Ctenosaura similis (black spiny-tailed iguana)

Pictures

Picture	Title	Caption	Copyright
Title Adults Caption Ctenosaura similis (black spiny-tailed iguana); male (centre) with two females. Barra Honda national park, Costa Rica. February 2007. Copyright ©Christian Mehlführer-2007, Vienna, Austria/via wikipedia - CC BY 2.5	Adults	Ctenosaura similis (black spiny-tailed iguana); male (centre) with two females. Barra Honda national park, Costa Rica. February 2007.	©Christian Mehlführer-2007, Vienna, Austria/via wikipedia - CC BY 2.5
Title Female Caption Ctenosaura similis (black spiny-tailed iguana); female. Barra Honda national park, Costa Rica. February 2007. Copyright ©Christian Mehlführer-2007, Vienna, Austria/via wikipedia - CC BY 2.5	Female	Ctenosaura similis (black spiny-tailed iguana); female. Barra Honda national park, Costa Rica. February 2007.	©Christian Mehlführer-2007, Vienna, Austria/via wikipedia - CC BY 2.5
Title Male Caption Ctenosaura similis (black spiny-tailed iguana); male. Caye Caulker, Belize. March 2016. Copyright ©Simon Pierre Barrette-2016/via wikipedia - CC BY-SA 4.0	Male	Ctenosaura similis (black spiny-tailed iguana); male. Caye Caulker, Belize. March 2016.	©Simon Pierre Barrette-2016/via wikipedia - CC BY-SA 4.0
Title Male Caption Ctenosaura similis (black spiny-tailed iguana); close view of a male. Barra Honda national park, Costa Rica. February 2007. Copyright ©Christian Mehlführer-2007, Vienna, Austria/via wikipedia - CC BY 2.5	Male	Ctenosaura similis (black spiny-tailed iguana); close view of a male. Barra Honda national park, Costa Rica. February 2007.	©Christian Mehlführer-2007, Vienna, Austria/via wikipedia - CC BY 2.5

Identity

Preferred Scientific Name

• Ctenosaura similis (Gray, 1831)

Preferred Common Name

• black spiny-tailed iguana

International Common Names

• English: black ctenosaur; black iguana; black spiny-tailed iguana; common spiny-tailed iguana; ctenosaur; Gray’s spiny-tailed iguana
• Spanish: iguana-espinosa rayada

Local Common Names

• Belize: wish willy
• Colombia: ishilly
• Costa Rica: garrobo; iguana negra
• Germany: gemeiner schwarzleguan; schwarzer leguan
• Nicaragua: garrobo negro

Summary of Invasiveness

Ctenosaura similis, commonly known as black spiny-tailed iguana, is the largest and most widely distributed ctenosaur. Its native range stretches from southern Mexico to Panama where is exploited for food and traditional medicine. It generally is found in seasonally dry, lowland habitats but can occur in sites up to 900 m elevation. It is often found in close proximity to human activity. The capacity of C. similis to co-exist in altered environments coupled with its high fecundity has contributed to its successful establishment and spread in Florida, USA, where it consumes valuable horticultural plants, invades dwellings, and threatens imperilled native species through predation and usurpation of burrows. The invasive range of C. similis also includes Venezuela and the Colombian islands of Providencia and San Andres.

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Metazoa
•         Phylum: Chordata
•             Subphylum: Vertebrata
•                 Class: Reptilia
•                     Order: Sauria
•                         Family: Iguanidae
•                             Genus: Ctenosaura
•                                 Species: Ctenosaura similis

Notes on Taxonomy and Nomenclature

Based on morphological and genetic characteristics, Köhler et al. (2000) proposed grouping the 13 Ctenosaura species into 3 subgenera:

Subgenus Ctenosaura Wiegmann, 1828, status nov. - C. acanthura, C. hemilopha, C. pectinata and C. similis.

Subgenus Loganiosaura nov. - C. bakeri, C. melanosterna, C. oedirhina and C. palearis.

Subgenus Enyaliosaurus Gray, 1845 status nov. - C. alfredschmidti, C. clarki, C. defensor, C. flavidorsalis and C. quinquecarinata.

This scheme was subsequently considered untenable, and 18 species of Ctenosaura are now recognized (Iguana Taxonomy Working Group, 2011). According to the Iguana Taxonomy Working Group (2011), “A well-resolved phylogenetic hypothesis of all taxa in this genus is sorely needed.”

Hybridization between C. similis and C. bakeri has been documented on Utila Island in Honduras (Gutsche and Köhler, 2008; Pasachnik et al., 2009).

Description

Considered to be a “giant” iguanid lizard, with adult males approaching 500 mm snout-vent length (SVL) and females approaching 350 mm SVL in their native range (Fitch and Henderson, 1978).

Hatchlings are 43-60 mm SVL (Fitch and Henderson, 1978; Avery et al., 2014). In their native range, marked individuals grew at rates of 1.8-3.4 mm per week (SVL), and first-year males and females averaged 188 mm and 177 mm SVL, respectively (Fitch and Henderson, 1978).

C. similis eggs are elongated and soft-shelled, measuring about 27 x 18 mm. Fat reserves contribute directly to yolk deposition, follicle growth, and clutch size (Fitch and Henderson, 1978). In Florida, USA, the amount of abdominal fat in female ctenosaurs ranged from 0.5-114.5 g, or 0.3-13.4% of body mass (Avery et al., 2014).

Distribution

The native range of C. similis extends on the Atlantic versant from the Isthmus of Tehuantepec southward to northeastern Nicaragua, and on the Pacific versant from the Isthmus of Tehuantepec southward to Panama (Ko¨hler, 2003). It occurs in Mexico and throughout much of Central America (The Reptile Database, 2016).

This species was first introduced to Florida, USA, during the 1970s and has expanded its range from Gasparilla Island onto the adjacent mainland and islands and throughout the southwestern coast of Florida from Manatee County southward to Collier County. At the same time it was introduced on Key Biscayne in Miami (Butterfield et al., 1997; Krysko et al., 2009). C. similis is also found elsewhere on Florida’s southeastern coast in Martin and Broward counties (Krysko et al., 2011b), as well as in the Florida Keys on Big Pine Key and No Name Key, Monroe County (Enge et al., 2006; Krysko et al., 2011a; Anderson and Enge, 2012).

C. similis has been introduced to Colombia, Carribbean islands of (San Andres and Providencia islands) (Tamsitt and Valdivieso, 1963) and Venezuela (Barrio-Amorós and Rivas-Fuenmayor, 2008; Flores and Esqueda, 2008).

Distribution Table

The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

History of Introduction and Spread

Results of the genetic analysis by Nuñez (2016) are consistent with multiple introductions within Florida, USA. The Gasparilla Island population possibly originated from three animals kept as pets and then released around 1979 (Krysko et al., 2003). The population, which comprises a single haplotype, now extends from Sarasota south to Naples 180 km along Florida’s southwest coast. Ctenosaurs move freely off Gasparilla Island via causeways, are carried off the island incidentally by vehicles, and are taken purposely off the island by people. C. similis now occurs on Keeywadin Island, Little Marco Island, Cayo Costa, and probably other islands as a result of purposeful transport and releases by residents (Krysko et al., 2003).  In Collier County a private landowner released between 5-30 lizards on his property on the southern end of the Keewaydin Island in 1995 (Krysko et al., 2003). The Key Biscayne population in Miami-Dade County originated from animals escaped or released from the Crandon Park Zoo when it closed in 1979 (Townsend et al., 2003b; Krysko et al., 2010). This population seems to have remained on Key Biscayne and not spread to the mainland or nearby islands. The dates and invasive pathways of other introductions in Florida are not known. Genetic analyses indicate that ultimately all Florida populations derived from Honduras (Nuñez, 2016).

Origins of introduced populations in Venezuela and on the Colombian islands of Providencia and San Andres are uncertain. In Colombia, first museum collections occurred in before 1933 (Barbour and Shreve, 1934); but introductions by humans to these islands likely occurred many years prior (Dunn, 1945; Rueda-Almonacid, 1999; Forero-Medina et al., 2006). On the other hand, Hedges (1996, p.105) concludes that these island populations “originated by dispersal from Central America in the Quaternary”. In Venezuela, C. similis was first noticed in 1998 inhabiting gardens, parking lots and parks.

Introductions

Risk of Introduction

Over time, the number of C. similis imported into the USA has declined; just 76 were imported in 2008 (Stephen et al., 2011). However, the presence of thousands of free-living ctenosaurs in Florida means that they can be trapped and moved to new locations. Deliberate releases have occurred on Keeywadin Island, and likely have occurred on Cayo Costa and other islands as well (Krysko et al., 2003).

The recent appearance of ctenosaurs in Venezuela (Barrio-Amorós and Rivas-Fuenmayor, 2008; Flores and Esqueda, 2008), presumably through hitchhiking on ships, demonstrates the on-going potential for the species to spread to new areas.

Habitat

Dry lowland scrub, woodland, grassland, beach dunes. Especially dependent on riparian woodlands in Guanacaste Province, Costa Rica where the 4-6 months dry season restricts vegetation and moisture to the riparian zones (van Devender, 1982). Also frequents mangroves in intertidal areas. Adapts well to disturbed areas, parkland, and human habitation (Henderson, 1973; González-García et al., 2009).

Habitat List

Biology and Ecology

Genetics

A phylogeographic study of C. similis throughout its native range is in progress (AS Pasachnik et al., Institute for Conservation Research, San Diego Zoo Global, California, USA, unpublished data).

Genetic analyses of C. similis in Florida, USA, revealed a set of 5 haplotypes (Nuñez, 2016). One haplotype was uniquely associated with the ctenosaurs population in southwestern Florida that ranges from Sarasota southward to Naples, and one came from a specimen found on No Name Key in the Florida Keys. The remaining haplotypes were from populations in Broward and Miami-Dade counties, in southeastern Florida. Each of the 5 haplotypes in Florida was most closely aligned with Honduran populations in the native range of C. similis.

Hybridization between C. similis and C. bakeri has been documented on Utila Island in Honduras (Gutsche and Köhler, 2008; Pasachnik et al. 2009).  

Reproductive Biology

In the introduced population on Gasparilla Island in Florida, USA, testes size peaked mark­edly during January to April, after which there was rapid tes­ticular regression (Avery et al., 2014). Similarly, during January to April, 67.3% of all females contained vitellogenic follicles or oviducal eggs, compared to 3.8% during the rest of the year. During April to June, 81% (199 of 245) of females of reproductive size (>185 mm snout-vent length, SVL), possessed either oviducal eggs, vitellogenic folli­cles, or a convoluted oviduct indicating previous laying. Among females with SVL ≥ 235 mm, 99 of 100 exhib­ited evidence of reproduction. There is no evidence of multiple clutches per year. Clutch size, as determined by the number of oviducal eggs, averaged 31.6 (SE = 1.7). Clutch size with­in size classes did not differ from that recorded in Nicaragua (Fitch and Hen­derson, 1978). Clutch mass (mean ± SE) as a per­centage of female body mass was 18.0% ± 0.7%, and did not vary with SVL.

On Gasparilla Island, hatchlings (minimum 43 mm SVL, 2.75 g body mass) were observed from the last week of July until early October, with 97.5% of the hatchling iguanas encountered in August and September (Avery et al., 2014). Hatchlings exhibited a growth rate of approximately 6.8 mm/month.

Reproduction is seasonal. Egg-laying takes place in the dry season, and hatching occurs just before the onset of the wet season. Thus, in Nicaragua, eggs are laid during February to March and hatching is mostly in June (Fitch and Henderson, 1978). In Florida, USA, egg-laying occurs in April-May, with hatching in July to August (Avery et al., 2014).

Populations of C. similis are female-biased (Fitch and Henderson, 1978). In Florida, 57% of 1321 animals necropsied were female (Avery et al., 2014). One female collected on 27 March 2002 on Key Biscayne had 82 well-developed eggs in her oviducts (Krysko et al. 2003).

Most females reach sexual maturity in their second year at about 200 mm SVL. Clutch size increases steadily with age and size; a maximum clutch of 88 eggs has been reported (Fitch and Henderson, 1978). In Florida, USA, clutch size averaged 32 (n = 56), with a range of 6-62 (Avery et al., 2014).

Physiology and Phenology

The very concentrated breeding season (February to April) characteristic of the species in its na­tive range (Nicaragua; Fitch and Henderson, 1978) oc­curs 2 months later (April to June) in the introduced range (Florida; Avery et al., 2014). C. similis appears to grow more slowly in Florida, USA, than in Central American sites 1500-1800 km farther south. Fitch and Henderson (1978) report that in Nicaragua hatchlings grew to >140 mm SVL and >80 g body mass within 6 months. Hatchlings on Gasparilla Island, Florida grow much more slow­ly, averaging just 91.0 mm SVL and 21.7 g body mass after 6 months.

The Florida population is subjected to a very different tem­perature regime than native C. similis. In par­ticular, mean minimum monthly temperatures in Managua, Nicaragua during winter (December to February) range from 19.7-21.2 °C, whereas on Gasparilla Island during the same months mean monthly temperatures range from 10.9-12.8 °C. The colder winter temperature regime not only reduc­es activity but also limits growth rate and could retard sexual maturation.

Longevity

In captivity, 22.4 years (The Animal Ageing and Longevity Database, 2016). Estimated maximum observed age of 9 years among free-ranging females (Fitch and Henderson, 1978).

Activity Patterns

Winter activity of introduced populations in Florida is greatly reduced relative to warmer parts of the year.

Population Size and Density

Reportedly common in appropriate habitat throughout most of its range, although some populations can be locally depleted due to exploitation for food and medicinal uses (Savage, 2002). In Parque Nacional Palo Verde, Costa Rica, Mora (2010) reported the population density to be 7.1 ctenosaurs/ha.

Nutrition

The species exhibits an ontogenetic dietary shift as individuals become increasingly more herbivorous with age (van Devender, 1982; Krysko et al., 2009). The shift to plant-dominated diet is accompanied by changes in dentition (Montanucci, 1968). In C. pectinata, this ontogenetic dietary shift is also reflected in the gut morphology of the species (Durtsche, 2000), but similar research has not been conducted on the physiology of C. similis. Regardless, C. similis is a dietary opportunist and will eat whatever it can catch. Even though diets of larger, older individuals consist mostly of plant materials, they are carnivorous when given the chance (Avery et al., 2009).

Natural Food Sources

Climate

Latitude/Altitude

Natural Enemies

Notes on Natural Enemies

Hatchlings are preyed upon by conspecifics (Krysko et al., 2009; ML Avery, National Wildlife Research Center, Gainesville, Florida, USA, unpublished data). Boa constrictor and other snakes prey on adult and juvenile ctenosaurs (Mora-Benavides, 1987; Farrollo et al., 2010). In Costa Rica, lyre snakes, Trimorphodon biscutatus, appear to specialize as predators of C. similis (van Devender, 1982).

Means of Movement and Dispersal

Natural Dispersal

Daily movements tend to be <100 m (Henderson, 1973; Fitch and Henderson, 1978), but movements up to 200 m by juveniles occur (van Devender, 1982). It is clear that ctenosaurs can disperse short distances on land, and C. similis is capable of swimming. Also, sightings of ctenosaurs along causeways connecting Gasparilla Island to mainland Florida, USA, lend credence to the idea that the species is capable of dispersing unaided from the island.

Accidental Introduction

Short- and long-distance dispersal of C. similis has been facilitated, and probably continues to be, by hitchhiking on vehicles and ships (Krysko et al., 2003; Barrio-Amorós and Rivas-Fuenmayo, 2008). The rate or frequency of such events is unknown, but even a single introduction event can potentially produce serious long-term consequences.

Intentional Introduction

Ctenosaurs have been intentionally transported from their native range to offshore Caribbean islands (Dunn, 1945; Rueda-Almonacid, 1999; Forero-Medina et al., 2006) and to Florida, USA (Krysko et al., 2003). Furthermore, within Florida, subsequent purposeful transport and introductions have exacerbated the potential impacts of ctenosaurs on native fauna (Krysko et al., 2003). Recent genetic analyses confirmed that multiple independent introductions were responsible for the current distribution of C. similis in Florida (Nuñez, 2016).

Pathway Causes

Pathway Vectors

Impact Summary

Economic Impact

Economic impacts are localized and consist principally of damage to ornamental plants used as landscaping at private residences and community buildings. No estimate of direct economic loss is available. Since 2008, Florida residents in areas beset by populations of C. similis have self-imposed taxes totalling in excess of $250,000 to hire professional trappers to control these invasive animals.

C. similis has caused extensive damage to melon seedlings and chicks (Tasmitt and Valdivieso, 1963).

Environmental Impact

Impact on Biodiversity

C. similis on Gasparilla Island, Florida (USA) negatively impacts state threatened gopher tortoises (Gopherus polyphemus) through direct predation (Avery et al., 2009), occupation of tortoise burrows (Engeman et al., 2009), and utiliza­tion of food resources (Krysko et al., 2009). Other burrow- or ground-nesting species of concern, such as the Florida burrowing owl (Athene cunicularia floridana), a species of Special State Concern in Florida, and the federally threatened least tern (Sterna antillarum), are potentially vulnerable to predation, although direct impacts to these species remain to be documented (Krysko et al., 2003).

Threatened Species

Social Impact

None documented at this time.

Risk and Impact Factors

Impact mechanisms

• Competition - monopolizing resources
• Hybridization
• Predation

Impact outcomes

• Conflict
• Negatively impacts agriculture
• Threat to/ loss of endangered species
• Threat to/ loss of native species

Invasiveness

• Abundant in its native range
• Benefits from human association (i.e. it is a human commensal)
• Capable of securing and ingesting a wide range of food
• Gregarious
• Has a broad native range
• Has high genetic variability
• Has high reproductive potential
• Highly adaptable to different environments
• Highly mobile locally
• Is a habitat generalist
• Proved invasive outside its native range

Likelihood of entry/control

• Difficult/costly to control
• Highly likely to be transported internationally accidentally

Uses

This species is of limited value in the pet trade as it can be aggressive and “difficult” (Stephen et al., 2011). Total imports of C. similis to the US peaked in 2004 (3296 animals), but decreased to 76 by 2008 (Stephen et al., 2011). Illegal international trade involving this and other species of Ctenosaura on the internet is common (Stephen et al., 2011).

In many parts of its native range, C. similis is hunted and sold for food or for traditional medicinal uses. This species is not usually raised in captivity for meat production because, in contrast to Iguana iguana, successful captive rearing of C. similis is very difficult (Stephen et al., 2011).

Uses List

General

• Botanical garden/zoo
• Laboratory use

Human food and beverage

• Meat/fat/offal/blood/bone (whole, cut, fresh, frozen, canned, cured, processed or smoked)

Medicinal, pharmaceutical

• Traditional/folklore

Similarities to Other Species/Conditions

For years, most populations of C. similis in Florida, USA, were misidentified as Ctenosaura pectinata until field collections and close examination of museum specimens revealed the true identity (Townsend et al. 2003b). The two species can be distinguished by three morphological characters (Köhler et al., 2000; Townsend et al., 2003b):

1. C. similis has 0-12 (mean 0.4) scales separating the dorsal and caudal crests, while C. pectinata has 0-20 (mean 7.2).
2. C. similis has 2 complete rows of intercalaries between whorls of enlarged caudal scales near the base of the tail, compared to 3 rows in C. pectinata.
3. C. similis has dark dorsal crossbands which are absent in C. pectinata.

Prevention and Control

Prevention

Prevention is the best approach to invasive species management. Species such as C. similis which are very fecund and dietary generalists are especially problematic as just a few introduced animals can transform into a large, established population in short order.

Control

Physical/mechanical control

The C. similis population on Gasparilla Island, Florida (USA) grew without restraint for over 30 years before being subjected to control measures by professional trappers. Removal strategies included placing bait­ed live traps in shaded areas frequented by ctenosaurs, de­ploying unbaited live traps in the mouths of gopher tortoise burrows where ctenosaurs had entered, and placing unbaited live traps along the base of walls or other bar­riers where ctenosaurs travelled. Trappers employed com­mercial live traps with 2.54 × 2.54 cm or 1.27 × 2.54 cm mesh and baited with fresh cantaloupe or watermelon. Trappers dispatched captured lizards with a pistol load­ed with birdshot or .22 caliber bullets. Field personnel shot iguanas when prudent and safe during regular pa­trols of the study area using a .22 caliber rifle or air rifle (Avery et al., 2014). These removal efforts were highly successful as >20,000 ctenosaurs have been removed since 2008, and control efforts remain in effect.  

Chemical control

Possible methods for controlling C. similis using chemical toxicants were investigated. Captive ctenosaurs readily ingested sweet, red artificial baits into which capsules of toxicant were placed. Zinc phosphide was the most efficacious material tested, but application of a field baiting strategy employing toxic bait is dependent on development of target-specific delivery methods which are currently not available (Avery et al., 2011).

Monitoring and Surveillance

Increased biosecurity measures to detect and remove ctenosaurs potentially hitch-hiking on cargo ships would help prevent accidental introductions such as recently occurred in Venezuela (Barrio-Amorós and Rivas-Fuenmayor, 2008). Additionally, increased scrutiny of international trade in C. similis and similar species is needed to counteract illegal movements of these animals across borders.

Gaps in Knowledge/Research Needs

Rather than devote more resources, time, and personnel to studying the introduced populations there should be greater emphasis on removing them. This seems particularly true in Florida, USA, and Venezuela where populations are still manageable and eradication/extirpation is possible with dedicated, persistent management efforts.

References

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Contributors

08/06/2016 Original text by:

Michael Avery, USDA/Wildlife Services, USA

Distribution Maps

• = Present, no further details
• = Evidence of pathogen
• = Widespread
• = Last reported
• = Localised
• = Presence unconfirmed
• = Confined and subject to quarantine
• = See regional map for distribution within the country
• = Occasional or few reports

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