Author: jjkolbe

Anole Invaders

The current distribution of plants and animals around the Earth has been strongly influenced by colonization, the ability of organisms to disperse short or long distances on their way to establishing new populations. Over the past several hundred years, humans have increased the rate and distance over which organisms colonize new lands. These human-mediated introductions have reshaped basic patterns of biogeography, a field that investigates the geographical distribution of plants and animals. Some of these invaders cause ecological or economic harm in their new homes and are known as invasive species. In cases where we lack detailed study of an introduced species, we often use high local abundance and rapid spread as proxies for invasiveness. Anoles are one of the most prolific groups of invaders with over 20 different species introduced outside of their native ranges. Most anole invaders originate from Caribbean islands and introductions occur to other islands in the Caribbean, Atlantic and Pacific as well as South Florida, Central America and Southeast Asia. Researchers are studying a diverse array of topics related to anole invasions including the origin of invaders in their native ranges, patterns of introduction and spread, and impacts on native species.

The brown or festive anole (Anolis sagrei) is native to Cuba and the Bahamas, but has been introduced widely around the Caribbean, Central America, and the continental U.S., where it has spread north from the Florida Keys into several other states. Photo by Day’s Edge Productions.

In my lab, we use DNA sequences and other types of genetic data to identify the origin of introduced anoles. Where did invaders originate from in their native range? How many introductions have occurred? How have they spread in their introduced range? You can think of each anole that invades a new area as having their native-range origin encoded in their DNA as if they were carrying a passport from their home region. Using this approach on over a dozen anole species, we in most cases identified the geographic source of introductions, commonly hubs of transport and commerce, which is consistent with anoles being transported by shipping. Many of these anole introductions occurred among Caribbean islands or from Caribbean islands to South Florida. We also found that anole invasions often originate from multiple, geographically and genetically distinct populations in their native range and when thrust together in their introduced range they mix together. This has important implications for evolution, including the potential to enhance adaptation by increasing genetic variation. Our DNA studies also reveal that well-established populations in introduced range can become sources for secondary introductions, which has been termed the bridgehead effect. Brown anoles (Anolis sagrei) are a particularly good example of this phenomenon as introduced populations in South Florida around Miami are likely the source of introductions to Bermuda, Grand Cayman, Grenada, and Hawaii. Similarly, the spread of green anoles (Anolis carolinensis) in the Pacific resulted from a stepping-stone pattern among islands after introduction to the region.

The green anole (Anolis carolinensis) is native in the southeastern U.S., where it competes with the invasive brown anole (Anolis sagrei). Elsewhere in the world, the green anoles have been introduced and are themselves the invaders! Photo by Day’s Edge Productions.

Islands have played a special role in illustrating some fundamental patterns of biogeography, including the well-supported observation that smaller and more remote islands harbor fewer species. However, human-mediated dispersal has increased the occurrence of long-distance and over-water dispersal, which were relatively rare prior to human travel on the sea and in the air. Recent studies of anoles on Caribbean islands show that humans have reduced geographic isolation among islands and economic activities now strongly influence patterns of anole biogeography. Islands also provide excellent arenas for experimental investigation of the consequences of anole invasions for native species. After invasive brown anoles arrived on small islands off the Florida coast, native green anoles shifted their habitat use to higher perches, presumably to reduce competition with lower-perching brown anoles. Within a mere 20 generations, green anoles adapted to their more arboreal circumstances by evolving larger toepads, which should allow them to cling better to the more risky higher perches. This is one of the best examples of rapid evolution caused by competition between an invader and a native species. Researchers are interested in studying anole invasions to gain insight both into basic ecological and evolutionary processes, such as species interactions and adaptation, and to mitigate the negative ecological and economic impacts caused by invasive anoles.

A. sagrei

Brown Anole

Appearance: Anolis sagrei may not be the most colorful or strikingly patterned anole, but owing to its widespread distribution and high abundance it may the anole most familiar to you. Adults commonly grow to 7-8 inches from head to tail with females being a few inches smaller than males. Their body color ranges from light tan to gray to dark brown, almost black. Unlike green anoles that can turn brown, these guys are never green! Males have red-orange dewlaps, sometimes with a yellow border or blotches. Females often have a pattern with diamonds, bars, or a stripe running down their back. When in south Florida don’t confuse brown anoles with crested anoles (Anolis cristatellus). Crested anole females only have a cream-colored stripe on their backs, and brown anoles do not have a light ring around the eye or a light stripe above their front limb. Brown anoles often have two dark bars above their eyes.

Ecology and Habitat: The brown anole is a trunk-ground habitat specialist or ecomorph. They are often found on the ground or perching low on tree trunks (up to 1.5 m high). Brown anoles have relatively long limbs for their body size, which allows them to run fast on the ground and tree trunks that they typically occupy. Brown anoles are usually sit-and-wait foragers, keying in on the movement of prey to jump from their perch and attack. Their diet consists of a wide variety of arthropods including most any spiders and insects they can fit in mouths. They will occasionally eat other invertebrates and small vertebrates including their own hatchlings. Brown anoles primarily occupy disturbed and edge habitats in both natural and human-modified landscapes. Their abundances suggest they are just as at home in the city as in natural areas. Like tourists at the beach, these guys love the sun – they are often active during the hottest parts of the day. All anole species display, but brown anoles are the champions – they are often seen fanning their brightly colored dewlaps, and doing head-bobs and push-ups with the intensity and stamina of an Olympic athlete. 

Geographic Range and Biogeography: The native range of the brown anole includes the Bahamas, Belize, Cuba, Cayman Brac, Little Cayman, Swan Island and the Gulf of Mexico/Caribbean coast of Mexico. Its evolutionary origin as part of the 15-species sagrei group is Cuba with multiple colonization events to Caribbean islands east and west of Cuba to round out its native range. One of the best natural colonizers of all anoles, brown anoles are also the undisputed champion of recent human-mediated introductions. Non-native populations are well established around the world including the southeastern United States (e.g. Alabama, Florida, Georgia, Texas), Atlantic islands (e.g. Ascension, Bermuda), Caribbean islands (e.g. Grand Cayman, Grenada, Jamaica, St. Vincent), and Pacific islands (e.g. Hawaii, Taiwan), among other locations with reports of new introductions every year. Genetic analyses reveal multiple origins of these introduced populations from numerous localities in Cuba as well as the Bahamas and Belize. A key discovery about these introductions is that genetic variation from these multiple native-range sources mixes within non-native populations, resulting in increased genetic variation compared to native-range populations.  

Research Highlights: 

If Anolis carolinensis (green anole) is the “lab rat” of the reptile world, then Anolis sagrei is undoubtedly the “field rat.” This species has been studied extensively in the field from populations on small islands in the Bahamas to the heart of the city in Miami. A well-studied natural history, high local abundance, and the ability to tag individuals for unique identification make this species ideal for field studies of natural selection. We have learned that natural selection acts on morphological, physiological and behavioral traits of brown anoles. For example, natural selection favors longer hindlimbs in the presence of terrestrial predators (curly-tailed lizards, Leiocephalus carinatus) and higher optimal performance temperatures in warmer habitats. Other studies show adaptive responses of limbs to local vegetation after being introduced to new islands. A recent study by Oriol Lapiedra and colleagues evaluated whether natural selection acts on variation in risk-taking behavior of brown anoles when they occupy small islands with and without the ground predator Leiocephalus carinatus. Brown anoles that avoided the ground in lab trials were favored in the presence of the predator, whereas more exploratory brown anoles survived better in the absence of the predator. Moreover, selection on behavior and morphology acted independently. This experiment is among the best studies yet showing that natural selection acts on behavior.  

AA post:  https://www.anoleannals.org/2017/06/27/evolution-2017-it-doesnt-pay-to-be-risky-when-predators-are-about/ 

Brown anoles are also a key species in studies of the consequences of global environmental change, including biological invasions, climate change, and urbanization. Recent studies of non-native brown anoles in Miami by Jason Kolbe and his lab discovered that brown anoles living in the city have consistently different personalities compared to their forest counterparts – city lizards are more tolerant of humans, less aggressive, more exploratory in new environments, and bolder after simulated predator attacks. Cities also dramatically alter the ecological niche space occupied by brown anoles. Urban heat islands produce warmer microclimates that allow brown anoles to more often attain their preferred body temperatures during the summer in Miami, likely increasing their ability to persist in some areas. Brown anoles prefer to perch on broad substrates like tree trunks, and urbanization typically results in the removal of smaller trees and the addition of broad diameter artificial structures, such as light posts and utility poles. This leads brown anoles to use broader perches in urban areas despite the fact these artificial substrates are often quite smooth and lizards slip and fall when using them.  

Video:  https://www.anoleannals.org/2016/09/08/lizards-running-and-slipping-in-ultra-slo-mo/ 

For more information:  

Animal Diversity Web: https://animaldiversity.org/accounts/Norops_sagrei/ 

SREL: https://srelherp.uga.edu/lizards/anosag.htm 

The Reptile Database: http://reptile-database.reptarium.cz/species?genus=Anolis&species=sagrei 

More microsatellite markers

Sequencing of the Anolis genome holds great promise for unlocking the genetic basis of anole phenotypic variation – such as dewlap coloration and limb length differences – and it also makes for a nifty way to discover new molecular markers, such as microsatellites.  Wordley et al. report in a recent article on mining A. carolinensis expressed sequence tags (ESTs) for repeats and then blasting the EST-derived sequences against the genome to obtain the genomic sequence and its location on assembled chromosomes.  From these sequences, they designed primers, tested them out in A. carolinensis, and, importantly, attempted to amplify them in multiple, phylogenetically diverse species.  They identified 8-25 new variable markers for apletophallus, carolinensis, distichus, porcatus, and sagrei, which can be added to the existing resources designed for carolinensis, cristatellus, distichus, luciae, roquet, oculatus, and sagrei, which also work for some related species.  Happy genotyping!

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