Author: Pete Zani

Anoles of South America Part 3: Introduced Anoles

Anolis porcatus collected in Brazil, and comparison with the native anole A. punctatus. A, male A. porcatus showing green coloration. B, male A. porcatus showing brown coloration. C, the pink dewlap of male A. porcatus. D, female A. porcatus. E, male A. punctatus, a native anole species. F, the yellow dewlap of male A. punctatus. Picture credits: A–D, Mauro Teixeira Jr.; E, Renato Recoder. For more information, see this Anole Annals post or read on below.

In North America, especially in places like Florida, invasions of introduced species have revealed novel insights into the competition between species as well as the formation of non-analog species assemblages during such events. Anolis lizards, in particular, have demonstrated a strong ability to invade novel environments possibly even displacing native species in many cities in the United States. Yet the same appears not to be true in South America possibly because similar invasions are only just now occurring. In previous posts I considered diversity of cis– and trans-Andean South American anoles. In this post, I will review what can be inferred about the introduced species of anoles in South America using data accumulated on iNaturalist.org.

Certain introductions have been reported, but not yet observed on iNaturalist, such as Anolis sagrei in Rio de Janeiro (Oliveira et al. 2018). However, there are a handful of examples of anole introductions in South America that can be explored in terms of their progress as becoming invasive. As it turns out, most of the anole introductions appear to be fairly early along in the process of invasion making their documentation a useful exercise for future study by anole enthusiasts.

Despite the lowland Amazon rainforest’s vast size, introduced anoles have not been an issue there until very recently. Where introductions do appear to be occurring in South America, they seemed to have started in coastal cities. The first area of interest are the coastal cities in the northeast of South America.

Anolis aeneus in Guyana


Anolis aeneus has been known from Guyana for several decades and is possibly over a century old (review by Williams 1977, see also Schwartz and Henderson 1991). This species appears to be established in Georgetown, Guyana, and has been observed on iNaturalist since 2012. By the end of 2024 there were still only 34 observations of this species in all of Guyana, but this includes at least a pair of observations away from Georgetown in the interior of that country. So, while A. aeneus is considered introduced in Guyana (see image below), it does not appear to be far along in the process of invasion at this point or moving especially rapidly.


Figure above shows all iNaturalist observations of introduced anoles in Guyana, Suriname, and French Guiana.

 

Anolis marmoratus and A. sagrei in Suriname and French Guiana

A larger ongoing invasion appears to involve the introduced Guadeloupean Anole (Anolis marmoratus) in French Guiana (see image above). Anolis marmoratus was first reported in French Guiana in 1975 as already well established in Cayenne, French Guiana (Hoogmoed and Lescure 1975). On iNaturalist, this species has been reported in the vicinity of Cayenne since at least 2004 but with only a handful of observations each year until very recently.

Annual observations of A. marmoratus from 2021-2023 were 7, 6, and 8, respectively, including one or two additional observations in Kouro about 40 km along the coast to the northwest. However, in 2024 observations jumped to 18 and came from every neighborhood of the twin cities of Cayenne and Remire-Montjoly as well as another seven observations in Roura 15 km to the south and eight in Kouro. While A. marmoratus appears to be established and spreading in French Guiana, it might begin experiencing competition from another well-known invasive anole that appears to be recently introduced in the area: Anolis sagrei.

The first report of Anolis sagrei in northeast South America came from Paramaribo, Suriname, in 2022 when a single observation was added to iNaturalist from within that city. This species was not among those listed as introduced to Suriname by Hoogmoed (1980), although other species of anoles were mentioned, but not yet observed on iNaturalist in Suriname.

In 2024, three observations of A. sagrei were made in the same area of French Guiana. In both cases, it is too early to tell if these are established populations and if they will spread, but the known invasive habits of A. sagrei in places like south Florida make this seem a likely occurrence.

Anolis sagrei in Guayaquil, Ecuador


In western South America, there have been occasional single observations of Anolis sagrei in places like Zamona, Ecuador, along the Napo River in the Amazon occurred in 2022. However, this observation has not been corroborated so far. Until additional observations are made, it is not possible to determine if we are seeing the beginnings of invasions in any of these places. Yet, A. sagrei already has a well-established foothold in another South American location that is worth examining: Guayaquil, Ecuador, reviewed recently by Narváez et al. (2024)(see image below).


Figure above shows all iNaturalist observations of Brown Anoles in the vicinity of Guayaquil, Ecuador. Image courtesy Jaime Camacho via iNaturalist.


Anolis sagrei was first observed on iNaturalist in 2007 in a park in Guayaquil, in southwestern Ecuador, but first reported in about the year 2000. This observation was of a lizard on the peninsula formed by the confluence of the Daule, Babahayo, and Guayas Rivers. It was not observed again until 2016 in that same park, but soon it became apparent it was probably there all along, which is why continued monitoring is essential for understanding invasions.

In 2017 Anolis sagrei was observed six additional times, but had now expanded to occur in several different places in the city having moved about 4 km from the original location and even jumped the Daule River. That same year also saw a single observation of A. sagrei in the northwestern Ecuador city of San Lorenzo, but this has never been repeated so it is hard to tell if this the beginnings of a new invasion or not. By 2018 it was at least 6 km from the original site and in 2019 A. sagrei was observed 17 times having reached the forested outskirts of Guayaquil to the west some 10 km from original location. Thus, even as late as 2019, A. sagrei appears to have been in the early stages of the invasion, but with a big leap forward in 2020.

In 2020, A. sagrei appears to have begun moving along the roads to the north and west of Guayaquil and tripled its occurrence to 49 observations in and around the city. Half of those observations in 2020 occurred on the peninsula where the species was first noted in iNaturalist. But 2020 also saw a leap for this species with an observation for the first time in Portoviejo nearly 200 km to the northwest of Guayaquil.

The next year, 2021 saw another 50 observations, with over half occurring outside the area where they were first noted. In that year, the suburbs of Guayaquil to the north and west of saw strong growth in observations. Similarly, 2022 added another 48 observations in the vicinity of Guayaquil. In 2022, the population had reached a zoo near Victoria 15-20 km to the north of Guayaquil. Thus, by the early 2020’s A. sagrei had become firmly established in the greater Guayaquil areas and this species took another leap forward in 2023.

In 2023, 85 observations spanned nearly every neighborhood of Guayaquil and its outskirts. Interestingly, even by 2023 A. sagrei had still not jumped the Guayas River to Duran on the east bank. Nor had it colonized the forested reserve of Isla Santay in the Guayas River between these two cities. However, in 2023 A. sagrei did make a big leap northwest to the coastal city of Manta about 20 km west of the aforementioned Portoviejo.

Guayaquil continued to be the center of observations for A. sagrei in 2024 with another 52 observations, but now observations are being reported elsewhere along the coast including Olon, which is about 100 km west of Guayaquil. However, the observations near Manta, Portoviejo, and San Lorenzo have not been repeated so far. Nor has the species appeared to jump the Guayas River to either Isla Santay or Duran.

Isla Santay is of particular interest because of the known population of the native Anolis festae that occurs there. If A. sagrei were to arrive on Santay it could lead to the extirpation of A. festae at that site or at least a novel competitive environment for that native anole. This is where directing a community of observers, such as on iNaturalist, might be of service to monitor these areas for the novel spread of A. sagrei as well as the persistence of A. festae.

If we take 2007 as the initial year of introduction, A. sagrei appears to be expanding its range at about 5 km per year, but with some leaps out beyond that probably via human-aided dispersal. And even though community data such as provided by iNaturalist are not controlled for in terms of effort, they do provide a fascinating window into an ongoing invasion by a known invader. However, A. sagrei isn’t the only invader of South America.


Figure above shows iNaturalist observations of Brown Anoles in Guayaquil, Ecuador binned in three-year intervals.

 

Anolis porcatus in Santos, Brazil

A lesser-known invasion of South America appears to involve Anolis porcatus in southeastern Brazil. This invasion appears to be even more recent than A. sagrei in Guayaquil and is worth detailing. Anolis porcatus was first observed on iNaturalist in Santos, Brazil (a coastal city just south of São Paulo) in March 2016 at about the same time it was reported in the literature (Prates et al. 2016, Samelo and Barella 2016). This was the only observation of this introduced species until 2018 when five additional observations recorded this species in three separate locations in São Vincente 8-10 km to the west of the original location.


Figure above shows all iNaturalist observations of Cuban Green Anoles in the vicinity of Santos, Brazil. Image courtesy Nereston Camargo via iNaturalist.org.


In 2019, an additional nine observations of A. porcatus occurred in Santos and São Vincente. Thus, by the end of 2019 it appears A. porcatus had established in four different locations. However, observations were still only occasional at this time.

In 2020 A. porcatus jumped to Cubatão, a city another 5 km to the north of São Vincente, and in 2021 it jumped to Praia Grande 5-6 km along the coast to the southwest of São Vincente. By 2023 there were another 21 observations of this species in and around the area including the forested outskirts to the north of Cubatão. In 2024 another 18 observations of the species suggested A. porcatus was now firmly established around Santos.

In addition to the above observations, in October 2024 A. porcatus was noted for the first time southwest along the coast near Itanhaém about 45 km from the original source location in Santos. Thus, similar to A. sagrei in Ecuador, A. porcatus appears to be making leaps from city to city, possibly via human-aided dispersal.

If we take 2016 as the initial year of introduction, A. porcatus appears to be expanding its range at about 5 km per year and should reach the outskirts of São Paulo in another year or two if this rate holds. This introduction does not yet appear to be affecting any of the native anoles since the closest natural populations of any Anolis occur in Rio de Janeiro several hundred km to the northeast.


Figure above shows iNaturalist observations of Cuban Green Anoles in Santos, Brazil binned in three-year intervals.


When A. porcatus does reach Rio, it could begin competing with the native Amazon Green Anole (A. punctatus), which looks and behaves very similarly. The protected forested corridor along the coast means that there is no meaningful barrier to movement from São Paulo to Rio for the introduced A. porcatus at the moment. Likewise, it seems only a matter of time before A. porcatus moves southwest along the coast toward Curitiba some 200 km away.

Summary


To summarize, there have not been many instances of introduced anoles in South America for whatever reasons. Nearly all of these start near the coast, likely because invaders are hitching rides in shipping containers or other ocean transport. Right now, the most advanced invasion seems to be taking place in Guayaquil, Ecuador with more recent invasions likely taking place in French Guiana and near São Paulo. Each of these introductions appears to be in the early days of invasion with establishment only clear in and around those locations where originally observed. The expansion of introduced species seems to involve expanding within local areas of these cities and only then expanding by leaps and bounds to nearby cities after several years of establishment. With the rise of community-based observations on platforms such as iNaturalist we can make predictions about when these introduced species will arrive in nearby cities. Likewise, we can leverage the local populace for use in environmental monitoring to better understand potential invasive dynamics of these introduced species for which anoles serve as a useful model system. Whether highly invasive anoles such as A. sagrei can ever invade species-rich environments such as the Amazon or other non-coastal areas remains to be seen. Yet platforms such as iNaturalist may allow us to observe such events in real time if the community of observers can be cultivated and maintained.

Literature cited


Hoogmoed, M.S. 1980. Introduced species of reptiles in Surinam. Notes on the herpetofauna of Surinam VIII. Amphibia-Reptilia 1:277-285.

Hoogmoed, M.S., J. Lescure. 1975. An annotated checklist of the lizards of French Guiana, mainly based on two recent collections. Zoologische Mededelingen 49:141-171.

Narváez, A.E, F. Ayala-Varela, S. Cuadrado, K. Cruz-García, M.H. Yánez-Muñoz, L. Amador. 2024. Updated distribution of the brown anole in continental Ecuador: a case of urban spaces favoring the establishment of an opportunistic exotic species. BioInvasions Records 13:373-384.

Oliveira, J.C.F., T.M. Castro, M.C. Drago, D. Vrcibradic. I. Prates. 2018. A second Caribbean anole lizard species introduced to Brazil. Herpetology Notes 11:761-764.

Prates, I, L. Hernandez, R.R. Samelo, A.C. Carnaval. 2016. Molecular identification and geographic origin of an exotic anole lizard introduced to Brazil, with remarks on its natural history. South American Journal of Herpetology 11: 220-227.

Samelo, R.R., W. Barrella. 2016. Geographic distribution: Anolis porcatus (Cuban Green Anole). Herpetological Review 47:256.

Schwartz, A., R.W. Henderson. 1991. Amphibians and Reptiles of the West Indies: Descriptions, Distributions, and Natural History. Univ. Florida Press, Gainesville.

Williams, E.E. 1977. Anoles out of place. The Third Anolis Newsletter. pp. 110-118.

Anoles of South America Part 2: Diversity South of the Andes

Anolis phyllorhinus. [Photo from iNaturalist by Christopher Borges](https://www.inaturalist.org/observations/145912245)

In a previous post, I reviewed the diversity of Anolis lizards north of the Andes in South America. In this post I continue on that theme with a review of the diversity of anoles in the Amazon and south of the Amazon throughout South America. Much of this review must include the caveat that this understanding is ever-changing as new taxa are described.

I start with a list of native anoles south of the Andes in South America, which is a surprisingly short list considering the large area included and that this list also includes the Amazon basin.

Widespread throughout both Amazon and Atlantic rainforests
A. punctatus, A. fuscoauratus, A. ortonii

Widespread, but limited to the Amazon
A. trachyderma, A. transversalis, A. chrysolepis, A. tandai, A. bombiceps

Occurs in Amazon and/or llanos northwest of Amazon
A. scypheus, A. inderenae, A. vanzolinii, A. huilae, A. ruizii

Limited distribution in northern llanos/Amazon and north of Andes
A. planiceps, A. auratus

Range restricted in western and southwestern Amazon
A. phyllorhinus, A. cuscoensis, A. dissimilis, A. lososi, A. boettgeri, A. soinii, A. fitchi, A. podocarpus, A. orcesi

Widespread in sub-Amazonian (southeastern) dry forests of South America
A. meridionalis, A. brasiliensis

Limited distribution in sub-Amazonian (southeastern) Atlantic rainforests of South America
A. nasofrontalis, A. pseudotigrinus. A. neglectus

There are roughly 102 anole species that occur in all of South America. However, 67 or so of these occur north and west of the Andes (the so-called trans-Andean region) leaving perhaps 35-40 species south of the Andes (cis-Andean). The numbers don’t quite add up because a small handful of species (e.g., A. planiceps, A. auratus) occur in both regions depending on how these regions are defined geographically. So, despite the great diversity of anoles in the Caribbean and Central America, or even the respectable diversity of anoles north of the Andes, the cis-Andean region is relatively diversity poor. Causes for this lack of cis-Andean anole diversity includes the lack of meaningful geographic barriers that may make speciation less common in anoles. Alternatively, these animals are less commonly used as study subjects so subtle molecular diversity may not yet have been identified in these lineages. Either way, despite occurring throughout the Amazon basin, the llanos of Colombia and Venezuela to the northwest of the Amazon, the dry transitional forests and grasslands of Brazil to the southeast of the Amazon, as well as in the coastal rainforests of Brazil along the Atlantic coast, only several dozens of species of anoles occurn south of the Andes mountains, many of which are range restricted in a few small areas up against the mountains themselves.

Most of the anoles in this area of South America are found in the Amazon region, which makes sense given the vast extent of the rainforest ecosystem in the northern half of the continent. Despite this, there are currently only 28 or so species of anoles in the entire Amazon observed on iNaturalist.org, all but three of which are considered native (non-introduced) species. Over half of these species (16 or so) are endemic to the Amazon region (broadly defined) and found nowhere else [note, in this context, I use iNaturalist’s definition of “Amazon region” which includes the dry edges [llanos in the northwest, cerrado in the southeast] and not just the rainforest ecosystem proper, including extending all the way to the Orinoco in the north). So, clearly the Amazon is the center of diversity for cis-Andean anoles even as the number of species (25 or so native species) is not especially impressive given that there are currently over 300 species of lizards described from the Amazon. Perhaps it is this lizard diversity itself that accounts for the lack of species richness in anoles in that many of the niches occupied by anoles elsewhere (e.g., crown giants) are likely occupied by competing species (e.g. Tropiduridae, Polychrotidae). Or if not competitors, perhaps the great diversity of predators is what accounts for the lack of anole diversity (iNaturalist currently includes observations of over 300 species of snakes in the Amazon alone). Either way, anoles are clearly not the dominant taxon south of the Andes that they appear to be elsewhere. However, they do still occur nearly everywhere in South America.

The natural distribution of anoles in South America reaches south to about the latitude of São Paulo (23.6°S), or just outside the Tropic of Capricorn. This corresponds roughly to the latitude of Miami just outside the Tropic of Cancer in the northern hemisphere. However, the native Anolis carolinensis in the north has a natural range at higher latitudes, suggesting the southern extent of the anole radiation is not as great as it is in the north. As with that northern range margin for the clade, the diversity of anoles dwindles greatly along the Atlantic coast near the range edges with only perhaps one native species occurring south of Rio de Janeiro (22.9°S) (Anolis porcatus is introduced in São Paulo, but introduced species will be discussed in a future post). Even inland, there is only perhaps one species of anole that ranges as far south as Paraguay or far northern Argentina. So, even though widespread, the pattern of diversity is such that anoles appear less widespread in southern hemisphere than in the northern hemisphere.

Perhaps, one of the more interesting patterns to emerge when considering anole diversity in South America is the slow, but steady description of new species, many of which are range limited to small areas in the southern and western parts of the Amazon. The greatest number of species in the Amazon can be described as range restricted with perhaps nine species occurring in small local enclaves or areas. These include some of the more enigmatic species such as Anolis phyllorhinus with the long fleshy proboscis in the males. Thus, as with the trans-Andean anoles, the geographic complexity of the Andes has likely created conditions for some version of allopatric species to take place, whether it be true allopatry or some type of peripatry on a range’s periphery. I fully expect that additional species will be described from the Andean foothills or the western and southwestern Amazon.

Finally, it is interesting to consider the near complete replacement of species on either side of the Andes with very little crossover; clearly the Andes are a fairly effective barrier to gene flow for anoles. A few species appear to occur on both side of the Andes in the area south and east of Bogotá, Colombia. However, this crossover appears to be limited in terms of diversity and extent of species. Instead, the biggest exchange of species appears to have occurred around the northern terminus of the Andes in Venezuela where two species in particular span the cis- and trans-Andean regions: Anolis planiceps and A. auratus. Anolis planiceps occurs along the Caribbean coast of Venezuela as well as south into Colombia, Guyana, and Roraima, Brazil. However, Anolis auratus as currently defined occurs much more broadly ranging from Costa Rica and Panama, throughout cis- and trans-Andean regions of Colombia and Venezuela, along the Atlantic coast to at least Amapá, Brazil, and south into the states of Roraima and Pará, Brazil. In Brazil in particular A. auratus seems to be limited to areas of geologic uplift that have created the grassland habitat with which this species is associated. The highlands to the north and south of the Amazon river were created when the Guiana Shield and Brazilian Shield, respectively, were forced upward by geologic uplifting. Hence, the Amazon river roughly runs in the low areas in-between the uplift of these two large blocks on the South American continent. Just south of the Amazon River, an isolated population of Anolis auratus occurs in a place called Alter do Chão (translation from Portuguese is literally ‘altar of ground’) where a small piece of the raised uplift creates savanna-like grassland/forest habitat surrounded by lowland Amazon rainforest to the south and east and bordered by the Amazon and Tapajós Rivers to the north and west, respectively. The presence of these lizards in as a small and isolated enclave 1000 km from their nearest neighbors generates more questions than answers. How long have they been there? How did they get there? How distinct are they? How stable is this population? These are all good and unresolved questions remaining to be addressed biologically.

I saved the above observation for last to make the point that on the surface the Amazon may appear to be a vast tract of uniform habitat with little barriers to gene flow leading to biological uniformity, but the reality is that this view is overly simplistic in the context of how species are actually distributed on the landscape. Rather, it is likely the lack of available data that prevent us from truly understanding the biogeography of lizards in the Amazon and other areas of South America. This makes the continual addition of new species and biogeographic patterns useful and exciting given the advances in molecular biology that continue to further our understanding. Hence, even in the relatively anole-poor regions of South America, anoles continue as a model system for understanding patterns of biological diversity.

Anoles of South America Part 1: Diversity North of the Andes

Although anoles are much more well studied and understood in the Caribbean and United States, there is a decent amount of diversity of these lizards throughout South America as well. In consultation with Jonathan Losos, I have agreed to write a series of posts outlining the South American diversity (parts 1 and 2), invasives (part 3), and conservation (part 4) of these animals. This post (diversity part 1) will describe the species that occur in the regions of South America north and west of the Andes along the Caribbean and Pacific Coasts, respectively, while diversity part 2 will focus on the Amazon and everything south and east of the Amazon.

In case you don’t know me, I had experience as a graduate student studying lizard functional morphology of toes and tails in North, Central, and South America. More recently, I have become way too involved in identifying lizards on iNaturalist.org, with a focus on lizards of the New World. To satisfy my own particular obsessiveness, I have tried to identify every lizard observed in the Amazon (all 300+ species of them!) and have expanded those lizard-identifying efforts into other areas as well. It was these efforts that gave me the idea that perhaps the AnoleAnnals might be interested in what I’ve noticed from this relatively understudied part of the world. As such, I will draw liberally from data available on iNaturalist to illustrate my points.

Anole diversity along Caribbean and Pacific Coasts

As of this writing, there are currently nearly 12,000 observations on iNaturalist of 102 species Anolis in South America (“research grade” observations filtered by “wild” and “has photos”). Compare this to the 238,000 observations of 270 species from North America (on iNat this region includes all of Central America, Mexico, the United States, and the Caribbean). Only eight of the species of South American anoles are introduced non-native species (more on this in part 3). This suggests that there are nearly 100 species of Anolis endemic to South America. Clearly, the anoles from South America are not as diverse and not as well observed as those from other areas.

Anoles along the Pacific Coast range from southern Ecuador (and perhaps northern Peru, but not well documented) north to the junction with Central America and continue along the Caribbean Coast east to the mouth of the Orinoco River in Venezuela (note: according to iNaturalist, Guyana, Suriname, and French Guiana are part of the Amazon region even though they may not be part of the Amazon watershed). This region is separated from the Amazon by the Andes Mountains and Orinoco River and very few species cross these particular barriers. Hence, many of the anoles of northern South America still have some degree of endemism and it is this endemism (and the resulting beta diversity) that I highlight below. Keep in mind that new species are being described on a regular basis, so the list below is almost certainly incomplete, but still illustrates my general point.

The species of Anolis that occur in north and west of the Andes include (organized by their geographic distribution):

Western (non-Amazonian) Ecuador (n=9)

A. bionotatus, A. bitectus, A. fasciatus, A. festae, A. lyra, A. neomonteae, A. parilis, A. peraccae, A. proboscis

Western (non-Amazonian) Colombia (n=18)

A. anchicayae, A. antioquiae, A. antonii, A. calimae, A. danieli, A. eulaemus, A. gorgonae, A. heterodermus, A. maculigula, A. macrolepis, A. mariarum, A. medemi, A. quimbaya, A. solitarius, A. sulcifrons, A. tolimensis, A. urraoi, A. ventrimaculatus

Western Ecuador/Colombia (n=13)

A. aequatorialis, A. anchicayae, A. chloris, A. dracula, A. fraseri, A. gemmosus, A. gracilipes, A. granuliceps, A. limon, A. lynchi, A. maculiventris, A. parvauritus, A. princeps

Northern Colombia/Venezuela (n=3)

A. jacare, A. onca, A. tetarii

Northern Venezuela (n=4)

A. anatoloros, A. annectans, A. squamulatus, A. tigrinus

Aruba, Bonaire, and/or Curaçao (A-B-C islands) (n=2)

A. bonairensis, A. lineatus

Found in western Colombia as well as crossing the Darién Gap into Central America (n=8)

A. biporcatus, A. gaigei, A. latifrons, A. poecilopus, A. purpurescens, A. triumphalis, A. tropidogaster, A. vittigerus

Along Caribbean Coast as well as crossing into the Amazon (n=2)

A. auratus, A. planiceps

Non-native anoles (n=8)

A. aeneus, A. cristatellus, A. marmoratus, A. porcatus, A. richardii, A. sagrei, A. trinitatis, A. wattsii

Several things become apparent when considering the above diversity.

—First, there are roughly 67 species of anoles that occur north of the Amazon. Given that there are only about 102 anole species that occur in all of South America, perhaps 2/3 occur in the area adjacent to the speciose Caribbean. That leaves only 1/3 of the South American anole species occurring south and east of the Andes Mountains.

—Second, of the 67 anole species found north of the Amazon, 47 species (70%) are endemic to this region of mainland South America, two species (3%) are endemic to the A-B-C islands, eight species (12%) cross from mainland South America into Central America, and two species (3%) are found extensively in the Amazon as well. So, not as diverse as North America, but with outstanding regional endemism. This regional endemism translates into an impressive beta diversity with geographic replacement probably the rule rather than the exception. In other words, when travelling from one valley to the next, species replacement (beta diversity) occurs at a relatively high rate so that nearly an entirely different set of species may be present in adjacent areas, depending on the nature of the barrier between them. Making this even more of a challenge, in some instances the barrier appears to be the low-lying valley instead of a mountain ridge. For example, in Colombia, Anolis heterodermus Duméril 1851 was recently split into eight distinct species (Morena-Arias et al. 2023). Both the various mountain ranges (Cordillera) as well as the Magdalena River valley appear to act as barriers isolating the various lineages into distinct species. Hence, in terms of trying to identify the Anolis in some of these areas, accurate geographic location is a must.

—Third, there are only eight species (12%) of non-native anoles in all of South America (including the A-B-C islands and Trinidad and Tobago). All eight occur in northern South America and only one or two (A. porcatus and perhaps A. sagrei) occur elsewhere in South America. Contrast this with the 16 or so species of introduced anoles in North America, which have made extensive inroads in places like the southeastern United States, southern Mexico, and certain of the Caribbean Islands. These eight introduced species will be the focus on part 3 of this series.

One of the interesting things about the high degree of endemism of the anoles of northern South America is how quickly it probably occurred. Andean orogeny began perhaps 20 million years ago (mya) and lasted until about 8 mya. Hence, the diversity differences between the occidental (western) and oriental (eastern) regions of countries such as Ecuador, Colombia, and Venezuela are notable for their near completeness. For comparison, the Laramide orogeny that created the Rocky Mountains occurred between 80-55 mya; the Alleghanian orogeny that created the Appalachian Mountains occurred between 325-260 mya. Hence, the temporal arrival of the Andes created spatial barriers to gene flow and geographic complexity that is reflected in the high beta diversity of Anolis both within and between regions of South America.

In conclusion, northern South America is the diversity hotspot for anoles. The anole species north of the Amazon are dominated by native, highly endemic species that often have geographic ranges limited to a single country, a single valley, or even one slope of a single mountain ridge. Moreover, there are relatively few introduced anoles in South America with most of those introduced to the A-B-C islands and Trinidad and Tobago. Given that almost no species of anoles cross the Orinoco River or Andes Mountains and very few species occur extensively in the Carribean or Central America, the anoles of northern South America represent a fairly separate and distinct fauna when compared to other geographic regions within the distribution of Anolis.

 

Literature cited:

Moreno-Arias, R. A., Méndez-Galeano, M. A., Beltrán, I., & Vargas-Ramírez, M. 2023. Revealing anole diversity in the highlands of the Northern Andes: New and resurrected species of the Anolis heterodermus species group. Vertebrate Zoology, 73, 161-188

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