Hello to everybody, I’m an italian naturalist that visited Cuba last December 2016.
I’m mainly a birder, but I like to give a name to all the creatures I meet. So, I’m going to post 20 pictures of lizards photographed in Cuba: for some I have hypotheses about the identification, but I need confirmation. For some others, I’m completely lost!
Can anybody help me??
Sexual dimorphism, or differences in size or appearance between the sexes, was used by Darwin to explain sexual selection in On the Origin of Species. Interestingly, sexually dimorphic traits, like antlers in deer or showy feathers in peacocks, often do not present themselves until the animals are reproductively mature. Juveniles are often sexually monomorphic, or relatively similar in appearance. Few studies have investigated how sexual antagonism, when different sexes have different optimal strategies, of these traits may develop in the wild over the course of the animal’s maturation. So Aaron Reedy from the Bob Cox Lab at the University of Virginia decided to tackle this question with brown anoles (Anolis sagrei) in Florida.
After tracking thousands of lizards for several generations, Aaron found that selection changed throughout an animal’s life. Adult lizards had selection pressure for large males and small females in one year, but reversed the next, which was surprising, but in both cases was still sexual antagonism. For juveniles, on the other hand, larger body sizes were better regardless of sex. This is an example of sexually concordant selection, where both sexes have similar optimal strategies. He also found that there was selection pressure on the dewlap (an important ornament of anoles in courtship displays) to be smaller in one year, but then reversing the next so that males with larger dewlaps had better chances of survival. This year-to-year variation in selection is interesting and hopefully we’ll learn more from this system in the future.
Anoles are probably best known for the ecomorph story: the presence of specialized species adapted to the same sets of structural microhabitats on different islands. Anoles in the Greater Antilles have contributed hugely to our understanding of both the evolutionary history and the contemporary ecology of communities of specialists.
While they are better known for specialization of species in communities, anoles have also contributed to our understanding of within-species ecological diversity. Around the same time that Ernest Williams was developing the ecomorph concept, Roughgarden (1972) used data from Lesser Antillean anoles to introduce a new framework for investigating the extent to which a population’s niche width (i.e. the diversity of habitats it uses or prey it eats) is determined by variation among individuals versus variation within individuals. For example, individuals in a population of Anolis roquet differ in the size of prey they consume, mainly because larger individuals can catch and ingest larger prey items. While Roughgarden’s early work set the stage for an explosion of studies of individual specialization over the past decade or two (reviewed in Araújo et al. 2011), surprisingly little work has been done to revisit individual specialization within species of anoles. In particular, we don’t know enough about how much individuals specialize in important aspects of microhabitat that differentiate ecomorphs, especially perch height and perch diameter.
Anole Annals contributors Ambika Kamath and Jonathan Losos have helped to fill this gap with a study just published online in Evolution. Ambika and her team spent a summer observing microhabitat use of a population of brown anoles (Anolis sagrei) in a forested park in Gainesville FL. They marked lizards with colored beads, and repeatedly recorded individual lizards’ perch height and diameter, compiling a total of over 1000 observations of 80 anoles. They grouped perch heights and perch diameters into classes, then compared the distribution used by each individual to the distribution used by the whole population (or to the distribution available to that individual) using a proportional similarity index. The mean value of this index gives a measure of the overall degree of individual specialization in a population, as lower overlap values tell us that individuals are specializing on a subset of the available perches.
Continue reading Anoles Are Habitat Specialists at the Individual Level Too
I’ve recently learned that famous nature micro-photographer Piotr Naskrecki observed an aquatic anole catching prey underwater. Here’s what he had to say on his blog, The Smaller Majority:
The other aquatic iguana
In a couple of days I am heading off to the Galapagos Islands, where I hope to be able to see the incredible marine iguanas, the world’s only truly marine lizards. Other lizards enter water occasionally, but aquatic lifestyle is quite rare among these reptiles, and few species live and feed under water. But in rainforest streams of Central America there is one little known species of iguana that does just that.I first saw the aquatic iguana (Norops aquaticus) in the southern part of Costa Rica in 1994. These lizards swam and dove in a fast-flowing stream, catching water insects. But when I told a herpetologist friend about it, she refused to believe me.
It took me 13 years to find the aquatic iguana again, and this time I had a camera with me. It was in a different part of Costa Rica (Est. Pitilla in Guanacaste), but the animal and its habitat were the same. I watched it for a couple of hours, following the lizard among slippery boulders, hoping to document its hunting behavior. Eventually I got lucky, but alas, the actual catching of the prey happened underwater, when the iguana cornered a nymph of an aquatic blattodean (a yet undescribed species.) Next time I will definitely try to get a photo of the underwater action.
Update (2 Sept 12): Turns out that the aquatic Norops iguanas that I saw in southern Costa Rica and those from the northern part of the country, shown here, are different species. The animal in the photos is Norops oxylophus, not N. aquaticus. You can read more about the amazing aquatic behavior of N. oxylophus here. (Thanks to Annemare Rijnbeek for pointing me in the right direction regarding the ID of these animals.)
Incidentally, it appears that these lizards are once again being placed in the genus Anolis, where they historically belonged.
This post was written by Brittney Ivanov, research technician in Michele Johnson’s lab at Trinity University.
Catalina Mantilla, a Ph.D. candidate at Florida International University working with Tonia Hsieh of Temple University, is interested in how anoles use their toepads and claws when they run. For most animals, movement on vertical perches such as tree trunks or buildings usually requires specialized morphologies to adhere to these substrates. While many species have evolved adaptations for moving through complex arboreal habits (e.g., prehensile tails or feet, sticky pads, spines), anoles evolved enlarged toepads and distinct claws, presumably to allow for better adhesion. The morphologies of these specialized structures can greatly impact performance; for example, greater toepad area is associated with greater clinging ability. Catalina wanted to better understand how toepads and claws work together to enhance running performance.
Catalina collected 17 males from four Anolis species (A. carolinensis, A. sagrei, A. cristatellus, and A. distichus). Each male was tested in four different running courses to test performance at difference inclines and on different substrates. Two of the courses were positioned at a 45° incline and two at a flat (0°) incline. Plexiglass covered one course at each incline to allow the use of toepads and eliminate the use of claws. Nylon mesh covered the other course at each incline to test the use of both toepads and claws. Performance was evaluated using mean relative sprint speed, relative stride length, and stride frequency.
Catalina found, unexpectedly, that when the lizards ran on the level plexiglass, they ran slower, took shorter strides, and increased their stride frequency compared to when they ran on the inclines. These results suggest that anoles are less stable when they can’t use their claws! in addition, these data support the idea that the combination of toepads and claws is important for their running performance. In the future, Catalina hopes to increase the number of species in this study to determine the effect of ecomorph on claw and toepad interactions during running, and to evaluate limb function changes when running across different inclines.
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Regular readers of AA will be familiar with the differences in microhabitat use that define the Anolis ecomorphs, but do species with such distinct structural habitats move differently on their specialized perches? In other words, does muscle function differ between the ecomorphs? In the very last session at this year’s SICB, Kathleen Foster, currently a postdoctoral researcher at the University of Ottawa studying the biomechanics of fish locomotion (come back to anoles, Kathleen!), presented a portion of her graduate work in Tim Higham’s lab at the University of California, Riverside, to address this question. She used high speed video to film five species of anoles running on broad and narrow perches at two angled inclines, combined with electromyography to record fore- and hindlimb muscle activity during running.
Kathleen found that all five species had greater motor unit recruitment on steeper inclines than on horizontal perches, and that muscle activity is shorter but begins more abruptly on inclines. Further, recruitment of the gastrocnemius (a “calf” muscle) was greater on broad perches, because the way lizards sit on narrow perches limits the function of this muscle. If you’ve seen how anoles position their feet on both sides of narrow perches, it’s easy to understand how this posture prevents effective propulsion by ankle extension. Kathleen also found several intriguing differences that distinguish trunk-ground species’ muscle function from trunk-crown and crown-giant species. The activity of the caudofemoralis (a limb retractor muscle in the hindlimb) changes more in trunk-ground species as a function of incline, and trunk-ground species use the biceps and gastrocnemius more in the early stance phase of propulsion than trunk-crown species.
Overall, these data help us understand how specialization in neuromuscular function can allow different anole species to successfully move through their varying habitats, and offer insight into how behavioral differences depend on the muscles that underlie them.
Above: Albert Chung presented on how testosterone influences sexual signals and energetics at the annual Society for Integrative and Comparative Biology meeting in New Orleans, LA.
Many researchers are curious about how sexual signals evolve and if those signals are influenced by an individual’s energetics. Signal expression (such as anole dewlap size or color) and energetics might have the same physiological underpinnings, and signal expression may be limited by energetic state. Albert Chung, a graduate student working with Dr. Christian Cox at Georgia Southern University, along with graduate student, Aaron Reedy, and Dr. Robert Cox at the University of Virginia tested this hypothesis with brown anoles (Anolis sagrei) in the lab and in a wild population in the Bahamas.
The researchers had three different groups in this experiment. In one group, the males were castrated, so they wouldn’t produce natural testosterone. These males were each given a small implant that released synthetic testosterone designed to have the same effects on the body as natural testosterone. In the next group, males went through a procedure to mimic castration, but were effectively left intact, and implanted with an empty implant. These males still produced natural testosterone. In the final group, males were castrated and each one was given an empty implant, so they did not have testosterone in their bodies.
Wild anoles were recaptured after two months. The researchers measured the size and color of the dewlap, how much fat was stored (which provided the researchers with an estimate of energetic state), and body condition. They took the same measurements in the lab population.
There were clear differences between the males that had testosterone (both natural and synthetic) and males that did not have testosterone. Castrated males had smaller dewlaps compared to intact males and testosterone-treated males in both the field and lab populations. Dewlap size of testosterone-treated males was similar to that of intact males in the wild population, but in the lab population, testosterone-treated males had larger dewlaps than intact males.
Castrated males in the wild population had brighter, more saturated dewlap coloration than testosterone-treated males and intact males. In the lab population, dewlap coloration did not differ among the treatments.
In the wild population, castrated males had higher body fat mass than intact males and testosterone-treated males. Wild testosterone-treated males were similar in body fat to intact males. Castrated males in the lab population also had higher body fat mass than intact males and testosterone-treated males.
The researchers also looked to see if either fat mass or body condition were correlated with dewlap size or color within each treatment group. None of these variables were correlated with one another.
Overall, while dewlap expression was not dependent on an individual’s energetic state, both energetics and dewlap expression were directly influenced by a common hormone: testosterone.
Urban environments are widespread and expanding across much of the earth, and this urbanization likely affects the flora and fauna in its path. Anoles are no exception and are frequently observed perching on anthropogenic structures. Thus, Chris Thawley, a post-doc in Jason Kolbe’s lab at the University of Rhode Island, and colleagues wondered how the abiotic and biotic changes in urban areas influence anole traits.
Thawley compared populations in urban and natural habitats of two species that we’re quite familiar with on Anole Annals – the Brown Anole (Anolis sagrei) and the Puerto Rican Crested Anole (Anolis cristatellus). Thawley found that A. sagrei prefers warmer temperatures than A. cristatellus, but that urban anoles do not differ in thermal preference than natural anoles for either species. Alternatively, urban male A. cristatellus and both sexes of urban A. sagrei were larger than their natural counterparts. As for parasites, A. sagrei had a higher parasite prevalence than A. cristatellus, but urban anoles did not differ from natural anoles in either species in parasite prevalence. However, for the A. sagrei that were parasitized, urban A. sagrei had higher parasite loads than natural A. sagrei.
These findings show that urbanization can influence anole morphology and parasite ecology. Thawley has just begun this work, and I look forward to seeing his future research on anole adaptation to urban environments!
Most animal learning studies have been conducted in the lab with the assumption that those findings are representative of behavior in the field. However, assessing behavior in the field increases ecological relevance. In addition, birds and mammals have received much of the attention in cognitive studies. Yet we on Anole Annals know that these lizards can be quite clever.
Levi Storks, a Ph.D. student in Manuel Leal’s lab at Mizzou, set out to address these issues by designing a method for testing behavioral flexibility in brown anoles (Anolis sagrei). Wild lizards in the Bahamas were allowed to feed unrestricted on a maggot placed in the middle of a testing apparatus in order to acclimate lizards to the structure. Storks then used a clear plastic tube to block the direct route to food, requiring lizards to move to either end to gain access. Lizards that successfully completed this task were then tested to see if they could associate unique patterns on the ends of the tube with single openings.
Storks found that a subset of lizards could successfully complete the first detour task, and lizards made fewer errors over the course of solving the detour task. These findings suggest brown anoles can learn and exhibit behavioral flexibility. Stay tuned for more of Levi’s work as he’ll be applying these methods to assess differences in behavioral flexibility between populations that vary in ecology!
The beautiful island of Dominica used to be home to only one anole (Anolis oculatus), but about 20 years ago, the Puerto Rican crested anole (Anolis cristatellus) showed up. Jeanel Georges, a graduate student in Matt Watson’s lab at Midwestern State University in Wichita Falls, Texas who is originally from Dominica, noticed that while A. oculatus occurs in all the ecological zones of the island, A. cristatellus is absent from the cooler, wetter uplands. With an international group of collaborators, Jeanel examined the thermal habitat use, sprint speed, and bite force of both species to determine what may limit the spread of A. cristatellus across the island.
At a lowland site where the two species co-occur, both species had higher body temperatures that the operative temperatures randomly available in the environment. In the much cooler upland site, A. oculatus had much higher body temperatures than the operative models, but these body temperatures were cooler than that species experiences in the lowland site. Jeanel also found that the two species had stronger bite forces and higher sprint speeds in the lowland site than A. oculatus had in the upland site. These data suggest that A. cristatellus and A. oculatus are partitioning the thermal environment of Dominica, and as climate change alters the temperatures available to lizards on the island, the interactions between these two species may change.
This post was written by Brittney Ivanov, research technician in Michele Johnson’s lab at Trinity University.
Austin Hulbert, an undergraduate in Dan Warner’s lab at Auburn University, presented a poster on the behavior of brown anoles (Anolis sagrei) in a novel environment: a few very hot greenhouses in Auburn, Alabama. Brown anoles are an invasive species, most notably in Florida, but some populations have been found farther north in states including Louisiana, Alabama, Mississippi, and Georgia. As ectotherms migrate to higher latitudes, they often have to deal with different thermal environments and must alter their behaviors accordingly. Austin was interested in determining the activity patterns of a population of brown anoles inhabiting a group of greenhouses in Alabama.
During the summer he found that temperatures inside the greenhouses were consistently higher than those outside. Temperatures drastically increased each morning, up to peak temperatures between 11am and 3pm (on average, 45°C inside the greenhouse and 37°C outside). In the evenings, the temperatures again cooled. Austin surveyed the greenhouses and the surrounding areas for anoles during the morning, peak, and evening hours and determined the type of substrate each individual was using (i.e. brick or concrete, ground, metal, or wood). On average, brown anoles were more abundant inside the greenhouses than outside during the morning and peak times. He also found that more of the brown anoles perched on wooden substrate in the morning and evening. During peak hours more lizards perched on the ground. Because temperatures are often cooler closer to the ground, the lizards may be altering their behavior to deal with the extreme heat in the greenhouses during the hottest part of the day. While the visual survey focused on lizards perched in the open areas visible to the surveyor, there may have been individuals hiding under undisturbed objects as a means to keep cool during peak hours. In the future, Austin would like to compare the thermal tolerance of this group of brown anoles to those of populations in Florida to determine if inhabiting these greenhouses has resulted in adaptions to tolerate their more extreme temperatures.
This post was co-written by Maria Jaramillo, an undergraduate in Michele Johnson’s lab at Trinity University.
A mother’s experience during gravidity may alter her offspring’s development, particularly through altering hormone levels in the yolk of her eggs. Stress hormones such as corticosterone (CORT) alter various aspects of offspring phenotype following in ovo exposure, and physical exercise elevates CORT in many vertebrates. In the work he presented at SICB, Jerry Husak and colleagues used exercise and food restriction to manipulate female Anolis carolinensis CORT, and to then determine whether the increased CORT was transferred to the females’ egg yolks.
Jerry assigned females to one of four treatments with different combinations of exercise and food restriction: 1) no exercise, regular diet; 2) no exercise, restricted diet; 3) exercise, regular diet; and 4) exercise, restricted diet. He found that maternal exercise increased maternal CORT (as expected), but surprisingly did not result in higher CORT in the eggs. Further, diet restriction did not affect maternal CORT, but moms with restricted diets laid eggs with reduced CORT.
This study suggests that anole mothers may manipulate the environments of their eggs in ways we don’t yet understand – the mechanisms by which CORT is transferred to eggs is an area ripe for future study!
Every year since 2013, the Division of Ecology and Evolution (DEE) hosts the Huey Award Symposium at the annual SICB meeting. The Huey award is given for the best student presentation in DEE, in honor of Ray Huey, professor emeritus at the University of Washington. Ray’s career featured a lot of key research on anoles, and so there is often good representation by anole biologists at the Huey award. At this year’s symposium, Ariel Kahrl, a graduate student in Bob Cox’s lab at the University of Virginia, presented her research on pre- and postcopulatory selection in Anolis lizards.
We know that male competition for mates occurs both before copulation (mating success) and after copulation (sperm competition). Her research focuses on investigating the evolutionary connection between these two phases of competition. She found that larger males have smaller relative testis size, indicating a tradeoff between pre- and postcopulatory success, as larger males will have better success gaining access to females, but less sperm available for mating.
When she looked at testis and sperm morphology in greater detail, a few interesting patterns emerged. First, she found that testis size evolves faster than body size, consistent with other studies showing that reproductive organs evolve faster than other body traits. She also found that the midpiece section evolved faster than the head and the tail of the sperm. Importantly, the midpiece section of the sperm was strongly associated with male condition and sperm swimming speed, whereas the head and tail of the sperm were not associated with male condition or sperm swimming performance. She further hypothesized that sperm count may be a more important target of selection than sperm morphology.
The effects of climate change and urbanization on reptiles and amphibians has been a major topic at this year’s SICB. Both are expected to cause drastic changes in the climate, which will likely be severely detrimental to many species. We hope that many species will be able to tolerate these changes by adapting or acclimating, either by thermoregulating or changing their physiology. Adults of many species are able to acclimate in this way, but Josh Hall of the Warner lab wanted to test if eggs (which are unable to move to thermoregulate) are able to acclimate their physiology in response to higher average temperatures and to spikes in temperature that you might find in urban environments. Josh collected two populations of wild A. cristatellus from Miami, an urban population and a forest population, brought them back to the lab, and collected their eggs.
He then put the eggs into five different thermal conditions 1) higher “urban” temperatures, 2) cooler “forest” temperatures, 3) “urban” temperature with a large temperature spike on day 8, 4) “forest” temperature with a large spike on day 10, and 5) “forest” temperature with a small spike on day 10. He predicted that eggs that had a higher baseline temp (i.e. the urban eggs), would be able to tolerate spikes in temperature better than eggs at lower temperatures and would have higher survival and less of a physiological stress response due to the temperature spike. Contrary to his hypothesis, he found that high temperatures, and spikes were both detrimental to the survival of eggs and hatchlings, and affected baseline and max heart rate in embryos. This is somewhat concerning because it suggests that even a single short burst of heat can kill embryos, and have lasting affects on juveniles. The bursts, which you might expect in urban areas, have a more profound affect when the background temperature is higher, which will likely happen due to climate change.
Marathon runners and elite sprinters, like Usain Bolt, have dramatic differences in their muscle physiology that allow them to specialize in their respective track-and-field events. Whereas sprinters have lots of muscle fibers that produce high force but fatigue quickly, marathon runners have lots of muscle fibers that produce less force but allow much longer activity because of their reliance on aerobic respiration. Might this be true for our beloved Caribbean anoles, too? Faith Deckard of Michele Johnson’s lab at Trinity University tried to answer that very question. She studied six species of anoles in the Dominican Republic to test whether anoles that have higher rates of dewlap extension and extend their dewlap for a longer duration have dewlap muscles with a higher proportion of slow-twitch muscle fibers that can be used for endurance. Surprisingly there was no significant correlation between the two behavioral traits and the proportion of slow-twitch fibers! However, this scrutinizing attendee feels pretty strongly that there is a relationship that is just yet to be teased apart statistically. The raw data Faith presented looked very convincing to me, so we’ll see what the future holds for this question. Faith’s results are a very interesting clue to the still-elusive mechanisms that underlie anole behavioral diversity.
All of the gumbo, Po boys, and beignets consumed by attendees of SICB 2017 have to go somewhere after consumption. Much of the energy contained in those delicious foods is used for very important maintenance functions in your body: metabolism, cell repair and replacement, and your immune system. What’s left over after maintenance costs can then be divided amongst other tasks, such as reproduction, movement, and wide variety of other tasks. Unlike humans, anoles do not have unlimited access to gigantic portions of gumbo, so their energetic investments require much harder decisions. Once energy from a cricket, for example, has been put into the immune system, it can no longer be used for making eggs or patrolling a territory a little bit longer. Andrew Wang of Jerry Husak’s lab at the University of St. Thomas was interested in what mechanisms are involved with anoles making these investment “decisions.” He did this by forcing allocation of resources to an energetically expensive trait (endurance running) by exercise training lizards to see what would happen to everything else that they might invest in.
Previous work showed that exercise training and diet restriction results in dramatic trade-offs with reproduction and the immune system. He suspected that what might explain this suppression was the hormone leptin, which is made by fat cells (yours make it, too). Since bigger fat cells means more leptin in the body, leptin can be thought of as a signal to the brain and body of how much resources are available for investment. Indeed, without sufficient leptin, reproduction grinds to a halt from the brain downward. Much like elite athletes, Andrew’s marathon lizards have little to no fat stores in their body, thus suggesting a role for leptin. To address this question, he supplemented half of the lizards with leptin (the rest got only saline as a control) to see if he could “rescue” immune function and reproduction. Interestingly, he found that leptin did rescue his measure of immunity, but it did not rescue reproduction. He attributes this latter finding to either (1) a lack of energetic resources to produce eggs even if there is a leptin signal or (2) the stress of the leptin injections over-rode the leptin signal in the brain where reproduction is controlled. His results suggest some very complex interactions in physiological pathways that can result in the trade-offs observed in many animal species.
What does it take be a good sprinter? How about a marathon runner? One might think that the traits responsible for such performance traits would be the same in males and females. If you are a green anole, that just isn’t true. Annie Cespedes, working in Simon Lailvaux’s lab at the University of New Orleans, explored the multivariate predictors of seven performance traits (sprint speed, bite force, cling force, exertion, endurance, jump power, and climbing power) in male and female green anoles. Annie explained how animals in nature rely on lots of different performance traits in their daily lives, and the large difference in body size and shape between male and female anoles might mean that the two sexes use different means to be successful in life. To add to this complexity, some individuals are just better overall at ALL performance traits than others (imagine a couch potato versus a very fit athlete), and one must account for this to understand what shapes anole performance.
Multivariate statistics allowed Annie to show that males and females do indeed differ in performance, but only in clinging ability, sprint speed, bite force, and jump power. Even more interesting, the suites of morphlogical traits that explained performance ability differed substantially between the sexes. For example, small females with large leg muscles were better sprinters and jumpers than females who are smaller and are better biters and endurance runners. What is especially important about Annie’s research is her approach. When considering how animals evolve, one must do so by simultaneously looking at a multitude of traits that might impact their survival and reproduction. By knowing how morphology predicts performance, we can begin to better understand how evolution will shape that morphology when selection acts on those performance traits.
Frequent readers of Anole Annals are likely to recall the amazing convergent evolution of morphology related to habitat use in Caribbean anoles that coincides with similarly striking convergent evolution of social behavior. Most of what we know about behavior of Caribbean anoles is how males behave: there are major differences among ecomorphs in how often males use their colorful dewlaps and how often they mate with females. Such male-typical behavior seems intuitively linked to species differences in testosterone signaling. Previous work has shown, though, that these differences do not seem to be related to levels of testosterone in the blood, so Miguel Webber of Michele Johnson’s lab at Trinity University examined whether the receptors for testosterone varies in a manner consistent with the behavior for six Dominican Republic species of anoles and one U.S. species.
Hormones can only cause effects on tissues that have receptors for them, so Miguel looked at receptors for testosterone (androgen receptors) in the muscles responsible for moving those fabulous dewlaps (the ceratohyoid muscle), expecting to find a correlation across species between the number of androgen receptors in the muscle and the rate of dewlap extensions. Although the data are still preliminary, there was a trend for males with higher dewlap extension rates to have more androgen receptors in the ceratohyoid muscle. His next steps are to look for an association between rates of copulation and androgen receptors in the muscle used by males to copulate (retractor penis magnus muscle – yes, it does what you would guess based on the name…). He also wants to see if there is a correlation among species in the behavioral traits and androgen receptors in regions of the brain that are important for social behavior regulation.