Climate change on earth is accelerating. These changes will have important impacts on all species, but some types of organisms are predicted to be affected more strongly than others. One such group is ectotherms which use the temperatures available in surrounding habitats to regulate their body temperatures. Another such group is mountaintop endemics. These species are restricted to one or several mountain peaks by climate and/or competition with other organisms. As such, they cannot easily disperse to other areas if climate makes their current habitat unsuitable!
Mountaintop endemic species may be particularly vulnerable to climate change (Chand Alli, CC BY SA).
Hispaniola contains several high elevation areas home to mountaintop endemic species, including anoles (NASA).
Many studies use correlative modeling approaches (often termed ecological niche models [ENMs] or species distribution models [SDMs]) to assess a species’ current distribution and predict its future distribution by projecting it into simulated future climate scenarios. This approach has some advantages including ease of implementation across many species. However, it has at least two potential drawbacks: the environmental data used in building such models are often measured at a fairly coarse scale that does not represent how many organisms use their environments, and the models do not explicitly include biological processes such as physiology and behavior.
Anolis armouri in a montane rock meadow (Reptile Database).
Vincent Farallo, a post doc at Virginia Tech, and his advisor, Martha Muñoz (both moving to Yale in a few weeks!), investigated whether incorporating physiology and behavior into modelling might affect predictions of climate change impacts on two mountaintop endemic anoles of Hispaniola, Anolis armouri and Anolis shrevei. Correlative SDMs (via BioMod2) predicted both species would lose much or all of their suitable habitat under climate change, perhaps leading to extinction. However, when Vincent constructed mechanistic niche models (via NicheMapR) that included knowledge about the thermal physiology and habitat use behavior of these species to predict activity time, they showed that habitat would increase in suitability under climate change, the opposite result! Interestingly, these models also predicted increased suitability for a widespread anole, A. cybotes. This result suggests that while climatic changes may not be a direct threat to these mountaintop anoles, increased competition with another anole, an indirect impact of climate change, may be.
Activity time of Anolis shrevei is predicted to increase across its range in Hispaniola with climate change (Farallo and Munoz).
As a whole, Vincent and Martha’s work shows that incorporating more mechanistic knowledge into models, including physiology and behavior, may be critical to predicting the impacts of climate change on organisms and making sound conservation decisions.
Periodically here on Anole Annals, we have posts about three-legged lizards. The most recent such post was last year from Miami. Here’s another lizard, with a twist: it’s got four legs, sort of. Looking at the floppy left hindleg of this lizard, caught in the Bahamas two years ago. An x-ray confirms that this is odd–there’s no bone in most of that limb! I’ve never seen anything like it, and wonder how it happened.
Despite this seeming impediment, the lizard looked quite healthy, and as the video shows, could run quite adeptly up a note pad.
And here she is when we released her back at the place where we caught her. Pretty nimble!
It seems hard to believe almost a year has passed since the last Evolution meeting. Last year we brought you coverage of the anole talks and posters in Montpelier, France. This year, we’re coming to you live from Providence, Rhode Island from June 22nd – 25th! This year there are eight talks and eight posters scheduled *(searching the schedule for keywords “anole” and “Anolis“). There’s some pretty fascinating topics on the schedule – here’s what you have to look forward to each day:
Saturday:
Habitat use, competition, and phylogenetic history shape the evolution of claw morphology in Lesser Antillean anoles (Yuan, Jung, Wake, Wang)
The effects of volcanic activity on the phylogeographic history of the Plymouth Anole, Anolis lividus, on Montserrat (Poster board #72) (Jung, Yuan, Wang, Frederick)
Identification and assembly of an anole sex-chromosome: Rapid degeneration since autosomal fusion? (Poster board #160) (de Mello, Hime, Glor)
Effects of urbanization on toe pad shape and lamellae size in Anolis cristatellus (Poster board #174) (Howell, Hagey, Winchell)
Monday:
Using archival DNA to elucidate anole phylogeny (Mayer, Gamble)
Comparative landscape genetics and epigenetics of two trunk-ground anoles (Wang, Wogan, Yuan, Mahler)
Ancient hybridization in the adaptive radiation of Anolis lizards on Puerto Rico (Wogan, Yuan, Wang)
Urban adaptation in anole lizards of the Greater Antilles (Poster board #7) (Winchell)
Cities in the Spotlight: Does Tolerance of Artificial Light at Night Promote Urban Invasions? (Poster board #97) (Thawley, Kolbe)
Adaptive radiation in the multidimensional phenotype (Bodensteiner, Muñoz)
Patterns of morphological diversity in Draconura clade anole lizards (Huie, Prates, Bell, de Quieroz)
Did we miss any? If so, let us know in the comments so we can be sure to add it to our schedules! We will be live blogging the meeting as usual, so check back starting June 22nd to hear about the latest in anole evolutionary research. And if you are attending the meeting, consider blogging a talk or poster for us (new contributors welcome!). Just send me an email and I will fill you in on all you need to know.
Anolis (Phenoacosaurus) heterodermus, a mainland anole that co-occurs with few or no other anole species
One of the most important questions in ecology and evolution is about the role of biotic interactions in driving phenotypic and behavioral changes across species. The insular Anolis species are a good model to address this kind of question due to their high abundance and pervasive ecological interactions across islands. Some insular species, however, live in isolation on small islands across the Pacific and Caribbean islands (21 species). These species have evolved similar morphologies across islands. For instance, Poe et al (1) found that body size evolved by exaptation (remember the classic Gould and Vrba 1982 paper) to colonize these small (and depauperate) islands successfully. By contrast, Poe et al. (1) showed that sexual size dimorphism (SSD) evolved by adaptation likely after island colonization to minimize intraspecific competition.
In brief, these solitary insular anoles evolved phenotypic (body size and SSD) traits by two different processes. Cool! But, what happens in mainland areas? Much work has been devoted to Caribbean species, but the mainland offers many more species and very little research has been conducted there to understand ecological and evolutionary processes. So, we decided to establish whether solitary ecology can be extended to mainland species or whether it is an island ecological phenomena.
The first problem that we had to resolve was trying to establish whether mainland species tend to live in geographical/ecological isolation as insular species. We adopted a novel concept in macroecology (the diversity field concept) developed by Mexican macroecologists (Hector Arita and Fabricio Villalobos see 2, 3) implemented here using extensive distributional information for almost all known Anolis species (377 spp), which I generated during my Ph.D. thesis (see 4 for an example using these maps). The diversity field concept allows us to establish how many species co-occur with a given species across its geographic range.
We calculated how many congeners can co-occur within the distributional area of each Anolis species using the range maps (see figure below). We divided mainland species into two groups: those co-occurring with few congeners (i.e., “solitary-like”, I had to say that his term did not like to reviewers, so we used a “species-poor” forms in the paper). Then, we test whether these “solitary-like” mainland species are different from other mainland species using a randomization approach. Our results revealed that “solitary-like” mainland species exhibit different traits from random mainland assemblages. These unique traits (i.e., uniform body size and greater SSD) suggest that solitary ecology from insular anoles can be extended to mainland settings.
Figure. Diversity fields for some Anolis species. Note that the diversity field is the set of richness values of co-occurring anoles inside each distributional area.
The next question was focused to establish whether similar (ecological and evolutionary) processes affected body size and SSD patterns in a similar way. We found that the phylogenetic position of body size and SSD shifts did not coincide and also with the evolutionary transitions to solitariness (i.e., reduced level of sympatry). We suggested that both traits are decoupled across the entire Anolis radiation and likely that both traits evolved exaptatively. In other words, it is possible to think that “solitary-like” species retained body size and SSD from their most recent common ancestors to facilitates the lonely life.
The paper is very short (less than 2500 words) and was published in the May number of Biology Letters(5).
In the most recent issue of Nature Ecology & Evolution, first-author Rob Pringle gives the inside skinny on the recent paper about the interaction of predation and competition among lizards (see the video description of the study).
Herewith, the essay:
I was heavily influenced by a handful of papers that were published during my first few years of graduate school. Some of these — Fine et al. (2004) on how herbivores promote habitat specialization in trees, Rooney et al. (2006) on food-web structure and stability — resonated because I could connect them to problems that I was working on. Others, such as Schmitz et al. (2004) on the ecological importance of predator-avoidance behavior, made an impression because they seemed to herald seismic shifts in the outlook of community ecology. And then there was a set of papers that captivated me with their sheer elegance: beautifully designed and executed field experiments that inspired me and made me jealous.
A string of papers by Tom Schoener, Dave Spiller, and Jonathan Losos belonged to this last category. There was a new one each year, with titles like “Predator-induced behaviour shifts and natural selection in field-experimental lizard populations” (2004) and “Island biogeography of populations: an introduced species transforms survival patterns” (2005). These studies used tiny cays in the Bahamas as arenas for a simple yet powerful experiment. On some islands, the investigators had introduced top predators — curly-tailed lizards (Leiocephalus carinatus), which occurred naturally on larger islands just a few stone-throws away from the experimental islands. A major aim of this work was to understand how the introduced predators affected populations of brown anoles (Anolis sagrei), which were native to the experimental islands.
The results were dramatic. Curly-tailed lizards are stocky, ground-dwelling animals, and they devastated brown-anole populations. The brown anoles that survived did so by climbing into the vegetation, beyond the reach of the curly-tails, and this behavioral shift was associated with natural selection on hindlimb length — an evolutionary consequence of predator-avoidance behavior. When I was a kid, we used to play a game called ‘the floor is lava’; if your feet touched the ground, you were dead. It seemed kind of like that for the brown anoles on these islands.
When I started a post-doc at the Harvard Society of Fellows in 2009, I met Losos and we started discussing ideas for a new experiment. I thought that a minor innovation on the earlier experiments could open up new conceptual territory. Losos said that he’d been wanting to do the same thing for years. To wit: if we introduced not just curly-tailed lizards, but also a second species of anole — green anoles (Anolis smaragdinus) — then we could ask questions about predation, competition, and the interaction between the two. Among other things, this design would enable us to test classic ideas about how predators affect the ability of competing prey species to coexist.
The risky thing about this idea was that so much of it had already been done to one degree or another. Previous work had painted a rich picture of the interaction between curly-tailed lizards and brown anoles — our odds of discovering something new on that front were low. And there were dozens of studies about competition between sympatric Anolis lizards. The novelty of our approach hinged on the interaction between predation and competition, which was a thin thread on which to hang such a massive undertaking. But I felt supremely confident that the experiment would work. Todd Palmer, Rowan Barrett, and a raft of other collaborators must have been confident too, because they joined me in setting up and monitoring the experiment.
By 2013, two full years into the study, my confidence was giving way to panic. I had started a job as an assistant professor in 2012; I was anxious about my professional survival, and I had ploughed large amounts of time and money into an experiment that did not seem to be working after all. The introduced curly-tailed lizards were firmly established in their new homes, and the brown anoles were responding by becoming more arboreal, as previous work had indicated they would. But the introduced green-anole populations seemed to be struggling. It looked as if they might die out, in which case our experiment would amount to a very expensive confirmation of the earlier work by Schoener, Spiller, and Losos. Our project had some original twists — Tyler Kartzinel was spearheading an effort to monitor the lizards’ diets using DNA metabarcoding — but it wasn’t at all clear that we would discover anything new or noteworthy.
Our break came in 2014, when it became clear that the green-anole populations were indeed thriving on some islands — just not on any of the islands with curly-tailed lizards. When we returned to the Bahamas in 2015, buoyed by the emerging results and freshly funded by the US National Science Foundation, we found that green anoles had disappeared on one island with curly-tailed lizards (the largest such island). Sometime during 2016, a second green-anole population vanished, this time from the smallest island with curly-tailed lizards. That left just two islands where green anoles still persisted in the presence of curly-tails, and one of those populations looked like it might soon join the list of casualties. Was this because the curly-tailed lizards were simply eating the green anoles to extinction on those islands? Probably not. The green anoles were highly arboreal; they rarely descended to the ground and instead moved by scampering through the twigs and leaves in the canopy. The chunky curly-tailed lizards, by contrast, lumbered across the ground, rarely climbing higher than 50 centimeters — and then only on the thickest of tree trunks. Indeed, the curly-tails didn’t seem to be eating many lizards of any kind. We saw them feasting on cockroaches, and occasionally snacking on fallen fruits and dead hermit crabs, but it wasn’t until 2016 that we finally saw one eat a small female brown anole. Isotopic analysis revealed that curly-tailed lizards actually occupied a slightly lower trophic position than did either anole species, which suggested that the top predator was subsisting more on insects than on other lizards.
The more plausible explanation for our results was that the presence of curly-tailed lizards intensified competition between the two anole species within the predator-free arboreal refuges, and that this competition — not direct predation — was the primary reason why the introduced green-anole populations failed to increase on islands with curly-tailed lizards. Molecular analysis of fecal samples subsequently reinforced this impression. DNA metabarcoding produced evidence that curly-tailed lizards exacerbated the competition between brown and green anoles for insect prey. And a quantitative PCR assay — conducted by Charles Xu at the behest of one of the four very thoughtful reviewers for Nature — detected the DNA of brown and green anoles in just 4% of the curly-tailed lizards that we sampled. Curly-tailed lizards really were the top predators; they just didn’t catch anoles very often.
We concluded that indirect effects of the top predator destabilized the coexistence of competing prey species. In the landscapes of fear created by curly-tailed lizards, the clear niche partitioning exhibited by brown and green anoles on predator-free islands — a product of adaptive radiation — was no longer evident. Instead, these species were trapped together in the top story of the small islands, competing for the same space and food, afraid of getting burned by the hungry predators on the ground. Green anoles, despite being better adapted to arboreal life, got the shorter end of the stick (both literally and figuratively). This might be because brown anoles, as the incumbents on the islands, had greater strength in numbers. If we had introduced both brown and green anoles at identical starting numbers, would the green anoles have come out on top? Or, now that the combination of competition and predation has greatly diminished brown-anole populations, might green anoles stage a comeback? In 2018, we reintroduced green anoles on the two islands where they had been extirpated, with the hope of answering this question.
In any event, our findings ran counter to one of the motivating hypotheses of the project. Early studies, notably Bob Paine’s classic experiment in the rocky intertidal habitats of Makah Bay, suggested that predators tend to ameliorate competition between species at lower trophic levels by preventing any one species from becoming too abundant and excluding the others. Many ecologists, myself included, love this idea of ‘keystone predation’. Not only is it an elegant concept, but it also validates top predators as linchpins of ecological integrity. But when can we expect predators to play this role? In rocky intertidal communities, where keystone predation is a powerful force, sea stars feed on sessile invertebrates; but prey that are attached to the substrate have a limited ability to escape predators in space. In predator-prey interactions involving fast-moving prey that can rapidly adjust their behavior to avoid predators, I would expect keystone predation (sensu stricto, as opposed to the broader concept of ‘keystone species’) to be infrequent, and competition for enemy-free space to be both frequent and strong.
It has now been almost a decade from the conception to the publication of this work. What started out as a post-doc project has become an enduring annual ritual, and one that I have (usually) been able to enjoy thanks to a talented group of collaborator-friends: Palmer, Barrett, Kartzinel, and Xu, along with Tim Thurman, Kena Fox-Dobbs, Matt Hutchinson, Tyler Coverdale, Josh Daskin, Dominic Evangelista, Kiyoko Gotanda, Naomi Man in ‘t Veld, Hanna Wegener, and Jason Kolbe — and, of course, Schoener, Spiller, and Losos.
The interdisciplinarity of this team enabled what is to me the most satisfying feature of our work. We were fortunate to have access to a replicated set of small cays on which to manipulate species composition. That is a rare opportunity and would have made for a nice study in itself. But by also integrating molecular assays (to quantify diet composition and intraguild predation) and stable-isotope analyses (to quantify trophic position and food-chain length), we were able to gain deeper insight into the mechanisms underlying the population dynamics. Indeed, without these additional assays, our suppositions about the relative importance of consumptive and non-consumptive effects would have been equivocal at best. Molecular techniques have fully entered the mainstream of ecology over the past decade, yet they are still rarely paired with the kind of manipulative field experiments that so inspired me as a first-year graduate student. The fusion of sound natural history, rigorous experimentation, and forensic mechanistic exploration offers tremendous power to resolve the kind of messy complexity that has long frustrated ecologists.
New paper in Nature examines the interaction between green anoles and brown anoles, and how the presence of the predatory curly-tailed lizard changes the balance. See also the commentary by Os Schmitz.
Green anoles were trained, marked, released, and tracked in New Orleans. Photo by Jerry Husak.
In the US, we spend a lot of money trying to stay fit. This isn’t necessarily a bad thing, since there is a major problem with obesity and type II diabetes in the country. In humans, investment in increased performance abilities via the exercise response is also associated with numerous health benefits, such as decreased incidences of metabolic syndrome, cardiovascular disease, obesity, and diabetes, and aerobic capacity is considered to be an important predictor of longevity. However, it is these “side effects” that make exercise so interesting to an evolutionary biologist, because those wide-ranging, multi-system responses can tell us something about the evolution of animal life histories.
Superior locomotor performance has been shown to be advantageous to a variety of organisms in terms of male combat success, survival, and fitness. In addition, one of the most striking aspects of exercise physiology is how similar the response to exercise is across vertebrate animals, suggesting that the response to exercise is both ancient (yes, even fish respond to exercise!) and adaptive. However, until now, no studies have tested whether non-human animals that invest in increased athletic performance through exercise realize a fitness advantage in nature.
Jerry Husak and Simon Lailvaux set out to test whether superior performance after exercise training would increase survival probability in green anole lizards. Previous work with green anoles showed that they respond to different forms of exercise training, and that enhanced performance results in tradeoffs in other systems, such as reproduction and immnuocompetence. Why? Because performance abilities are energetically expensive to build, maintain, and use.
Urban islands in New Orleans where the study was conducted. Photo by Jerry Husak.
Jerry and Simon conducted their study in a New Orleans urban park that they cleared of existing lizards. They trained 30 lizards (15 male, 15 female) for endurance on a treadmill, 30 lizards for sprinting with weights on a racetrack, and had 30 untrained controls. All were released into isolated, urban islands in New Orleans, LA, USA and monitored for survival over an active season, over winter, and through the next active season. They predicted that training would enhance survival during the active season, but that the associated maintenance costs of training would decrease survival overwinter compared to controls.
This male made it a year in the wilds of New Orleans, but it looks like it was a rough year. Photo by Jerry Husak.
Contrary to expectations, they found that sedentary controls realized a significant survivorship advantage over all time periods compared to trained lizards. Trained lizards had reduced immune systems and lower fat stores, suggesting that in an environment with limited resources, it does not pay to exercise too much. These results suggest that locomotor capacity is currently optimized to maximize survival in green anoles, and that forcing additional investment in performance moves them into a suboptimal phenotypic space relative to their current environmental demands. We as humans can get away with it because we are not food limited. On the other hand, this is why doctors suggest consultation before going on a diet and doing intensive exercise training.
“I got an idea and I can’t get rid of it. I go to sleep and it comes right back at me. Never had anything give me so much trouble. It’s kind of a big idea. Maybe it’s full of holes.” – Adam Trask in East of Eden.
John Steinbeck (1962): The year he won the Nobel Prize in Literature.
So, what was Adam Trask’s big idea in John Steinbeck’s magnum opus “East of Eden”? And, more importantly, how does it relate to anoles? The kernel of an idea that would eventually revolutionize the salad industry—and link anoles to a literary legend—can be found in the fictional dialogue written by Steinbeck in 1952.
“… they’ve dug up a mastodon in Siberia. Been in the ice thousands of years. And the meat’s still good.” said Adam Trask.
“Mastodon?” inquired Will Hamilton.
“Yes, a kind of elephant that hasn’t lived on the earth for a long time.”
“Meat was still fresh?” asked Will.
“Sweet as a porkchop” proclaimed Adam.
Steinbeck was born in the Salinas Valley of Central California, known as “America’s Salad Bowl” for its prodigious production of leafy greens. He spent many summers, while away from college at Stanford, working in the vegetable fields near Salinas. Steinbeck’s fondness for his birthplace and working knowledge of the agriculture industry is a cornerstone to many of his novels, especially “East of Eden.”
“… in the cold parts of the country, don’t you think people get to wanting perishable things in the winter—like peas and lettuce and cauliflower? In a big parts of the country they don’t have those things for months and months. And right here in the Salinas Valley we can raise them all the year around.” declared Adam.
“Right here isn’t right there,” said Will. “What’s your idea?”
“… if you chop ice fine and lay a head of lettuce in it and wrap it in waxed paper, in will keep three weeks and come out fresh and good.” said Adam.
“Go on,” said Will cautiously.
“Well, you know the railroads … they’re pretty good. Do you know we could ship lettuce right to the east coast in the middle of winter?”
The perennial availability of perishable vegetables in the United States is now commonplace, but in the early 1900s, it made literary characters like Will Hamilton exclaim to Adam Trask to “let your damned idea die.” In fact, America’s most popular lettuce variety (iceberg) was originally called crisphead, until Salinas Valley growers began packing it with crushed ice and shipping it nationwide. The genesis of Adam Trask’s business plan was obviously fictional, but the idea of shipping lettuce with ice was successful and revolutionary in the early 1900’s; however, the method never quite kept vegetables fresh for long enough.
“What arrived in New York was six carloads of horrible slop with a sizable charge just to get rid of it.” – East of Eden by John Steinbeck.
In the pursuit of profitable ways to ensure lettuce does not turn into “horrible slop,” the next advance in production came from the humble bag. Lettuce can last for days on ice, but a bagged salad can last for a couple of weeks. It’s always difficult to establish the original (or best) anything in the food industry (vis-à-vis famous rivalries such as Pat’s versus Geno’s for cheesesteaks or Pepe’s versus Sally’s for pizza), but the late 1980s in the Salinas Valley is believed to be when and where the first bagged salads were packaged, distributed from, and then sold nationwide. The bagged salad turned a commodity crop whose predictability was in the capricious hands of nature into a consumer good as constant on the shelves of stores as shampoo or Twinkies.
Over the next decades, prepackaged leafy green vegetables boomed. To keep up with demand, growers invented creative ways to automate aspects of the production process, such as mechanically harvesting leafy greens. They also ramped-up the speed across the entire supply chain, such that lettuce could be bagged in the field within minutes of harvest and then sent overnight to supermarkets nationwide. These overlapping vignettes of industrial prepackaged salads provide the backdrop for a distinctly modern human-wildlife interaction: Small wild animals found by customers in prepackaged produce.
In our recent paper, we attempted to shed light on this poorly understood phenomenon by surveying online news articles for reported incidents. In doing so, we found that this is a much more common occurrence than one might think and that incidents encompassed representatives of several vertebrate groups. Most incidents involved amphibians (treefrogs and toads), and then reptiles (lizards and snakes), mammals (rodents), and birds. Anoles were the most common lizard that we could identify from the pictures and descriptions provided in the reports. The anole incidents included Green Fruit Loop, the aptly named Green Anole that became a class pet at Riverside Elementary in New Jersey. We suggested that the likely source of Green Anoles among the incidents was Florida because not only is the species is common there, but by 2012 the state was the third largest producer of leafy green vegetables in the United States, behind only California and Arizona.
Figure 1 from Hughes et al. (2019): Taxonomic and temporal breakdown for 40 incidents of extemporaneous wild animals found by customers in prepackaged produce items purchased in the United States. A) Vertebrate diversity among incidents; B) Annual distribution of incidents; and C) Monthly distribution of incidents.
An interesting social element emerged from my deep-dive into the trenches of the internet. I found that these incidents were shrouded in uncertainty and thus reporters often relied on anecdotes to discuss and describe them. One common urban myth was that these incidents almost never happen and the second was that if they happen, then it was because the produce was organic. In contrast to these popular views, we found that at least 40 incidents were reported since 2003—so, not exactly rare—and that less than 30% of incidents involved organic produce—most actually came from conventionally grown crops. For greater context and more details, see the Discussion of our paper where we address: 1) why these unfounded views may have persisted; 2) spatial, taxonomic, and seasonal patterns to our findings; 3) our results in the context of competing demands imposed upon the produce industry; and 4) the biosecurity concerns relating to the unintentional translocation of wild amphibians.
Modern agriculture has taken significant steps towards industrialization since the time that John Steinbeck penned Adam Task’s revolutionary idea (see Epilogue). Industrialization of food production will help to address the problems associated with feeding 9 billion people, a figure that is projected for the human population by 2050. Wild vertebrates in prepackaged produce, however, may be one symptom of an overburdened and overstretched produce production system. Any solution to this problem will not likely come from greater controls for wildlife, such as the currently employed “scorched earth” approach, but rather from the decentralization of agriculture. We suggest that the best approach would be to first invest in research aimed at studying a wide segment of biodiversity near agricultural lands, which will help growers assess potential intrusion risks of more species, and second to adopt quality control methods that account for a greater diversity of wildlife to improve screening at more stages in the produce supply chain.
Epilogue:
The birth of Adam Trask’s plan was fictional, but the growth of that idea, as depicted in the novel, is a great example of John Steinbeck’s (often overlooked) scientific mind. While many people my age read “Of Mice and Men” in high school and got to know Steinbeck the literary genius, they may not know Steinbeck the scientist. Ed Ricketts was a marine biologist that became a lifelong friend to Steinbeck when he moved to Monterey in the 1930s. The relationship between the writer and the scientist was one of mutual respect and admiration. At one point, they even undertook a six-week specimen-collecting expedition to the Gulf of California, which resulted in two published books. Not only was Ricketts the basis for Steinbeck’s character “Doc” in several novels (e.g., “Cannery Row”), but the influence he had on Steinbeck is unmistakable in many of his other works, including “East of Eden.” Adam Trask, for example, spawned his idea for preserving lettuce with ice from a scientific expedition that found a frozen mastodon in Siberia, and he read about this finding, refrigeration science, and bacterial growth in articles from “Atlantic Monthly,” “National Geographic,” and “Scientific American.” The mentioning of these specific journal titles in “East of Eden” was by no coincidence as they would have been the same ones that Steinbeck saw, and likely read, in Rickett’s lab, a place that he visited frequently. At the time of Ricketts death in 1948 (which sent Steinbeck into a depression), the two were planning another collecting expedition to British Columbia and another book.