Anolis cristatellus (Puerto Rican crested anole)

Pictures

Picture	Title	Caption	Copyright
Title Adult Caption Anolis cristatellus (Puerto Rican crested anole); adult, basking. Bahia Honda Key, Florida, USA. March 2015. Copyright ©RobinSings/via wikipedia - CC BY-SA 4.0	Adult	Anolis cristatellus (Puerto Rican crested anole); adult, basking. Bahia Honda Key, Florida, USA. March 2015.	©RobinSings/via wikipedia - CC BY-SA 4.0
Title Adult Caption Anolis cristatellus (Puerto Rican crested anole); adult, basking. Picard, Dominica. February 2012. Copyright ©Postdlf/via wikipedia - CC BY-SA 3.0	Adult	Anolis cristatellus (Puerto Rican crested anole); adult, basking. Picard, Dominica. February 2012.	©Postdlf/via wikipedia - CC BY-SA 3.0
Title Adult Caption Anolis cristatellus (Puerto Rican crested anole); adult, basking. Picard, Dominica. March 2012. Copyright ©Postdlf/via wikipedia - CC BY-SA 3.0	Adult	Anolis cristatellus (Puerto Rican crested anole); adult, basking. Picard, Dominica. March 2012.	©Postdlf/via wikipedia - CC BY-SA 3.0
Title Adult Caption Anolis cristatellus (Puerto Rican crested anole); adult, basking. Picard, Dominica. February 2012. Copyright ©Postdlf/via wikipedia - CC BY-SA 3.0	Adult	Anolis cristatellus (Puerto Rican crested anole); adult, basking. Picard, Dominica. February 2012.	©Postdlf/via wikipedia - CC BY-SA 3.0
Title Head Caption Anolis cristatellus (Puerto Rican crested anole); adult, showing close view of head. Picard, Dominica. February 2012. Copyright ©Postdlf/via wikipedia - CC BY-SA 3.0	Head	Anolis cristatellus (Puerto Rican crested anole); adult, showing close view of head. Picard, Dominica. February 2012.	©Postdlf/via wikipedia - CC BY-SA 3.0
Title Adult Caption Anolis cristatellus (Puerto Rican crested anole); adult, displaying extended dewlap. Picard, Dominica. February 2012. Copyright ©Postdlf/via wikipedia - CC BY-SA 3.0	Adult	Anolis cristatellus (Puerto Rican crested anole); adult, displaying extended dewlap. Picard, Dominica. February 2012.	©Postdlf/via wikipedia - CC BY-SA 3.0

Identity

Preferred Scientific Name

• Anolis cristatellus Duméril & Bibron, 1837

Preferred Common Name

• Puerto Rican crested anole

Subspecies

• Anolis cristatellus cristatellus

Other Scientific Names

• Anolis cozumelae Smith & Taylor, 1950
• Anolis cozumelae Smith, 1939
• Anolis cristatellus Boulenger, 1885
• Anolis lindeni Ruthven, 1912
• Ctenonotus cristatellus Guyer & Savage, 1986
• Ctenonotus cristatellus Liner, 1994
• Ctenonotus cristatellus Schwartz & Henderson, 1988
• Xiphosurus cristatellus O’Shaughnessy, 1875

International Common Names

• English: crested anole

Summary of Invasiveness

A. cristatellus is a medium-sized brownish anole lizard characterized by a crest on the tail and body of adult males. Native populations occupy a wide range of habitats on Puerto Rico and the British and US Virgin Islands, from open habitats to rainforest, excluding only high altitude forests. Introduced populations occur in the Dominican Republic (since the 1910s), Florida (USA, since the 1970s), Costa Rica (1970s), Dominica (2000) and St. Maarten in the Netherlands Antilles (2008). All these introduced populations are spreading, although restricted to urban and other disturbed areas. This species may displace native Anolis species, but no other damaging effects on species or ecosystems have yet been recorded.

Taxonomic Tree

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

Notes on Taxonomy and Nomenclature

The taxonomy of the A. cristatellus has been uncontroversial in the 180 years since it was first described by Dumeril and Bibron, 1837. Only one change has been proposed in recent years, the general revision of the genus Anolis by Guyer and Savage (1986) reallocating the species to Ctenonotus cristatellus, but this usage disappeared after about 10 years and the species is currently known as Anolis cristatellus. The higher taxonomy is more controversial, Anolis being variously referred to the family Dactyloidae (The Reptile Database, 2017), Polychridae or Iguanidae.

There are two subspecies, A. cristatellus cristatellus and A. cristatellus wileyae. A. cristatellus cristatellus is widespread on Puerto Rico and its satellite islets whereas A. cristatellus wileyae occurs on the US Virgin Islands, the British Virgin Islands and other islands east of Puerto Rico (Isla Vieques, Isla Culebra, Isla Culebrita). All introduced populations are apparently the subspecies A. cristatellus cristatellus (Henderson and Powell, 2009).

Description

A. cristatellus is a moderately sized anole, snout-vent length (SVL) up to 75 mm in males and 73 mm in females, although most females are much smaller (Powell et al., 2015). Diagnostic scalation details are given by Schwartz and Henderson (1991). Colour is variable, with bronze, greenish grey head and body with irregular brownish spots and greenish yellow belly. Juveniles often have a light mid-dorsal stripe, retained in some females. The throat is whitish and the dewlap greenish centrally, reddish orange at the edges, or completely yellow-tan or deep orange. The main distinguishing feature is a compressed tail with a high fin in males, extending at lower height on to the body and increasing again on the nape of the neck. The iris is dark brown (Powell et al., 2015).

Distribution

The two subspecies of A. cristatellus have differing distributions; A. cristatellus cristatellus is widespread and native to Puerto Rico and its satellite islets, in a wide variety of habitats whereas A. cristatellus wileyae is native to the US Virgin Islands, the British Virgin Islands and other islands east of Puerto Rico (Isla Vieques, Isla Culebra, Isla Culebrita) (Henderson and Powell, 2009). Introduced populations of A. cristatellus cristatellus occur in the Dominican Republic, Florida, USA, Costa Rica, Dominica and St. Maarten in the Netherlands Antilles.

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

All introduced populations of A. cristatellus are the subspecies A. cristatellus cristatellus (Henderson and Powell, 2009).

The first introduction of A. cristatellus took place in the Dominican Republic and the port of La Romana on the south coast, probably before 1920 (Williams, 1977; Fitch et al., 1989). By 1989, it had become well established and abundant over a coastal area about 13 km in length and 12 km² in area, although confined to the town and adjacent disturbed areas. A. cristatellus coexists there with two other species of Anolis (A. distichus and A. chlorocyanus), but has an exclusive distribution with a third species (A. cybotes), which it may have displaced. Powell et al. (2011) record A. cristatellus as still localised in the Dominican Republic and Powell and Inchaustegui (2011) note that it is restricted to the La Romana urban area. Zani et al. (1993) and Kolbe et al. (2007) have determined the source areas in Puerto Rico for the introduction in the Dominican Republic. Distribution maps of A. cristatellus in the Dominican Republic can be found in Fitch et al. (1989).

A. cristatellus was first recorded by Schwartz and Thomas (1975) in Florida, USA in the early 1970s into two areas; on the island of Key Biscayne and on the mainland south of Miami. On the mainland, separate populations were found to exist at two localities in Broward and Dade Counties and there were other early populations around Miami which apparently died out (Bartlett and Bartlett, 1999). The introduced populations were tenuous in their first decade, but since expanded and today, A. cristatellus is now common around human settlements (Lever, 2003). Kolbe et al. (2007) have determined the source areas in Puerto Rico for the introduction in to Florida and it has been suggested that A. cristatellus entered Florida through the pet trade and noted it as a stage 5 introduction, being widespread and dominant in the area. The population on Key Biscayne was the third most frequently observed herpetofaunal species on the island in the period 2000-2006 (Krysko et al., 2010), after two other introduced reptiles (A. sagrei and Iguana iguana).

A. cristatellus was first recorded in Costa Rica by Fitch (1975) from his surveys in the period 1967-1974, at Parque Vargas and other places in the city of Puerto Limon, Limon Province. This has been the most successful introduction in terms of the area occupied and its spread beyond coastal areas. Savage (2002) shows A. cristatellus at five separate locations in Puerto Limon, Cahuita, Guayacan and Valle de Rosas (Limon Province) and Turrialba (Cartago Province), spread over an area of about 50 x 100 km. Savage (2002) provides maps of the distribution in Costa Rica.

Powell and Henderson (2003) identified A. cristatellus in Dominica based on an unpublished ms of RS Thorpe. Malhotra et al. (2007) describe the invasion initially from the coastal town of Roseau sometime between 1997 and 2002, to include about 10km of adjacent coastal regions. Powell et al. (2011) record A. cristatellus as still localized in Dominica, in disturbed habitats (Ackley et al., 2009) and maps of its distribution in Dominica can be found in Malhotra et al. (2007).

The latest introduction of A. cristatellus occurred in 2008 at the Port de Plaisance Resort and Casino, an islet in the Simpson Bay Lagoon, St. Maarten, in the Netherlands Antilles (Breuil et al., 2009). By 2012 it had spread about 500 m and was within 100 m of suitable habitat on the main island of St. Maarten. Yokoyama (2012) suggested that the invasion could still be eradicated at that stage, before the species crossed the bridge to the main island. Powell et al. (2011) record A. cristatellus as still localized in the Netherlands Antilles. Powell et al. (2015) suggested that this introduction had come in with ornamental plants from Florida, i.e. as a secondary introduction, not direct from the native Puerto Rico.

Introductions

Risk of Introduction

A. cristatellus is widespread in natural and disturbed habitats on its native Puerto Rico and most introductions into countries outside of its native range are thought to be a result of hitchhiking by adults or eggs in shipped cargos or in horticultural material. The introduction into Florida is likely to have been due the pet trade. There is therefore a high risk of future introductions through all three routes (cargo, horticulture, pet trade) and Bomford et al. (2009) include A. cristatellus (in 14^th place out of 596 species) on the list of the most successful invasive reptile and amphibian species, the highest ranking Anolis species.

Habitat

A. cristatellus occurs virtually island-wide in Puerto Rico, its native range, at altitudes up to 980 m, only excluding elfin woodland (Miranda-Castro et al., 2000). Henderson and Powell (2009) characterise A. cristatellus as mesophilic and xerophyllic, i.e. occupying moderate as well as dry habitats and adapting well to altered habitats. It is less abundant in high altitude rainforest, being replaced there by A. gundlachi (Henderson and Powell, 2009). The relative abundance of A. cristatellus in disturbed rural habitats increased during succession, from 0.04 in pasture, 0.08 in young forest, 0.60 in intermediate forest, to 0.57 in advanced secondary forest (Herrera-Montes and Brokaw, 2010). A. cristatellus was the dominant reptile species after pasture abandonment, suggesting that it is likely to be able to move out from urban areas into disturbed rural regions. In urban environments, A. cristatellus is able to use artificial, smooth surfaces effectively and are typically found on them 73% of the time. This is likely to induce selection for smaller body size and longer limbs in artificial habitats (Kolbe et al., 2016), as found by Winchell et al. (2016).

Habitat List

Biology and Ecology

Genetics

Natural selection within A. cristatellus has occurred in urban environments in Puerto Rico, with populations show 2% longer legs and 3% more toepad lamellae (plates used to grip surfaces when climbing). These differences were confirmed to be genetic after common-garden rearing experiments and reflect the use of smooth artificial surfaces (Winchell et al., 2016). In undisturbed habitats on Puerto Rico, A. cristatellus shows more genetic variation by ecology/habitat (39%) than by geography/distance (16%), than Anolis species in general (17 species, mean 18% by ecology and 36% by geography) (Wang et al., 2013). Thus individuals transported as hitchhikers from ports are likely to be already adapted to urban environments and possibly better able to compete with native anoles in such habitats. Revell et al. (2007, 2010) show that genetic variation between populations of A. cristatellus also affects phenotypic plasticity, allowing short-term adaptability as well as purely genetic responses.

Reproductive Biology

The clutch of A. Cristatellus is a single white egg, like all Anolis species (Andrews and Rand, 1974), size 10-11 x 6-7mm, laid under cover or in debris such as that around the base of banana plants (Fitch et al., 1989; Henderson and Powell, 2009). Eggs may be laid at intervals of two weeks during suitable conditions and females can store sperm for up to 84 days, producing up to six viable offspring over this period (Eales et al., 2008, 2010). Females may mate with multiple males and multiple paternities were found in 30% of progeny groups produced by females over time (Eales et al., 2010). Eggs may be laid communally by many females in the same location, with Rand (1967) noting 11 under one log. Reproduction is strongly seasonal in xeric (dry) habitats, restricted to the wet season (Licht and Gorman, 1975; Lister, 1981), but year-round in mesic habitats. Female reproductive cycles can differ over short (microlandscape) distances in A. cristatellus, between open (xerophilic) and closed (mesophilic) forest habitats (Otero et al., 2015).

Physiology and Phenology

A. cristatellus is active year-round, although non-reproductive in the dry season in xeric habitats (Licht and Gorman, 1975). The mean body temperature (T_b) in the native Puerto Rico is 29-30°C, with a critical thermal maximum of 38-39°C (Lister, 1976; Huey and Webster, 1976). The mean T_b in the introduced population in the Dominican Republic is 29.1°C (Fitch et al., 1989). A. cristatellus is a behavioral thermoregulator in open habitats, but a thermoconformer in closed forest (Huey, 1983). The optimum T_b for sprint performance is 27.3°C (Hertz et al., 1993). Low-temperature intolerance may cause the absence of A. cristatellus from upland rainforest in Puerto Rico (Gorman and Hillman, 1977). The Miami population of A. cristatellus has evolved a critical thermal minimum T_b some 3°C lower than that of the source population in Puerto Rico, within 35 years of the species invasion (Leal and Gunderson, 2012). Although tropical ectotherms are often thought not to be able to adapt to temperature changes over relatively short timescales, this study showed adaptation during the course of an invasion. However, a second introduced population, at Key Biscayne, showed no change in thermal acclimation ability (Kolbe et al., 2012).

The water loss rate of A. cristatellus is 0.08%/hour, lower than that of Anolis species from wetter, high elevation habitats (Hertz et al., 1979). Lizards from mesic and xeric sites have been shown to differ in water loss rates, with those from xeric (dry) areas losing water more slowly (Perry et al., 1999; Gunderson et al., 2011). There was, however, no short-term acclimation of water loss rates to dry (30% relative humidity) or moist (60% RH) conditions (Gunderson et al., 2011).

Longevity

The life expectancy of a medium-sized anole is about 2 years (Losos, 2009).

Activity Patterns

A. cristatellus is naturally diurnal, but may be active around artificial lights at night in urban areas (Garber, 1978; Schwartz and Henderson, 1991; Perry et al., 2008). Males display preferentially during periods of low environmental noise (vegetation movement) to increase the detectability of their signals (Ord et al., 2011).

Population Size and Structure

The population density of A. cristatellus in native populations is about 1.0 male and 2.6 females per tree (Philobosian, 1975), or up to 4000-7200 per hectare (Lazell, 1991).

Nutrition

A. cristatellus is a sit-and-wait predator (Perry, 1999), consuming a wide variety of insects and other small invertebrates including snails and earthworms. They also occasionally consume smaller lizards, including juvenile Anolis and fruits, berries and flowers (Henderson and Powell, 2009).

Environmental Requirements

A. cristatellus is a trunk-ground anole, with mean perch height of 1-1.2 m and perch diameter of 10-13cm (Losos, 1990), spending only 0-7% of the time on the ground (Hertz, 1992). It is a non-basking thermoconformer in shaded lowland (mesophilic) habitats on Puerto Rico and is intolerant of high body temperatures. Global warming by 3°C would negatively impact the species, with body temperatures becoming outside the preferred range at the high end and a large reduction in predicted performance (Huey et al., 2009).

Climate

Natural Enemies

Notes on Natural Enemies

Wild A. cristatellus have a high frequency of tail injury, with 71% of individuals having lost part of the tail in the native Puerto Rico (Lovely et al., 2010) and 70-80% in the introduced population in the Dominican Republic (Fitch et al., 1989). This suggests that populations will be resistant to high predator pressure. Individual A. cristatellus change their behavior after disturbance (Huang et al., 2011) and so may reduce the risk of mortality from predators.

Known predators of A. cristatellus include the lizards Anolis cuvieri and Ameiva, the snakes Borikenophis, Epicrates and Magliophis, the introduced Indian mongoose (Herpestes javanicus) (Henderson and Powell, 2009) and the American Kestrel (Falco sparverius) (Powell and Henderson, 2008a), the pearly-eyed thrasher (Margarops fuscatus) (Henderson and Powell, 2009), the Puerto Rican Tody (Todus mexicanus) (Rios-Lopez et al., 2015) and the red-tailed Hawk (Buteo jamaicensis) (Powell and Henderson, 2008a).

A number of parasites have also been recorded from A. cristatellus and include Lueheia inscripta (Henderson and Powell, 2009), Mesocoelium monas (Goldberg et al., 1998), Oochoristica maccoyi (Goldberg et al., 1998), Parapharyngodon cubensis (Goldberg et al., 1998; Falk and Perkins, 2013) and Plasmodium azurophilum (Schall and Vogt, 1993).

Means of Movement and Dispersal

Natural Dispersal

A. cristatellus are sedentary with small home ranges, only moving 0.8% of the time, with a mean of 0.36 moves/minute (Perry, 1999). They do show homing ability if displaced up to 60 m outside the home range (Jenssen, 2002). Limited natural movements are also shown by their sleep site fidelity, with 98% of individuals located within 1 m on five consecutive nights (Clark and Gillingham, 1990). Nevertheless, the species rapidly invaded a high elevation site in Puerto Rico after the canopy was opened up following leaf loss after irradiation (Turner and Gist, 1970), showing some natural dispersal, probably (as in other Anolis species) from movements of juveniles.

Accidental Introduction

Perry et al. (2006) noted five juveniles of A. cristatellus in ornamental plants moved from Beef Island to Guana Island among the British Virgin Islands. Communal egg laying would increase the chances of a successful invasion through hitchhiking of eggs. Local physiological adaptation, documented for thermal biology, water loss and reproduction, suggests that hitchhikers moving between similar habitats (port to port or horticulture to gardens) would be suitably adapted, also increasing the chance of invasion success.

Intentional Introduction

Kraus (2008) noted that A. cristatellus may have been intentionally introduced to Florida, through the pet trade in exotic reptiles.

Pathway Causes

Pathway Vectors

Impact Summary

Environmental Impact

Impact on Biodiversity

A. cristatellus is only likely to directly impact other species of lizard, in particular species of Anolis. A. cristatellus is an aggressive species and interspecific aggression has been observed with other native species in Puerto Rico (Ortiz and Jenssen, 1982; Jenssen et al., 1984; Hess and Losos, 1991).

A. cristatellus has been found to compete with and displace A. cybotes in the Dominican Republic (Fitch et al., 1989) and A. oculatus in Dominica (Malhotra et al., 2007). A. cristatellus has also been noted as a predator of A. pulchellus and A. stratulus (Rios-Lopez et al., 2015). This competition may take the form of intra-guild predation on juveniles or more rapid population growth (Fitch et al., 1989). A study by Salzburg (1984) found that A. cristatellus affected the perch height of another introduced species, A. sagrei, in Florida, but no other negative effects were noted.

Risk and Impact Factors

Impact mechanisms

• Competition - monopolizing resources

Impact outcomes

• Conflict
• Reduced native biodiversity

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
• Has high genetic variability
• Highly adaptable to different environments
• Highly mobile locally
• Is a habitat generalist
• Tolerant of shade

Likelihood of entry/control

• Highly likely to be transported internationally accidentally
• Highly likely to be transported internationally deliberately

Uses

Economic Value

A. cristatellus may be of some economic benefit in terms of pest control in plantations (Borkhataria et al., 2012), as the diet includes beetles, flies, lepidoptera, hemiptera and snails (Henderson and Powell, 2009).

Social Benefit

In urban areas A. cristatellus may be of some social benefit as the diet includes cockroaches and flies (Henderson and Powell, 2009). They are also becoming more popular as pets. A. cristatellus has been a model organism in studies of ecology, ecophysiology, evolution, behavior (Charles and Ord, 2012) and recently in the impacts of climate change (Gunderson and Leal, 2012, 2015, 2016).

Uses List

General

• Botanical garden/zoo
• Laboratory use
• Pet/aquarium trade
• Research model

Detection and Inspection

A. cristatellus is likely to be first noticed as an unusual crested anole unlike existing species in an area. Most areas suitable for A. cristatellus introduction will already have native or other introduced Anolis species. Schwartz and Henderson (1991) provide a useful guide to identification, but this should be confirmed by a taxonomic specialist as there are hundreds of Anolis species potentially involved in introductions.

Similarities to Other Species/Conditions

There are approximately 400 species of anoles (Nicholson et al., 2012) and although males of A. cristatellus are distinctive, final confirmation of identity requires capture and checking of scalation details (Schwartz and Henderson, 1991).

Gaps in Knowledge/Research Needs

A. cristatellus has been a model organism in studies of ecology, ecophysiology, evolution, behavior (Charles and Ord, 2012) and recently in the impacts of climate change (Gunderson and Leal, 2012, 2015, 2016). These provide an excellent base for the study of invasion and adaptation in a lizard.

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Principal Source

Draft datasheet under review

Contributors

30/03/2017 Original text by:

Adiran Hailey, The University of West Indies, Trinidad and Tobago

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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