Bumble and Bumble: what’s black and yellow and maybe more than one species?

During the dark afternoons of December in New England, I like to scroll through my old field photos and think of all the green, growing things I’ve measured in beautiful places during those long-ago long-lit seasons. Yesterday I flipped through a couple field photos from a friend — “Photos of younger Jon! :)” he wrote in the email — and the same sunny feelings flooded in.

As a master’s student*, Dr. Jon Koch and his insect net chased bumble bees all over the western United States. He was studying bumble bee decline, but hit weird hurdle: a messy species boundary between two bumble bees. Taxonomists and field guides were torn on whether Bombus fervidus was or was not Bombus californicus. These two “species” in the Bombus fervidus species complex were nearly morphologically identical, except for their color patterns: B. fervidus is noted as usually mostly yellow with a little black, while B. californicus sports mostly black with some yellow in variable detail. They were maybe different species, maybe hybridizing, or maybe the same thing with different color morphs. As Jon explained to me, “If we don’t know what the species are, how will we manage them? Bumble bees are differentially sensitive to land use change, disease, etc. The bumble bees in the Bombus fervidus species complex are found to be impacted by one disease, Nosema bombi, but perhaps differently. Therefore, it is important to recognize what the species boundaries are because estimates of infection prevalence might be not be done correctly due to the inability to tell the species apart.” 

Jon wanted to bring some clarity to the species complex by providing some new molecular evidence with broader taxa sampling. His new PLoS ONE paper, “Phylogeny and population genetic analyses reveals cryptic speciation in the Bombus fervidus species complex (Hymenoptera: Apidae)” delivers on the broader taxa sampling — 320 specimens from 53 sites — but the clarity is a bit of a cliff hanger. During the fieldwork, Jon and his coauthors keyed out identifications for their bees based on the setal color, and also took a tarsal clipping from the mid-leg for DNA extraction and microsatellite genotyping. When they compared field identification to the genotypes, they had an ID rate of just under 94%. Jon and I agree that that’s a pretty good record for fieldwork with cryptic species** but he adds, “it’s also cool to think that 6.2% of the time we were wrong! These bees are great at fooling us.” 

The bees that were fooling Jon were B. fervidus dressed as B californicus and vice versa. In Pinnacles and Yosemite National Parks there were ten mostly black bees (the typical B. californicus look) that turned out to belong to the genetic cluster that usually wears mostly yellow. The rest of the bees with black setal coloration belonged to another clade based on genotype, though this clade also included some bees in yellow. I asked Jon, “What is going on with the bee costume parties in Pinnacles and Yosemite?” His wild speculation is that little black dress is the dominant phenotype for bees in these parks, and the typically-yellow-genotype wears black here because everyone else is doing it: “bumble bees are notorious for converging on a local phenotype, which can even make it very hard to tell distantly related species apart.” However, in the sites where both genetic clusters of the B. fervidus species complex overlap, they usually do not look alike, so they aren’t mimicking each other. 

Ultimately, Jon’s team determined that the species complex comprised two lineages, but that both lineages exhibit the yellow and black phenotypes depending on geography. So while the B. fervidus species complex is not a single species, B. fervidus and B. californicus are not NOT conspecifics. Jon explains, “those names [B. fervidus and B. californicus] might not even be valid! The holotype of B. californicus happens to be where the genotype assigned to the “B. fervidus” was collected in the Sierra Nevada.” In short, the original bee that taxonomists knew as B. californicus may actually be genetically on the B. fervidus side of the lineage, and eventually one or both names might need to be thrown out.

This “it’s complicated” conclusion might be depressing news for someone who dedicated so much time and energy towards disentangling the species complex, but Jon closes his email to me with a happy emoji “nature has so many surprises, and science is an ongoing process :)” In the meantime, this paper points out that even if we don’t have the right names in place, we know enough to recommend that managers use Jon’s non-lethal method of clipping a bit of mid-leg for genotyping, and monitor the two clades of the B. fervidus complex separately. This is a great reminder for all of us in conservation research: we need to keep the ongoing process in perspective, while also delivering our findings, however not-quite-as-clear-as-we-hoped or maybe-unnamed as they may be, to our partners in management and policy. 

References:Koch JB, Rodriguez J, Pitts JP, Strange JP (2018) Phylogeny and population genetic analyses reveals cryptic speciation in the Bombus fervidus species complex (Hymenoptera: Apidae). PLoS ONE 13(11): e0207080. 

*Now, old Jon and old Caitlin are David H. Smith postdoctoral fellows together :)

**see McDonough MacKenzie et al. 2017 — When I was a master’s student working with volunteer-collected data I would have killed for a 93.8% identification rate. One my species, Labrador tea, was correctly identified 27.3% of the time. This is not a cryptic species; it doesn't sometimes dress up as Diapensia. 

Reading, Walking, Wishing

June in New England is a long stretch of long-lit days. When I was a PhD student, my Junes were the peak of my field season and I spent the long days logging miles up and down Cadillac, Sargent, and Pemetic mountains. For four years, my Junes were hiking ridges, recording data, wearing holes in the toes of my trailrunners. Now, I’m revising the papers that were written on the heels of those leg muscles and it’s weird to be indoors in June, sitting at a computer, without the tight hamstrings or blackfly bites.

After a long slog through a cold spring, this June I’ve returned to reading, picking up #365papers again in earnest after slacking off on the literature for a few months. Last week, I read Liam Heneghan’s essay “Have Ecologists Lost Their Senses? Walking and Reflection as Ecological Method” in Trends in Ecology & Evolution. I was indoors, at my desk, with the AC whirring, reading about walking. I felt like a fish out of water, or more aptly a field ecologist out of nature. In the essay, Henegham makes the distinction between ecologists and naturalists, comparing word counts in the anthologies The Essential Naturalist: Timeless Readings in Natural History (2011) and Foundations of Ecology: Classic Papers with Commentaries (2012).

“Although the two disciplines ‘observe’ and ‘see’ things in equal measure, natural historians nonetheless report engaging all of their senses in the pursuit of observations of nature to a greater degree. Natural historians report touching, feeling, hearing, and smelling the things of the world to an extent that scientific ecologists do not. Indeed, ecologists, if this small sample is representative, have abandoned smelling in its entirety. Moreover, natural historians ‘walk’, ‘roam’, ‘climb’, ‘sniff’, and ‘listen’ to a degree their ecological colleagues do not.”

I am a roaming, climbing, sniffing ecologist. But I bristled at the thought that ecologists as a whole should be compelled to walk to prove some kind of connection to the true core of the discipline. Heneghan does not outrightly demand that all ecologists walk, roam, and climb — his main argument seems to be the gentle conjecture “ecologists may have overlooked the fact that scrutinizing nature can benefit from an engagement of all the senses” — but he doesn’t leave much space within the discipline for non-field ecologists.

Perhaps Heneghan’s essay title is misleading and he isn’t worried about all ecologists losing their senses, just the outdoor ones. The field-based, nose-to-the-ground, perambulatory science that Heneghan and I practice is clearly not universal to ecology — and it shouldn’t be! We need modelers and theorists and lab scientists! But I fell for this essay hard. I am the target audience. When I started as a master’s student at the University of Vermont’s Field Naturalist and Ecological Planning program, my Botany 311 class, the Fall Field Practicum series of weekly full-day field trips, listed 7 goals on the syllabus. Goal #7: “Visit bakeries and enjoy spending the day outdoors.” In Heneghan’s analysis of word counts in the Ecology vs. Natural History texts, “Breakfast” receives 0.72 mentions per page in The Essential Naturalist; it does not appear at all in Foundations in Ecology*. Just digging out my Fall Field Practicum syllabus conjured up memories of cider donuts and eskers, travel mugs of maple-syrup-sweetened coffee and ombrotrophic bogs. My UVM experience was steeped in the kind of sensory details that Heneghan would appreciate and savor.

‘Walking and Reflection as Ecological Method’ reminded me of a similar paper I’d read in another (sadly non-bakery-centered) UVM class: Craig Loehle’s 1990 ‘A guide to increased creativity in research — inspiration or perspiration?’ Loehle also identifies the benefits of walking as a part of the scientific process when he encourages students to “get bored” as a work habit. This is recommended alongside running, procrastinating, and surfing — allegories for carving out time to think deeply and engage in non-productive, non-routine activities. These pursuits, Loehle promises, will facilitate creative problem solving. When I went back to re-read Loehle this week, I was surprised to find the advice “Don’t read the literature” under his list of methods for releasing creativity. I am, traditionally, a big fan of reading the literature. I’m a reader: when I was asked to review a Tansley Insight manuscript for The New Phytologist, my first move was to download and read the 2015 editorial “Introducing Tansley Insights – short and timely, focussed reviews within the plant sciences.” I won’t admit how many other Tansley Insights I downloaded after. A lot, okay? Maybe all of them. But Loehle’s “Don’t read the literature” is not a blanket statement; he clarifies that the first step as a scientist begins mulling over a new idea should not be to run to Web of Science (or whatever researchers used to find papers back in the dark ages of 1990), but to work through it a bit on your own.

“[Reading the literature] channels your thoughts too much into well-worn grooves. Second, a germ of an idea can easily seem insignificant in comparison to finished studies. Third, the sheer volume of material to read may intimidate you to abandoning any work in a new area.”

I agree with Loehle on all three points, but I’d add that the habit of reading broadly in the literature — taking recommendations from twitter**, searching outside of the Table of Contents of your subdiscipline’s favorite journal, checking out how your pet methodology is applied in another country or ecosystem, or seeking out papers with your field site as a keyword by researchers who are not in your field — is a kind of antidote to the well-worn grooves.

This month I read papers from Agricultural and Forest Meteorology, Alpine Botany, Bioscience, Conservation Biology, Current Biology, Ecology, Ecosphere, Frontiers in Ecology and the Environment, Integrative and Comparative Biology, Journal of Applied Ecology, Journal of Geophysical Research: Biogeosciences, Nature Geoscience, New Phytologist, Ocean & Coastal Management, Palynology, Proceedings of the National Academy of Sciences, and Trends in Ecology & Evolution. I am a broadly trained field ecologist — thanks UVM! — but as my career has progressed I’ve naturally found myself engaged in narrower research pursuits, and reading broadly keeps me centered, provides context for the tedium of slicing a 4.09 m core of lake sediment into half centimeter subsamples, and makes my work feel connected to society, policy, and big-picture conservation.

I’ll likely never publish in Ocean & Coastal Management, but reading “‘Back off, man, I’m a scientist!’ When marine conservation science meets policy”*** resonated with my own experiences writing public comments and meeting with congressional staffers. In a way, reading broadly is a kind of indoor-walking for restless ecologists who are prone to wandering.

Loehle and Heneghan’s essays are endlessly quotable for natural history students. But while they strive to expand how scientists engage in the world — Shake off your routine! Get outside! Smell! — they present an ironically narrow picture of role models. The patron saints of creative, roaming researchers, name-checked by both Loehle and Heneghan, are Darwin and MacAthur. I feel very strongly that if your argument around what’s needed in the “culture of ecology” can be reduced to “be more like this white man who had the privilege to travel freely and comfortably in the outdoors” you are fundamentally wrong. In Heneghan’s case, in 2018, there’s no excuse for whitewashing field ecology. Priya Shukla’s amazing piece in Bay Nature Magazine beautifully lays out the importance of representation in contemporary ecology, and the urgent need to uncover and share the ways in which wild landscapes are not empty areas that blankly awaited manifest destiny and reflect only Anglo-European stories. She writes “We need an act of revisionist natural history to color in the environmental and conservation movements. We should remind every hiker, biker, birder, citizen scientist, and field researcher that innumerable diverse people have shaped our natural spaces.” In a series of profiles of diverse voices in outdoor recreation, James Edward Mills writes in Outside, “Organizations like Outdoor Afro, Latino Outdoors, and Out There Adventures have begun stripping away the presumption of a white, male, heterosexual experience. Even more importantly, by unapologetically presenting their unique points of view, they’ve shined a light on a rich heritage of adventure and environmental stewardship that has been there for generations.”

This diversity exists in field ecology and natural history writing too, and it is not hard to find. Sure, Darwin and MacArthur were great at walking and writing about walking with wonderful sensory detail — but have you read J. Drew Lanham’s essay ‘Birding While Black’ or his book The Home Place? Robin Wall Kimmerer’s Braiding Sweetgrass? Janisse Ray’s Ecology of a Cracker Childhood****? Hope Jahren’s Lab Girl —in which the titular "girl" (Jahren) spends long stretches outside of the lab writing lyrically about working in the outdoors?

Heneghan begins his essay in a bog, but his call to arms (hiking boots?) is not simply an #OptOutside manifesto. He follows his walking naturalists — his long list of old white men: Irish botanist Robert Lloyd Praeger, Henry David Thoreau, Charles Darwin, Robert McArthur, and E. O. Wilson — indoors to their writing desks. At the end of the piece, Heneghan is in the archives, reading Praeger’s papers and reflecting on his prodigious writing. “A day’s walk can furnish long hours back at the desk.” Heneghan muses, “Thus for every insight into nature, there is a hidden process by which that insight was achieved; every active life contains a hidden core of repose.”

So this is my indoor June, my hidden core of repose. My trailrunners lie neglected, but the writing & reading continues, as I adventure through the memories and field notes and spreadsheets on the heels of the illustrious white men, and the many, many equally bold, sure-footed, and thoughtful unnamed white women and people of color who have trod this path before me.

References:

Heneghan, L., 2018. Have Ecologists Lost Their Senses? Walking and Reflection as Ecological Method. Trends in Ecology & Evolution 1–4. doi:10.1016/j.tree.2018.04.016 

Loehle, Craig. 1990. "A guide to increased creativity in research: inspiration or perspiration?." Bioscience 40.2: 123-129.  

*I have a confession to make here. I read most of Foundations in Ecology while I was a PhD student. I had not even heard of The Essential Naturalist until I read this paper. So maybe I’m not such a great naturalist after all? ...Or maybe I’m an amazing naturalist, always outside tromping around, and I don’t have time to read natural history anthologies because I’m too busy smelling nature?

**I found Heneghan’s essay by way of @ChelskiLittle’s prolific #365papers tweets. Thanks Chelsea!

***I found this paper by way of @Drew_Lab’s #365papers tweets. Thanks Josh!

****I cannot say enough about Milkweed Editions. This independent, nonprofit literary powerhouse in Minneapolis publishes incredible environmental writing. My husband gifted me a Milkweed book subscription years ago and it's my absolute favorite piece of mail every month. Maybe 30% of my love for LacCore & the science they do there is a side effect of the fact that every time I visit LacCore, I get to take a side trip to Milkweed. 

Biodiversity Patterns in Melanesian Coral Reef Fish: New Research with Old Naturalists

Old naturalists are my jam. I dedicated my PhD dissertation to a 19th century botanist who had spent her childhood following Thoreau around the Concord woods. I have a soft spot for research that draws on the work of older ecologists, for data that was handwritten before the advent of ballpoint pens, for 21st century papers based on museum natural history collections. This nostalgia is well-timed: museum collections are increasingly digitized and freely available online, and the Biodiversity Heritage Library is doing the same for scientific literature on biodiversity.

Just as my kind of fieldwork no longer requires taking the steamship to downeast Maine and a buckboard on wild roads between logging communities, my scholarship is not dependent on scouring the library stacks for a particular volume or traveling to the archives of a natural history collection to comb through specimens for just the right sample. In the 21st century it is significantly easier to be an armchair laptop historical ecologist. Easier, but not easy.

“Natural history and collections seem to be a bit of a hard sell when it comes to the ecological literature, which surprised me,” says Dr. Kathryn L. Amatangelo. She and Dr. Joshua Drew just published a PLOS ONE paper using coral reef fish data from museum collections records, peer reviewed literature including fish check lists, and biological inventories. The biodiversity pattern they were attempting to analyze and understand — that reef fish diversity in the Indo-West Pacific decreases along a longitudinal gradient from species-rich Papua New Guinea to species-poor American Samoa — was described in 1906.

Amatangelo laments, “It seems almost passé to look at old collections and think about how and why long-dead historians collected their data. When you try to combine that with statistics and scientific analyses people seem to get a little squirrely.”

Drew and Amatangelo’s paper “Community Assembly of Coral Reef Fishes Along the Melanesian Biodiversity Gradient” applies modern ecological theory and big data statistical tools to observations recorded by David Starr Jordan, a Victorian-era ichthyologist who was both the founding president of Stanford University and a suspect in the possible murder of Jane Stanford. If that legacy is not problematic enough, he was also into eugenics.

Thanks to the efforts of Biodiversity Heritage Library (BHL), we can read Jordan’s 1906 paper “On a Collection of Fishes from Fiji” where he notes the diminishing diversity of fish as you travel across Melanesia. Drew remarks, “historical ecologists are always looking for old species lists, and it was super cool to find that he worked in my study system in Fiji.” Drew describes a Jordan as “an ichthyological hero of mine, a complex and not unproblematic figure”: Jordan’s writing on ichthyological biogeography and community change, his system for organizing ichthyological collections and his service on the US Fish Commission, a precursor of NOAA, provide a foundation for the kind of work that Drew and Amatangelo so beautifully execute here.

In the pursuit of quantitatively describing this biodiversity gradient, Drew and Amatangelo compiled presence/absence records for 396 fish species in five taxa across 7 countries. As Drew describes it, this dataset was created from “a massive literature search from collections-based and peer-review based lists that were then double-checked with FishBase.” They looked for agreement across all three datasets (collections, literature, and FishBase), which gave them more confidence in the data since it was not susceptible to the biases present in only one dataset. Amatangelo is a community ecologist with a plant background, she partnered with Josh Drew through a twitter connection, bringing statistical savvy to these new-to-her taxa and ecosystems. I asked her what it was like to work with unfamiliar study species in this project. “One downside was that things that were intuitive to Josh, such as why some traits are important, was a bit of a mystery to me. That could also be considered a positive, though, because it meant that Josh had to be able to explain WHY they were important, which helped in writing the paper.”

The paper’s ultimate goal was to illuminate the processes behind the reef fish biodiversity pattern to inform conservation efforts. Drew acknowledges that their conclusions are not ground-shattering — the biodiversity gradient was described 110 years ago, and likely broadly known before then in local communities. “But it’s nice to put a p-value on it,” he says. “Natural history and traditional ecological knowledge are not always recognized because they don’t come with a p-value, so here we did that. We probably could have told you the same result before, but this adds weight to the management recommendations.” Those management recommendations include collaborations across Melanesia to more efficiently share resources and partition the region into functional biodiversity groups.

Through the power of twitter, digitization, and online collections two modern ecologists were able to build on a paper from 1906 and study Melanesian coral reef fish diversity from their laptop screens in the United States. So much of this data would be instantly recognizable to Jordan, but so little of the actual process of collaborating, compiling and analyzing data, and writing a paper has remained constant since 1906.

Drew reflects on this revolution in his recent correspondence to Nature Ecology and Evolution: “Digitization of museum collections holds the potential to enhance researcher diversity.” He and coauthors write that “the advent of digitization (open access to images and specimen data) now makes a wealth of biodiversity information broadly available…Digitization allows access to museum holdings to those for whom collections have typically been out of reach.” The concentration of collections in the Global North is a reflection of our discipline’s role in the history of exploration and colonialism. Untangling this broader context of past research is perhaps the most impressive, thoughtful work that a historical ecologist could pursue.

In two papers this fall Drew has managed to both uphold the ichthyological legacy of Jordan, and articulately argue that the museum collections Jordan once organized in his spare time from being abhorrently racist, could be, in digital form, a force for increasing diversity in science. 

References:

Drew, Joshua A., and Kathryn L. Amatangelo. "Community assembly of coral reef fishes along the Melanesian biodiversity gradient." PloS one 12, no. 10 (2017): e0186123.

Drew, Joshua A., Corrie S. Moreau, and Melanie L. J. Stiassny. "Digitization of museum collections holds the potential to enhance researcher diversity." Nature Ecology & Evolution (2017):10.1038/s41559-017-0401-6

Leveraging the Power of Biodiversity Specimen Data for Ecological Research

A guest post from PLOS Ecology Reporting Fellow, Caitlin McDonough, on research from the Ecological Society of America Scientific Meeting in Ft. Lauderdale, Florida, August 7-11, 2016.

Leveraging the Power of Biodiversity Specimen Data for Ecological Research at ESA 2016 While ecologists spend their graduate days troubleshooting code, writing manuscripts, and fighting with dataloggers, they often trace their roots back to a love of natural history--an acknowledgement of a childhood curiosity sparked by museums, camping trips, and backyard bug collections. This curiosity ties us ecologists to a long line of scientists, taxonomists, and collectors; we imagine that we could have sailed on the Beagle, or climbed Chimborazo, or that we would have happily canoed the wild Allagash River to botanize with Kate Furbish. On Wednesday morning, a group of 21st century ecologists presented a modern twist on these natural history dreams, with research in collaboration with these taxonomists, botanists, and collectors of the past. (A video of the session will be posted here.) 

iDigBio (Integrated Digitized Biocollections) organized this session, which brought together a diverse array of ecologists who have leveraged the power of biodiversity specimen data to approach 21st century problems in taxonomy, conservation biology, and climate change research. Each project relied on some form of biodiversity specimen data — from herbarium specimens to insect collections to marine collections — for applications ranging from restoration ecology to unraveling cryptic speciation, or creating species distribution models to tracking patterns in phenology. Recent efforts to digitize biological specimen data have sparked a renaissance in their use — pressed plants and pinned bees that once sat neglected in a dusty corner are now accessible to researchers thousands of miles away. In many cases, the 19th century collectors would likely recognize these research goals as they too were interested in species distributions,  recorded phenological events, and made observations about interactions between herbivores and plants. But, Thoreau did not geotag his field notes, and Linneaus might be surprised to find his herbarium specimen available as a jpeg. The importance of making biodiversity specimen data digitally accessible was clear from the start of the session.

Pamela Soltis noted that there are over 1,600 natural history collections in the U.S. with somewhere between one and two billion specimens. But iDigBio estimates that only 10% of biodiversity specimens are digitized. Throughout the session, presenters noted both the benefits of accessing the digitized data and the challenges of working with taxa and trophic levels that were underrepresented in the digital specimen world. Katja Seltmann lamented the lack of digitized parisitoids collections, and called out a bias towards plants and pollinators. Joan Meiners, who uses digital natural history collection specimens to investigate native bee conservation, showed a graphic of the low proportion of digitized bee specimens at major U.S. insect collections. The next speaker, Francois Michonneau, topped both of their complaints with an example of a historic sea cucumber collection that had been preserved in pieces, the equivalent of an ornithologist placing a beak and talons in a glass bottle and calling it a bird collection.  It is clear that the biodiversity specimens that are digitized are inspiring new research. Emily Meineke shared the origin story of her herbaria research: her project began in her kitchen. While flipping through old specimen data online during a procrastination jag, she noticed herbivory damage captured in one of Linnaeus’ specimens. With a little more digging, she found evidence of herbivory in many specimens — leaf mines, chewing damage, and galls — as well as actual insects preserved in the old leaves. Another example of unintentional data captured in herbarium specimens is Amanda Gallinat’s fruit phenology study. She found over 3,000 specimens comprising 55 species in seven major New England herbaria that contained mature fruit pressed among the plant material. Just as Meineke realized that herbaria offer unprecedented opportunities to understand what factors drive herbivory rates across large spatial and temporal scales, Gallinat was able to assess patterns in fruiting across native and invasive species at a regional scale from the 19th century to the present. Meineke has begun surveying for herbivory damage in the Harvard University Herbarium collection, but she is also working to make this a citizen science project called Bite Marks in the Zooniverse. Soon everyone will have the opportunity to look at herbivory damage while procrastinating in their kitchens! 

In addition to the diverse research that has emerged from digitized biological specimens, this session provided some practical advice for all ecologists. Pamela Soltis presented Charlotte Germain-Aubrey’s project “Using museum data for species distribution modeling: The case of plants in Florida” and provided a thoughtful behind-the-scenes look at the building of a maximum entropy model. She deliberately explored the process behind decisions about climate data (e.g. average climate vs. climate data from the year of collection for each specimen), the area in which the model trains, smoothing response curves, and the number of background points. François Michonneau closed his talk with a great overview of his best practices for instituting data quality checks in R code workflow. While these skills are typically missing from our training, he stressed the importance of building a culture of documentation and replication, recommending courses from datacarpentry.org. Katelin Pearson showed that the collector community — a group that is regularly in the field, well-trained to recognize patterns and norms, and communicate with other experts — currently lacks the protocols and the semantics to document outliers in a consistent, meaningful way. This community has great potential to detect outliers in phenology, distribution, ecology, behavior, morphology, but at the present there is no direct feed between the collectors and ecologists who are tracking changes or outliers.

Finally, Libby Ellwood closed the session with an overview of iDigBio’s citizen science projects to engage the public in the work of digitizing the many, many biological specimens that are not yet a part of the digital record. 

Caitlin McDonough MacKenzie is a PhD candidate in the Primack Lab in the Biology Department at Boston University. She spends her field seasons in Acadia National Park, Maine studying leaf out and flowering phenology and patterns of historical species loss across plant communities. Her field methods include three ridge transects that are conveniently located adjacent to beautiful running trails and carriage roads. Away from Acadia’s granite ridges, she’s interested in underutilized sources of historical ecology data including herbarium specimens, field notebooks, photographs, and old floras; the potential for citizen science in phenology research; and the intersection of science and policy.  (Follow Caitlin on Twitter @CaitlinInMaine)