flora and fauna, and alter natural ecological processes.

The green iguana (​Iguana iguana​) has been anthropogenically introduced to new habitats in the

tropics and subtropics. This study examines how the​ ​biological niches of the green iguana, and

the ecological characteristics of the environments the species is introduced to, facilitate the green

iguana’s invasion. Results from a comprehensive literature review showed that the iguana’s

climatic compatibility, lack of predation, adaptability and diet all facilitate its ability to become

invasive in an introduced environment. Adequate knowledge on the effects of the introduction of

the species can help prevent local habitat degradation and promote better management practices.
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Introduction

Globalization and increased demand for commercial sale of ‘exotic’ plants and animals

has caused the introduction of foriegn species to ecosystems all over the world. Many

ecosystems now have a variety of exotic species, and countless suffer from exotic species that

have become invasive. A species becomes invasive when it begins to impact the native

ecological processes and competes with native species for resources. The introduction of exotic

species is most common in and around urban areas where people bring foriegn plants for

landscaping, and exotic animals as pets.

The green iguana (​Iguana iguana​) has been transported from its native habitat in South

America, as a result of the exotic pet trade of live reptiles. These iguanas have established

populations across the tropics and subtropics including South Florida, Puerto Rico, Japan and

Eastern Pacific Islands (Fujisaki, et al., 2010; Falcon, Ackerman, Curtis, 2013;

Burgos-Rodriguez, Aviles-Rodriguez, Kolbe, 2016). Given the species' prolific nature, green

iguanas often become invasive in new territory (Campbell & Maple, 2012). This study will

analyze the specific biological characteristics of the green iguana that facilitate its invasion, as

well as an analytical comparison of their natural habitat and their introduced environments. The

hypothesis tested in this study is: the biological niches of the green iguana (​Iguana iguana)​ , and

the ecological characteristics of the environments the species is anthropogenically introduced to,

facilitate the green iguana’s invasion.

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Methods

This assessment was carried out through a literature review. The review searched for

literature about the species’ diet and seed dispersal, adaptability, predator-prey relationships,

human interactions, habitat comparison and overall invasive potential. These research topics give

the insight needed in order to evaluate how the species can become invasive. This section

discusses the types of research procedures that were used in each of the articles reviewed.

Diet and Seed Dispersal​:

Govender, Muñoz, Camejo, Puente-Rolón, Cuevas and Sternberg worked together on

“An Isotopic Study of Diet and Muscles of the Green Iguana (​Iguana iguana)​ in Puerto Rico”

(2012). The study samples two sites, a natural and a managed site, along the northeastern coast

of Puerto Rico. A total of 42 iguanas were sampled, 16 females and 25 males. Data on individual

attributes (mass, length, sex etc.) was collected for each iguana in addition to tongue and muscle

tissue samples. The entirety of the digestive system was collected from all iguanas and inspected

microscopically. Govender, et. al. (2012) collected and analyzed the carbon (C) and nitrogen (N)

isotopes of leaves and crabs identified from gut content. An isotopic analysis was done by

placing samples into an automated elemental analyzer and pyrolized. The gases were then led

​ C and 15​
into a mass spectrometer and analyzed for 13​ ​ N abundance. Then an analysis of variance

(ANOVA) was run to determine the difference in individual attributes.

Furthermore, in a study titled, “​Effects of invasive Green Iguanas (​Iguana iguana​) on

seed germination and seed dispersal potential in southeastern Puerto Rico”​, scientists

Burgos-Rodríguez, Avilés-Rodríguez, & Kolbe, (2016) processed 258 green iguana scat samples
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to test the effects of digestion on seed germination and growth rate. Conducted near a mangrove

forest area at the Humacao Natural Reserve in SE Puerto, scat samples were mainly found on the

ground, and GPS coordinates were taken for each sample. After collection, samples were stored

outside in the sun and later dissected, using a dissecting microscope and magnifying glass, after

one week of drying out. The extracted seeds (native and exotic) were planted under the same

garden conditions and cultivated. Germination rate and the speed of growth were recorded

(Burgos-Rodríguez et al., 2016).

Another factor considered in ​I. iguana’​ s impact on seed dispersal is the size of an

individual’s home range, or the extent of area that they travel regularly. In the article “​Home

Range Size and Potential for Exotic Seed Dispersal by Green Iguanas (​Iguana iguana)​ in

Southern Florida”,​ researchers Campbell and Maple (2012) captured a total of 14 green iguanas

from two Florida sites: five from Morikami Japanese Gardens, located in Boca Raton and nine

from Palm Beach Zoo in West Palm Beach. After capture, a blood sample was taken and each

individual was tagged. Green iguanas were tracked and their home ranges were calculated using

ArcGIS and the minimum convex polygon method. Home ranges were compared using an

ANOVA analysis. After, researchers compared their data to primary source literature about the

species’ home ranges in their native environments (Campbell & Maple, 2012).

Adaptability​:

Balasko and Cabanac performed an experiment to identify the behavior of iguanas in a

situation of conflict between two resources. In “Behavior of Juvenile Lizards (​Iguana iguana)​ in

a Conflict between Temperature Regulation and Palatable Food”, Balasko and Cabanac (1998)
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housed three juvenile green iguanas bought from a pet store. The iguanas were housed in a

wooden terrarium inside a climatic chamber where the ambient temperature could be regulated.

At the ‘home’ corner of the terrarium, two large bowls of water and ‘standard’ food were always

present in addition to a 250-W infrared lamp to provide heat from 6am to 10pm. The ambient

temperature was maintained at 34-35 degrees celsius. Opposite to the ‘home corner’, lettuce (a

palatable food) was placed five days a week. The iguanas were given a ten week period to adjust

to the environment. During the first experiment, the time interval between lettuce offerings

ranged from 1 to 8 days and the ambient temperature stayed at a constant 20℃. The second

experiment decreased the ambient temperature to 20, 15, 10, 5, 0 and -5℃, while the home

corner maintained an ambient temperature of 34℃.

In the study “Genetic Evidence of Hybridization between the Endangered Native Species

Iguana delicatissima​ and the Invasive ​Iguana iguana​ (Reptilia, Iguanidae) in the Lesser Antilles:

Management Implications” ​Vuillaume, Valette, Lepais, Grandjean, and Breuil ​(2015) ​collected a

sample size of 133 iguanas (59 ​I. delicatissima, 4​ 7 ​I. iguana​ and 27 hybrids). The study defined

hybrid status by: “both parental species were observed during the fieldwork according to

diagnostic characters - and individuals that present various intermediate conditions, for the whole

set of characters and/or exhibit a mosaic of both parental or intermediate characters” (Vuillaume

et al. 2015). The tips of the tails of each iguana were cut and DNA was obtained from animals

caught in the field by hand or by nooses. The morphology of each iguana was described using

characters depicted by Breuil and pictures were taken.

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Predator-Prey Relationships​:

To better understand the relationship between carnivorous predators and herbivorous

consumers, the study, “Ecological Meltdown in Predator-Free Forest Fragments”, by ​Terborgh,

Lopez, Percy Nanez, Rao, & et, al. (2001)​ used vertebrate and invertebrate animal surveys,

conducted between 1994 and 1995, to quantify populations on 12 island fragments. The islands

ranged in size and were classified as “small”, “medium”, and “large”. At the same time,

vegetation and small tree growth were monitored by the scientists on the small and medium

islands, as well as on land. They utilized sheltered and exposed vegetation plots to test the

hypothesis that abiotic processes were impacting vegetation growth. Results were analyzed using

an ANOVA statistical test to determine if there was a significant difference between the

experimental groups (Terborgh, et. al., 2001).

Human Interactions​:

Carlos G. García-Quijano, Tomás A. Carlo and Javier Arce-Nazario conducted

interdisciplinary research between wildlife ecologists and anthropologists to uncover the

interactions between people and the introduced green iguana in Puerto Rico. The article “Human

Ecology of a Species Introduction: Interactions between Humans and Introduced Green Iguanas

in Puerto Rican Urban Estuary” (2011) focuses on the social-ecological changes, the human

perception and economic interactions resulting from the introduction of the species. The study

mapped the distribution of iguanas, conducted a literature review of history of the introduction of

the species, and interviewed residents. The mapping of distribution was used to perform an
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Ordinal Logistic Regression Analysis to model the iguana detections to the nearest vegetation

edge and the nearest human structure.

Habitat Comparison

In the source, “Biology and Impacts of Pacific Island Invasive Species. 10. Iguana

iguana, the Green Iguana (Squamata: Iguanidae)”, Falcón, Ackerman, Recart, & Daehler (2013)

reviewed primary source literature on a variety of environments where green iguanas are known

to exist, native and nonnative. The scientists also developed a model, using the Maximum

Entropy Niche-Modelling Algorithm method, to predict future impacts to areas the green iguana

have been introduced (Falcón, et al., 2013).

Invasive Potential​:

The authors of, “​Risk Assessment of Potential Invasiveness of Exotic Reptiles Imported

to South Florida”, ​Fujisaki, et.al (2009) considered specific factors that may be used to predict

the establishment success of nonnative reptiles. They identified 17 key variables including

taxonomic order, maximum temperature match between native ranges and S. Florida, animal sale

price, and manageability. The model the authors developed was then used to determine the

success of 33 reptile species that were recently introduced to Miami and St. Petersburg (Fujisaki,

et. al., 2009).

“Just Green Iguanas? The Associated Costs and Policy Implications of Exotic Invasive

Wildlife in South Florida” by Arthur Sementelli, Henry Smith, Walter Meshaka Jr. and Richard

Engeman (2008) conducts a literature review of the wide range of effects ​I. iguana​ has on South
GREEN IGUANA INVASIVE POTENTIAL 8

Florida’s environment. The study uses literature focused on population density, biotic damage

and dispersal, and transportation and infrastructure issues.

Results

Diet and Seed Dispersal​:

In the isotopic study of diet and muscles, Govender et al. (2011) found that there was a

significant difference of mass and snout-vent length (SVL), as males were larger than females.

​ C and 15​
There was no significant difference of 13​ ​ N isotopes between the sexes, and no found

​ C and 15​
correlation between size of the iguana and 13​ ​ N isotope values. The gut content revealed

that ​I. iguana​ eat mostly leaves however in some individuals, traces of insects, crabs and

occasionally snails, were identified. Both native and nonnative plant species were found in the

gut.

Additionally, the experiment conducted at ​Humacao Natural Reserve​ in 2016 f​ound that

digestion by green iguanas reduced the time needed for seeds of the ​Ficus spp.​ species to

germinate, had no effect on native ​Annona glabra​ seed germination and had variable impacts on

invasive ​P. pterocarpus​ and ​Pterocarpus spp. (​ ​Burgos-Rodríguez et al., 2016)​.​ For invasive

plant species, seed digestion decreased the time it took for seeds to germinate, but also prevented

some seeds from ever germinating.

Figure 1 shows the frequency of seeds found in scat samples at different distances from

two species of parental trees (a) ​Anona glabra​ and (b) ​Ficus spp.​ The researchers noted that the

size of the tree’s canopy (0.8m - 2 m) exaggerates the frequency of seeds found closest to the

tree, and that data should be disregarded. Green iguanas spread seeds up to 20 m or 50 m away

from parental canopies depending on the tree species (​Burgos-Rodríguez, et al., 2016)​.
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Fig. 1

Figure 1: This figure obtained from, “Effects of invasive Green Iguanas (Iguana iguana) on seed
germination and seed dispersal potential in southeastern Puerto Rico” by ​Burgos-Rodríguez et al. (2016),​ depicts the
frequency of seeds, found in 122 green iguana scat samples, at various distances from the parental tree. Graph (a)
refers to ​Anona glabra​ and (b) represents ​Ficus spp.

Meanwhile, when Campbell & Maple compared ​I. iguana​’s exotic and native home

ranges, or areas that they cover regularly, ​their results​ showed that average ranges in South

Florida (2,393.2 m​2​) were significantly smaller than what was found in literature about native

areas (9,605.2 m​2​). Since they tend to have smaller home ranges in S. Florida, researchers

concluded that here green iguanas probably do not spread exotic seeds far distances, but rather

spread a higher density of seeds in a smaller area (Campbell & Maple, 2012). However, their

results showed that home range sizes varied between study sites: Morikami Japanese Gardens
GREEN IGUANA INVASIVE POTENTIAL 10

had home range sizes that ranged from 46.42 m​2​ to 10,943.5 m​2​ and Palm Beach Zoo ranged

from 354.75 m​2​ to 12,604 m​2​ (Campbell & Maple, 2012), which suggests that there may be other

factors that have an impact on how far green iguanas travel day to day.

Adaptability​:

Balasko and Cabanac (1998) described the iguanas' behavior as “time budgeting”. The

first experiment found that ​I iguana​ didn’t increase the duration of their stay near the feeder

when the number of days without lettuce was increased. The second experiment showed that the

duration of lettuce meals and the number of lettuce meals decreased as the ambient temperature

decreased. The number of visits to the lettuce feeding station also decreased with the ambient

temperature.

In the study of genetic evidence of hybridization, Vuillaume et al (2015), found ample

evidence of the occurrence of hybridization between ​Iguana delicatissima a​ nd ​I. iguana​ in

habitats where the two species are syntopic. The study also found that hybrids are able to

reproduce with parental species as well between hybrids. Both F1 and F2 hybrids were identified

in the study, meaning that iguanas fall under the anthropogenic hybridization of complex

admixture when F1 hybrids are fertile. The results indicate that hybridization can occur in both

directions, and the level of hybridization differs across different habitats.

Predator-Prey Relationships​:

In their article, authors ​Terborgh, Lopez, Percy Nanez, Rao, & et, al. (2001)​ state, “​...we

have shown that the absence of ​predators ​consistently frees certain consumers to increase many
GREEN IGUANA INVASIVE POTENTIAL 11

times above "normal," unleashing a trophic cascade”. ​Specifically, their results suggested that the

recruitment of small trees and shrubs on these islands was significantly lower than recruitment

on larger land masses that host predators (​Terborgh, Lopez, Percy Nanez, Rao, & et, al., 2001).

Human Interactions​:

The results from the interdisciplinary research by García-Quijano et al. (2011) show that

species introduction is a human ecological phenomenon, that can be traced back to early

homosapien migration. The literature review found evidence that shows that historically, when

homosapiens migrated, they always transported fauna and flora with them. When the distribution

of ​I iguana​ was mapped, the study found that they are not distributed randomly. The distance to

the nearest vegetation edge and the distance from human settlements were good predictors of

sighting the iguanas in mangroves. The study found that there were higher concentrations of

iguanas along the edges of the mangroves, and almost none were present in pure mangrove

stands. There was a spatial association between iguana populations and human settlements as

they have the ability to adapt and survive in landscapes heavily altered by humans.

García-Quijano et al. (2011) found that the local people’s experiences with the green iguana vary

from disasters to the ecosystem (loss of biodiversity and livelihoods) to new economic

opportunities like tourism. The study identifies that given ​I iguana's l​ arge size, and its ability to

be easily tamed have brought tourism opportunities.

GREEN IGUANA INVASIVE POTENTIAL 12

Habitat Comparison:

Results showed that high climatic compatibility exists between many tropical Pacific and

Carribean islands and the​ I. iguana​’s native environments. In their review, ​Falcón et al. (2013)

found literature suggesting that​ conservation efforts in Puerto Rico for native plants, specifically

trees, had been thwarted by green iguanas feeding on young tree saplings (López-Torres et al.

2012). Specifically, gut contents have shown that the green iguanas selectively eat red and black

mangroves (Gómez-Carrasquillo, Pérez-Reyes, Hernández-García, & Thomas, unpubl. data), and

have often degraded these tree species in certain areas (Carlo and García-Quijano, 2008).

Additionally, in Fiji, green iguanas are known to consume various native plants and notably

plants that are vital to the local community as well as major exports (Harlow and Thomas 2010,

Thomas et al. 2011). While in South Florida, green iguanas have been found to consume

numerous plant species, native and exotic (Krysko et al., 2007).

Invasive Potential​:

Sementelli et al. (2008) found that ​I. iguana​ has had great success in areas of South

Florida where there are few natural predators and have less success in shared habitats. The study

identifies that areas that have a decrease in raccoons and foxes, have also had an increase in

green iguana populations. When predation is absent, bursts in ​I. iguana​ populations occur

quickly as a result of the species' early sexual maturity and large clutch sizes. This gives green

iguanas the potential to rapidly flood any sustainable habitat. Additionally, evidence was found

that the green iguana put native species at risk, like the Florida burrowing owl, gopher tortoises

and egrets. Sementelli et al. (2008) also identifies the damage that the intrusion of the species has
GREEN IGUANA INVASIVE POTENTIAL 13

on native plant communities as the green iguana acts as an invasive plant disperser.

Moreover, in their risk assessment, Fujisaki et. al. discovered that South Florida had the

highest climatic compatibility with native environments of the 33 species of exotic reptiles that

they analyzed (2009). However, the data they chose to focus on was from northern Florida.

Nonetheless, their results showed that insectivores had the best chance of establishment (88% as

juveniles or adults), compared to omnivores (33% as juveniles and 43% as adults), and

herbivores (25% and 31%, respectively) in this environment (Fujisaki, et. al., 2009).

Discussion

In order for a species to thrive in new environments it must be able to adapt easily and

quickly. The ​I. iguana ​has a variety of biological characteristics that allow it to assimilate to

introduced environments. The results show that despite the species being primarily herbivorous,

it is an “opportunistic omnivore” (Govender, et al., 2012) in nonnative environments. Its

flexibility in diet allows it to adapt easily to different habitat constraints, and could potentially

give the species an opportunity to evolve its diet entirely. However, the plant heavy diet also has

an effect on the native ecosystem, as the iguana contributes to seed dispersal. A study found that

the green iguana can affect the germination rate of some native species and even prevent

germination entirely in other plant species (Burgos-Rodríguez et al., 2016). The species also has

the ability to disperse seeds of exotic and invasive plants, although it is unlikely to spread them

outside of their home range (Campbell & Maple, 2012). The change in germination of local

plants, and the increase of invasive plants within the home range can drastically change the local

ecosystem. The spread of invasive plants within their home range may reinforce the
GREEN IGUANA INVASIVE POTENTIAL 14

establishment of exotic plant species, such as Brazilian pepper (often found in their diet), making

them harder to eradicate.

Nonetheless, its heavy grazing on local vegetation could be detrimental to local

ecosystems, especially if green iguana populations outgrow the resources available. ​I. iguana

populations have the ability to grow rapidly given the species young sexual maturity. Male

iguanas reach maturity at 20 months, females at about 31 months, and can lay clutches of up to

50 eggs (Sementelli et al., 2008). The species' prolific nature and long life expectancy of 15-20

years, gives the green iguana the potential to overpopulate any sustainable environment.

Additionally, the results showed that green iguanas can successfully reproduce with similar

species (​I. delicatissima)​ , an adaptation that guarantees its survival through hybrids. This can

result in direct impact to the native iguana species population, as was observed in the Lesser

Antilles (Vuillaume et al., 2015). The ​I. iguana​ populations in the Lesser Antilles are more

powerful than the local ​I. delicatissima​, and resulted in the displacement of ​I. delicatissima

males. Additionally, the large size of the hybrids can increase competition for the male ​I.

delicatissima​ and further hurt the endemic species population (Vuillaume et al., 2015). The green

iguana also displayed great ability to adapt when in conflicting conditions. The iguanas

demonstrate the ability to prioritize when temperatures drop, and their “time management”

behavior gives them the ability to stabilize and establish themselves (Balasko & Cabanac, 1998).

Along with analyzing the ecological niches of ​I. iguana t​ hat facilitate the species’

potential to be invasive, specific habitat characteristics about their native and nonnative

territories were compared. The results identified in this paper indicate environmental and

anthropogenic factors such as climatic compatibility, human interference, and lack of natural
GREEN IGUANA INVASIVE POTENTIAL 15

predators promote the rapid establishment of ​I. iguana i​ n introduced areas. Specifically,

García-Quijano et al. (2011) showed that populations of ​I. iguana ​are often associated with

degraded environments, as they can quickly adapt to habitat changes​. For this reason, they have

the potential to outcompete native animal species that are more sensitive to anthropogenic

impacts. Additionally, the experiment by ​Terborgh et al. ​(2001), demonstrated that herbivores

and opportunistic species can thrive and quickly overpopulate areas that have little to no natural

predators. Reduced home ranges in nonnative areas were largely attributed to the lack of

predators and the availability of food, because the green iguanas likely did not have to travel as

far to sustain themselves ​(Campbell & Maple, 2012)​. Lack of predation in South Florida, Puerto

Rico, and other introduced locations provide the necessary environmental conditions for ​I.

iguana​ populations to expand rapidly.

In light of the high invasive potential of the ​I. iguana,​ having adequate regulations in

place is vital to managing and limiting its impact. Educating the community and government

officials could reduce the influx of exotic pets that become invasive, and lead to better policy

regarding invasive species. Enforcing penalties for the release of green iguanas into natural

habitat can prevent the establishment of a population and discourage exotic pet trade (Sementelli

et al., 2008). Some management suggestions include sterilization of the species and destruction

of eggs in effort to stop population growth, construction of artificial nests (Falcón et al., 2013),

and trapping and removal from state parks (Sementelli et al., 2008). Without management of the

species, the green iguana is likely to establish itself in tropical and subtropical introduced

environments and heavily alter the native ecosystem.

GREEN IGUANA INVASIVE POTENTIAL 16

Moving forward, this study would benefit from more quantitative data comparing the

environmental conditions between native territories and introduced areas. One way to do this

would be to obtain information such as precipitation levels and temperatures from databases that

survey these areas. Doing this for South Florida may be reasonable, but accessing that

information for Pacific, Carribean, and South American territories may prove difficult. Ideally,

another way to expand this research question would be to conduct a field study in a S. Florida

location, and compare the results to the literature examined here. Critical and extensive data is

necessary to determine the full extent of the ​I. iguana s​ pecies’ impacts to native ecosystems in S.

Florida. Although invasive species management plays a crucial role in conservation and

preservation, the number of studies that focus on the potential invasion of the ​I. iguana​,

especially in S. Florida is lacking, and this paper aimed to bring awareness to that deficiency.
GREEN IGUANA INVASIVE POTENTIAL 17

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