QuestionOverbrowsing by abundant deer reshapes the composition of forests from palatable native species to browse-tolerant species, leading to simplified understories with reduced biodiversity. Here, we ask whether geologic refugia (e.g., tall boulders) will provide a haven for vulnerable plant species to escape browsing pressure from high populations of native deer.LocationHuron Mountains, Michigan, USA.MethodsWe compared the percent cover, species richness, and Shannon diversity of all plant species on tall boulders to adjacent fenced exclosures and to forest floor reference plots to quantify the degree to which deer impact one of the largest remaining tracts of old-growth forests in the Eastern United States.ResultsWe found that boulders contained 60% higher richness and diversity than reference plots, as well as a more diverse mix of midstory species, shrubs, and forbs. Seven-year-old exclosure and reference plots were nearly indistinguishable from each other and were dominated by Acer saccharum and other browse-tolerant species.ConclusionsThis study demonstrates that boulders are a highly effective, though rarely used, proxy or bioassay of the degree to which chronic overbrowsing creates impoverished understories composed of browse-tolerant or unpalatable species. Our findings highlight the significance of refugia as a haven from overbrowsing for vulnerable plant species and emphasize the importance of using exclosures alongside refugia.

Throughout the Eastern Deciduous Forest Biome, overbrowsing by white-tailed deer (Odocoileus virginianus) has reduced plant diversity and impoverished forest regeneration. However, some geomorphic features may provide refuge from browsing. For example, browse-sensitive species are often abundant on the tops of tall boulders and treefall tip-up mounds. If refugia are common, they might serve as viable seed-source populations for recovery of vulnerable plant species that have been locally or regionally extirpated. Here, we test the hypothesis that cliff edges and steep slopes provide refuge for browse-sensitive woody species. We quantified stem density or individual patch size, or both, of four woody species at six steep exposures across the Hemlock-Northern Hardwoods Association. Level areas adjacent to steep exposures served as deer-accessible reference sites. The combined density of our focal species was 16 times higher on cliff edges and steep slopes versus reference areas, and combined mean patch size was nearly six times larger. We also conducted a remotely sensed spatial analysis of slopes between 35 degrees and 75 degrees throughout the State of Pennsylvania to quantify the abundance and spatial aggregation of these potential refugia. Steep slopes occurred in all 67 Pennsylvania counties, representing up to 3% of land surface in rugged counties. Steep slopes were also often clustered or large, suggesting locally high gene flow may maintain genetically diverse and vigorous seedsource populations of browse-sensitive species. Our study is the first to quantitatively demonstrate that cliff edges and steep slopes have high conservation value by serving as browsing refugia for browse-sensitive woody plants in temperate forests.

Lianas (woody vines) are an important component of tropical forests and research on the liana community of the Barro Colorado Nature Monument (BCNM) has helped to elucidate their diversity, natural history, ecology, and role in tropical forest processes. More than half of the canopy trees in old-growth forests, such as the one on Barro Colorado Island (BCI), host lianas in their crowns. Both observational and experi -mental studies have demonstrated that lianas reduce tree recruitment, growth, reproduction, and survival, resulting in reduced forest biomass uptake and lower carbon storage. Because lianas have a stronger negative effect on the performance of some tree species than on others, they may influence tree community composition. Lianas also appear to promote animal diversity by providing resources, nesting sites, and connectivity among canopy trees. Research on the lianas of the BCNM has fundamentally advanced our understanding of liana ecology and their role in tropical forests.

Sometimes referred to as the “great unseen,” microbes (e.g., bacteria, fungi, protists) are by far the most abundant organisms on Earth. Although soil microbial communities and their impacts on plant communities have received extensive attention, comparatively little is known about leaf microbes and their interactions with plants and higher trophic levels, particularly in forests. We began a long-term research effort at the Barro Colorado Nature Monument in 2010 to assess the diversity, distribution, and impacts of foliar microbiomes – specifically bacteria that occur upon and within leaves –on seedlings and saplings of co-occurring tree species. We demonstrated that experimentally altering foliar microbiomes with commercial antibiotics had pervasive effects on plant traits, plant-plant interactions, and interactions with higher trophic levels. Taken together, our results suggest that these cryptic organisms may drive important trophic cascades and ecosystem-level patterns, including plant diversity, nutrient cycling, and net primary productivity in forest biomes.

Disturbance regimes, like low-intensity fire, canopy gaps and ungulate browsing, play a critical role in determining ecological composition and structure in temperate forests around the world. Each disturbance (or lack thereof) can lead to unique plant communities, but we do not understand how combined disturbances change plant diversity and the resulting soil seed bank. Changes in the soil seed bank, which depend on the plants that survive post-disturbance, can then influence future biodiversity and succession. We used a long-term experiment in West Virginia, USA, that factorially manipulated low-intensity fire, deer exclusion and canopy gaps. Thirteen years after disturbance initiation, we sampled the seed bank from each disturbance treatment. We found that low-intensity fire led to increased seed bank density, with additional canopy gaps and deer exclusion each creating unique seed bank communities. Combined fire, canopy gaps and deer presence led to high seed bank diversity and the most unique seed communities, while canopy gaps and deer had no effect on seed banks unless the area was previously burned. In contrast, combined fire, canopy gaps and deer exclusion led to the lowest seed bank diversity of all treatments, reflecting the continued legacy of extant plants that grew immediately after disturbance. Seed communities were also distinct from extant understory species over 13 years, regardless of disturbance treatment. Each reintroduced disturbance combination left a unique legacy in the seed bank that will likely influence future forest reorganization following disturbances, adding to our understanding of how multiple disturbances influence forest succession and organization. Synthesis. Forest disturbance regimes have changed around the world and are being restored or manipulated to support biodiversity. Reintroduction of disturbance leads to unique plant communities, but we do not understand how combined disturbances change the soil seed bank. Using an experiment that manipulates low-intensity fire, canopy gaps and deer exclusion, we find that combinations of these experimental treatments leads to substantially different seed communities. These disturbance-altered seed banks will likely influence future biodiversity and successional patterns, highlighting how the restoration of disturbance can strongly and indirectly influence temperate forest community dynamics.

Among-individual variation in predator traits is ubiquitous in nature. However, the role of intraspecific trait variation in trophic dynamics has been seldom considered in community ecology. This has left unexplored a) to what degree does among-individual variation in predator traits regulate prey populations and b) to what degree do these effects vary spatially. We address these questions by examining how predator among-individual variation in functional traits shapes communities across habitats of varying structural complexity, in field conditions for the first time. We manipulated Chinese mantis (Tenodera sinensis) density and trait variability in experimental patches of old fields with varying habitat complexity and quantified the impacts on lower trophic levels, specifically prey and plant biomass. Our mantis groups thus contrasted in density (six or twelve individuals) and levels of variation in a key behavioral trait, activity level (movement on an open field). Our metric of habitat complexity was the density of plant material. In complex habitats and at high mantis densities, behaviorally variable groups decreased prey biomass by 35.1%, while at low densities, low levels of behavioral trait variability decreased arthropod biomass by 27.1%. Behavioral variability also changed prey community composition. Our results are among the first to demonstrate that among-individual trait variation can shape open species-rich prey communities. Further, these effects depend on both predator density and habitat complexity. Incorporating this important facet of ecological diversity revealed normally unnoticed effects of functional traits on the structure and function of food webs.

Natural disturbances can maintain forest diversity by creating a heterogeneous resource landscape, which can contribute to coexistence of early-and late-successional species. However, almost all forested ecosystems are also subject to multiple anthropogenic stressors, which could inhibit plant diversity and forest recovery after natural disturbances. Here, we present the first large-scale replicated field experiment testing the interactions among two natural and two anthropogenic disturbances in a mature secondary temperate forest. Specifically, we test the extent to which: 1. a natural windthrow (tornado), 2. deer browsing, 3. salvage logging, a common forestry practice to generate revenue after natural disturbances, and 4. mechanical removal of the understory, drive subsequent plant community assembly, diversity, and abundance. Our results have two clear messages. First, the combined disturbances, especially salvage logging, increased plant species richness and abundance; we found 65 more species in 0.1 ha of the disturbed areas versus the same area of reference forest. Second, we found that it was critical to analyze plant growth forms separately; each major plant growth form responded to the distur-bances differently. Surprisingly, salvage logging, browsing, and vegetation removal had negligible effects on the diversity and abundance of tree seedlings or saplings. In contrast, salvaging increased herb richness by 30% (six extra species per 36 m2 plot), and shrubs were twice as abundant in salvaged versus unsalvaged plots. The combined disturbances created unique plant communities in comparison to adjacent undisturbed reference sites, but plant growth forms differed in which combination of disturbances shifted community composition the most. We did not find that non-native invasive species substantially increased in abundance in response to the dis-turbances, nor were invasive species unique to the salvaged areas. Together, our results demonstrate that a patchwork of combined natural and anthropogenic disturbances can promote plant diversity in a human-modified temperate forest. In particular, the combination of a natural wind disturbance and salvage logging benefited native herbaceous species, the plant group that represents the vast majority of vascular plant species (similar to 80%) in temperate forests across the world.

Throughout eastern North America, chronic white-tailed deer overbrowsing and widespread exotic plant invasions degrade native plant diversity and abundance. However, because deer densities are high across entire regions, it is challenging to find large areas where deer have been at low density for long periods. We predicted native plant diversity and abundance would be significantly higher during gap-phase regeneration where deer have been maintained at low densities ( 7 deer/km²) compared to where deer have been chronically overabundant. Simultaneously, we predicted that exotic plant diversity and abundance would be lower in gaps where deer have been maintained at low versus high densities. We also evaluated whether deer density interacts with gap age, gap size, and gaps in dry versus more mesic forest patches. We studied 41 gaps throughout an 88 km2 fenced military post where deer were maintained near pre-European-settlement densities for 67 years and in nearby forests where deer have been overabundant for decades. Native species richness, diversity, and cover were 37–65

Intraspecific phytochemical variation across a landscape can cascade up trophic levels, potentially mediating the composition of entire insect communities. Surprisingly, we have little understanding of the processes that regulate and maintain phytochemical variation within species, likely because these processes are complex and operate simultaneously both temporally and spatially. To assess how phytochemistry varies within species, we tested the degree to which resource availability, contrasting soil type, and herbivory generate intraspecific chemical variation in growth and defense of the tropical shrub, Piper imperiale (Piperaceae). We quantified changes in both growth ( e.g ., nutritional protein, above- and below-ground biomass) and defense ( e.g ., imide chemicals) of individual plants using a well-replicated fully factorial shade-house experiment in Costa Rica. We found that plants grown in high light, nutrient- and richer old alluvial soil had increased biomass. High light was also important for increasin g foliar protein. Thus, investment into growth was determined by resource availability and soil composition. Surprisingly, we found that chemical defenses decreased in response to herbivory. We also found that changes in plant protein were more plastic compared to plant defense, indicating that constitutive defenses may be relatively fixed, and thus an adaptation to chronic herbivory that is common in tropical forests. We demonstrate that intraspecific phytochemical variation of P. imperiale is shaped by resource availability from light and soil type. Because environmental heterogeneity occurs over small spatial scales (tens of meters), herbivores may be faced with a complex phytochemical landscape that may regulate how much damage any individual plant sustains.

White-tailed deer (Odocoileus virginianus) have been overabundant in eastern North America for more than five decades, re-sulting in depauperate understories and ricocheting effects on higher trophic levels. Even after deer populations are reduced, understory plant communities may fail to recover for an unknown length of time due to persistent legacy effects. We surveyed understory plant communities in six deer exclosures and paired reference plots in northwestern Pennsylvania to determine the degree to which 19 years of deer exclusion was sufficient for recovery of species richness, diversity, percent cover, and un-derstory structural complexity. We observed a 2.3-fold increase in tree cover and a 60% reduction in fern cover in the ground layer, as well as a 114-fold increase in foliage density between 80 and 200 cm above ground level, in exclosures compared to reference plots. However, the exclosures did not permanently support higher overall percent cover, species richness, or di-versity in the ground layer, nor did we detect any meaningful divergence in community composition between exclosures and reference plots. We conclude that 19 years of release from chronic over-browsing are sufficient to restore understory structural complexity, but recovery of diversity in the ground layer will require more time or direct intervention.

Epiphytes are a unique group of plants that live nonparasitically on other plants ("hosts") and constitute approximately one-fifth of Neotropical vascular plant diversity. However, the processes governing early epiphyte community assembly are poorly understood and have scarcely been experimentally tested. Here, we use an in situ experiment in the cloud forest of Santa Fe, Panama, to evaluate the extent to which host substrate texture regulates early epiphyte establishment We experimentally varied the surface roughness of native wood substrates, applied bromeliad and orchid seeds to the substrates, and monitored emergence of epiphyte seedlings and their persistence for a year. Rougher substrates facilitated higher initial abundance of epiphyte seedlings; after two months, 81% of the 1,934 total germinated epiphytes occurred on the substrates with experimentally added roughness. Via photo analysis, we also show that epiphytes disproportionately established early on in the experimental grooves, wherein 71% more epiphytes per unit area occurred within 1.5 mm of the grooves than on nearby smooth surfaces. While epiphyte cohort survival rates differed between rough and smooth substrates in the first six months, more than 99% of all seedlings died after one year, regardless of experimental roughness treatment. Only 10 seedlings survived through the end of the experiment. Our results suggest that while substrate texture explains some variation in early epiphyte emergence, roughness alone is not sufficient to explain epiphyte persistence to adulthood. Moreover, our results highlight the importance of removal processes (e.g., wind, rain, animals) in structuring early epiphyte community assembly. Variation in substrate texture may contribute to differences in epiphyte diversity and community composition withinand among-host tree species, but more experiments are needed to disentangle removal processes from substrate-mediated host affinity.

Healthy forests are vital components of terrestrial ecosystems for their raw materials, high biodiversity, cycling of nutrients, and potential to sequester carbon. However, these ecosystems are sensitive to disturbances, and anthropogenic activities pose a serious threat to forest ecosystems globally. For example, human activities have dramatically altered multiple historical disturbance regimes in forests, including suppressing fire, increasing the density of large herbivores, and reducing the size of canopy gaps, among other disturbances. Such disturbances can have dramatic impacts on microbially-mediated forest soil functions, but more research is needed to determine the collective impacts of these disturbances. In this study, we investigated the interactive effects of disturbances, namely the legacies of fire, large herbivore densities, and canopy gap creation, in a deciduous forest soil. We determined that forest floor and mineral soil carbon and nitrogen pools were shaped by multiple disturbances, but fire was more influential than the other disturbances. The abundance of several functionally-relevant microbial taxa were significantly impacted by fire, and the effect was more pronounced in the mineral soil than in the forest floor. Together, these findings demonstrate that multiple disturbances, especially a legacy of fire, exerts long-term control over soil carbon, nitrogen and microbial dynamics in a deciduous forest system.

White-tailed deer (Odocoileus virginianus) overbrowsing over the past several decades has caused substantial changes to plant communities in eastern deciduous forests. Deer-preferred species have declined or become locally extirpated in many areas while deer-tolerant species have greatly increased in abundance. Moreover, the abundance of nonnative invasive species has also increased over the last few decades, outcompeting many native species. Native shrubs such as Sambucus canadensis (American elderberry) are now much less common, particularly in urban forests and forests with high deer densities. Here, we performed a short-term study in which we introduced propagations of a native shrub into a replicated field experiment in which we factorially manipulated deer presence or absence and invasive species presence or absence by using exclosures and removing invasive species. We monitored metrics of S. canadensis fitness including leaf count and plant height over two growing seasons. Browsing substantially reduced survival whereas invasive species provided a modest degree of associational resistance. Browsing significantly decreased both leaf count and plant height. By the end of the study, individuals within exclosures had over twice the number of leaves, were five times taller, and had much lower mortality versus those exposed to browsing. Removing invasive species did not change plant height, however, S. canadensis individuals experienced marginally higher leaf counts inside exclosures where we had removed invasive species. This trend was opposite in individuals outside of fences such that those growing within patches where invasive species remained had slightly higher leaf counts, especially during the first growing season. Until the final census date, elderberry growing within patches of invasive species had slightly lower mortality versus those growing in patches where invasive species had been removed. The results from our short-term study indicate that browsing is more inimical to a native shrub than competition from invasive species and we did not find evidence of associational resistance to browsing from invasive cover.

Chronically elevated ungulate browse pressure in temperate forests worldwide often generates ecological legacies characterized by low plant diversity and contributes to the formation of dense, nearly monodominant, and highly recalcitrant layers of understory vegetation. Once established, these recalcitrant layers combined with continued browsing may jointly constrain tree establishment and diversity so completely that understory recovery may be unattainable without mitigating browse pressure, the recalcitrant layer, or both. Here, we investigate the independent and synergistic effects of both white-tailed deer (Odocoileus virginianus) browsing and hay-scented fern (Dennstaedtia punctilobula) competition on tree regeneration in a 10-year experiment. Specifically, we examine how tree seedling establishment, growth, and composition are filtered by fern cover versus fern removal (gaps), browser presence versus absence (exclosures), and their combined effects during 10 years at three hardwood forest sites in Pennsylvania, USA. Fern gaps enhanced establishment for multiple tree species, increasing seedling density and diversity, particularly in the first 3 years post-treatment, and enhancing richness (≤1 species) over the course of the experiment. Excluding deer for a decade increased the height growth of other regeneration and altered species composition, but had no effect on diversity, richness, and density. Notably, we observed higher Prunus serotina seedling densities outside exclosures, possibly due to greater secondary dispersal. We argue that browsing legacies in second growth forests established at the turn of the last century created two conditions inimical to diverse forest regeneration: an overstory dominated by two species, P. serotina and Acer rubrum (86% of basal area), and a dense recalcitrant understory layer dominated by a native fern. The first condition limits propagule supply, the second strongly filters seedling establishment, and both create impoverished forest understories composed of few individuals and species. In undisturbed forest understories, the inertia toward impoverishment was sustained across the decade even where browsing was eliminated. Consequently, stand replacing disturbances (whether natural or anthropogenic) that disrupt the understory layer and reinitiate succession may be necessary to propel forests out of their current stasis and down a pathway leading to greater diversity.

We provide an introduction to the process of applying to graduate school in ecology and evolutionary biology with information on how to prepare for graduate school, how to choose a program, how to gain admission, and how to find and select an advisor. We provide a basic step-by-step guide for the application process and for the prelude to that process. This is a much revised and updated version (Carson 1999) containing additional information on parenting during graduate school and information for underrepresented groups in ecology and evolutionary biology and first-generation college students. We hope that this guide will help students, and others get started down the right track and help them to ask more refined questions of their mentors about the whole application process. This guide applies primarily to graduate programs in ecology, evolution, and behavior, as well as other areas of organismal biology, particularly those where the prospective student will be directly admitted into a specific professor’s laboratory. It may also be useful for those applying to graduate programs in other subfields of biology such as molecular, cellular, or developmental biology. In general, students should understand at the outset that applying to graduate school in these disciplines is much different than applying to undergraduate programs or applying to professional graduate degree programs such as medical or law school. Graduate schools will often use some combination of up to five primary criteria or metrics to evaluate applicants: these are grades, your scores on the Graduate Record Exam (GRE), your research experience, your letters of recommendations, and your personal statement or essay. We discuss each of these below. While it is never too early to start thinking about graduate school, before applying, you should be pretty confident that graduate school and research are right for you. It can be a long haul (typically 5–6 years for a Ph.D.), and a serious commitment is required for success. If you are not certain, or if you feel “burned out” from school, take a year or two off, gain some additional research experience through postbaccalaureate research programs, seasonal fieldwork, laboratory technician positions, or get a job and bank some money, and then carefully consider postgraduate education. Delaying entry into graduate school will rarely hurt your chances of admission later and will improve them if you garner relevant research or life experience. Perhaps more importantly, it may reinforce your decision that graduate school is the right place for you. The use of the GRE by universities is waning, at least partly because of the mounting evidence that the examination only weakly predicts student success (Moneta-Koehler et al. 2017, Peterson et al. 2018) and often fails to predict the success of students, particularly minority students (Miller et al. 2019). Whether the GRE is required can vary by departments even within institutions, check to be sure. Although your grade point average (GPA) and your GRE scores are not always good predictors of success in graduate school, universities will use these metrics as one way to compare and evaluate applicants. Here is some advice: Many graduate schools have a 3.0 as a standard cutoff and the most competitive programs will look for GPAs that are much higher. However, if you have a low GPA, do not lose hope because there are a few things you can do. For example, make it clear in your personal statement (see Applying) if your grades improved steadily over time. Some programs will emphasize your GPA in the last two years of your degree program, or within your major; if your GPA is higher in these respects, emphasize this in your application. Also, if you had a semester or year when your GPA plunged relative to other years, explain why this may have occurred in your personal statement. These reasons, however, should be compelling and may include illness, caregiving, holding down a part-time job, or participation in scholarship athletics, but only you can provide relevant context for your GPA to an admission’s committee. The GRE attempts to evaluate your quantitative, verbal, and analytical abilities. While the use of the GRE as a performance metric is declining, if you score well (~75 percentile or above), this can make up for a low GPA. Thus, even if an institution does not require GRE scores, if you do well, you can still pass on the scores when applying or contacting a prospective advisor (see Selecting a potential advisor). A high score on the GRE can make up for a low GPA (or vice versa). If you decide to take the GRE, you should consult one of the many available preparation and study guides. Educational Testing Services, which administers the GRE, has low-cost tools to help you, including two free practice examinations (https://www.ets.org/gre/revised_general/prepare/). Practice taking the test under the actual conditions of the examination until you feel comfortable with the format and length of the test. In addition, see if your undergraduate institution offers free help and instruction through the library or career services center. Students can also request disability or health-related accommodations from Educational Testing Services (https://www.ets.org/gre/revised_general/register/disabilities). Note that some universities may require that you also take the GRE Biology Subject Exam. Regardless of all of this, we predict, and welcome, a continued decline in the use of GRE scores as a metric to vet applicants. One way to avoid the stress and cost associated with taking the GRE is to avoid applying to those places that still require it. There are many outstanding master’s programs out there. Thus, if your grades or GRE scores or both are relatively low, but your ultimate goal is a Ph.D., do not despair. Consider trying to find a master’s program (see Should you do a master's degree first?) that emphasizes research, where your chance for admission might be higher. In a master’s program, you can conduct interesting research and produce publications in peer-reviewed journals derived from that research. Along with enthusiastic letters of recommendation, this effort can more than make up for a modest GPA. Know that classes are only one part of your education. You should begin to obtain real hands-on research experience as early as your first year as an undergraduate. Research is the most important thing you can do to prepare yourself for graduate school. This is because it will teach you not only how to conduct research, but also whether you like research, and if so, what areas of research you enjoy the most. You can gain research experience working with professors, graduate students, and with scientists at nearby natural history museums, zoos, or botanical gardens. The vast majority of scientists have personal web pages or information about their research programs on their departmental website. On these sites, you can learn much about the professor, their graduate students, their research, and their scientific publications. There are a variety of ways to gain research experience including volunteering to help, working as a paid assistant, signing up for research credit, conducting independent research, or doing an independent study (library project that will require reading in the primary journal literature). Graduate students almost always need help with their research, and they are often short of funds, so seek them out if you are willing to volunteer time; they will be grateful, and you will learn a lot. Unfortunately, many undergraduate research opportunities are unpaid, which is inequitable because students who cannot afford to work without pay will have a harder time gaining research experience than their peers who have the financial support to volunteer their time (Fournier and Bond 2015). Students who seek research experience but cannot afford to volunteer may be able to apply for federal work-study funds to work in research laboratories or look for funding through university undergraduate research offices. There are also opportunities for undergraduates at universities across the United States and at field stations that offer well-paid summer immersion programs in research. Many of these are funded by the U.S. National Science Foundation (NSF), but others are funded separately by the host institution. If your university has an Office of Undergraduate Research, they also likely offer grants or fellowships to promote research or have additional information on how to find and apply for paid undergraduate research opportunities. These opportunities are a terrific way to get paid to do research. If you are unsure about how to begin the process of selecting an undergraduate research mentor or applying to an undergraduate research program, McGill et al. (2021) have produced a useful guide. A current listing of NSF funded Research Experience for Undergraduates can be found at https://www.nsf.gov/crssprgm/reu/list_result.jsp?unitid=5047. There are other resources for finding opportunities to conduct research. Texas A&M has a fairly comprehensive Job Board in ecology and wildlife biology (https://wfscjobs.tamu.edu/job-board/) as does the Ecological Society of America (https://www.esa.org/nextgencareers/resources/job-sites). After gaining experience by one of the above means, try to attend a scientific conference or research fair, and if possible, give an oral presentation or poster on your research. Presentations are short talks (12–15 minutes) on your research, while a poster displays your research with text and figures. Conferences can also be a great place to meet and visit with potential advisors or graduate students at institutions where you are considering applying for graduate school. Conferences also frequently have professional development workshops and panels that you could attend to learn about future career options, how to develop a CV, and even how to apply to graduate school. However, to make the most of a conference, you may want to reach out to schedule times to meet with scientists prior to the start of the meeting. Most meetings publish their conference programs online where they list those who are presenting. Many universities have research fairs specifically designed to allow undergraduates to present their work, although graduate admission’s committees weigh regional or national meeting participation more highly than research fairs at your home institution. Larger national meetings occur in most years and are hosted by scientific organizations such as the Society for the Study of Evolution (SSE) and the Ecological Society of America (ESA). Ask graduate students and professors for advice on attending meetings and work with your research mentors to polish your presentation or poster. A growing number of scientific societies offer funding for undergraduates to attend meetings; these include ESA’s SEEDS program (Strategies for Education in Ecology, Diversity, and Sustainability) and funding offered by SSE (http://evolutionsociety.org/content/society-awards-and-prizes/travel-awards.html). These societies, and others, often have dedicated travel awards to promote participation by underrepresented groups. Sometimes universities or individual scientists will have funds to defray the cost of attending a meeting. See if your institution has an Office for Undergraduate Research, which may provide travel funds. Even if you do not have independent research to present, you should still try to attend a conference or research fair. These gatherings will give you a glimpse of the diverse array of research that is out there, give you a chance to meet prospective advisors, and probably convince you that you can do interesting science. This could result from your independent research, a project you began in a class, or an independent library project; it will require the help of a professor, postdoctoral researcher, or graduate student. Do not think that this is beyond your ability but know that it will require dedication and perseverance. It is essential to plan ahead, because getting the research to a publishable level will be more time-consuming than you expect. Nothing impresses a prospective advisor or graduate school like a publication in a refereed scientific journal! This will no doubt help you get into a top program or is an excellent way to get past a modest GPA. If you are working on a publication, but it is still in the works after your application was due, make this clear when you apply. Finally, some applicants have earned co-authorship on publications; this also demonstrates commitment and achievement. And of course, many applicants are not authors on papers, and this does not preclude you from gaining entry to an excellent graduate program. The research experience itself will have been essential to prepare you for graduate school. A mentor is someone you can talk to about careers in science, ask for guidance, request letters of recommendation, and someone you can keep in touch with throughout your journey in academics or subsequent career. The earlier you find a mentor or even multiple mentors the better; however, there is no deadline. If you are a first-generation student, this is a great way to start researching possible career pathways with someone who has already navigated them. Your mentor does not necessarily have to be a professor, but could also be a graduate student, a postdoctoral researcher, or someone from the advising office. The most important thing you want in a mentor is someone who you feel comfortable meeting with and talking about your professional goals. It is important to know that recommendations that only include your performance in class will be considerably less influential than recommendations that evaluate your performance outside of class conducting independent research or completing an independent study. Recommendations are extremely important. Your professors are likely to be friends or colleagues of the professors that you are applying to work with. Potential graduate advisors will often trust the recommendation of a close colleague or scientific peer more than a GPA or GRE score. Participation in departmental events will contribute to immersing you into the culture of science. Examples of departmental organizations include biology or ecology clubs. Consider taking on a leadership role (e.g., president) or starting a relevant science club if one does not exist; doing this can help you learn important leadership and organizational skills. Perhaps most importantly, if your department has a weekly seminar series or journal club or both, make time to attend at least some of these gatherings. A journal club is an informal weekly meeting of scientists to discuss recent scientific papers. Seminars are hour-long presentations on current research by scientists from both within the institution, and more often, by those outside the department or university. Undergraduates are almost always welcome at these events; however, you may need special permission to attend a journal club. Just ask a grad student or professor to be sure. It is certainly possible that you could meet a future advisor (see Selecting a potential advisor) at one of these events. Graduate-level courses provide a window into the graduate school experience. They can expose you to the flavor and tone of graduate school and will allow you to interact on a regular basis with graduate students. Do not think these courses will be over your head; often, they are no more difficult than undergraduate classes. Primarily undergraduate institutions may not offer graduate courses; if so, talk with your professors about acquiring skills that will help you prepare for graduate school, such as analyzing data in R, learning ArcGIS, or reading and discussing scientific papers with a group. Consider organizing a group of students with similar goals to work on these activities together, and at some colleges, you can register these groups as student-led courses for credit. Graduate students at research universities typically plunge right into a Ph.D. program. Consider completing a master’s degree if you are unsure whether you want to commit to a lengthy Ph.D. program, if you are really unclear about your research interests, or if you are not sure that research is definitely for you. You will get additional experience and will be able to choose a Ph.D. program with much greater insight and clarity. Alternatively, there are often opportunities to become a full-time paid research technician where you can acquire important research experience (see the Job Boards mentioned above). Start the application process well before applications are due, which is typically from early December to early February for programs that begin the following Fall. A spreadsheet might be helpful with entries such as application deadlines, whether the GRE is required, whether there are internal fellowship opportunities at the institution, and whether there is an application fee. Note that only a small number of programs accept graduate students in the middle of the year; thus, it is a once-a-year process! You should ask your mentors to help with your application. Your university’s career center may also be helpful in helping you find fellowships, connect you to alumni who have pursued graduate school, and write your application statements. There are also experienced mentors from around the country that are willing to help anyone with the application process; we encourage you to seek out this help (see https://dynamicecology.wordpress.com/2018/08/21/call-for-mentors-and-mentees-for-eebmentormatch/. You will need to secure at least three letters of recommendation for your application. These recommendations should come primarily from faculty, but one may also come from a senior graduate student or job supervisor. Choose people who know your abilities well. It is fine to email professors to request a letter of recommendation and to ask them if they are willing to write you a positive letter. Attach your résumé and offer to provide the professor a list of bullet points that highlight your accomplishments that might not be obvious on your résumé. These could include fluency in a second language, how you overcame significant hardships, lead guitarist in a rock band, or anything else that would help the professor compose a more compelling letter. Make sure you give the professor at least a three-week window, if not more, to submit your letter on time. Most programs will have you provide the email address of each person who has agreed to provide a letter of support. These programs will often automatically send emails to those faculty requesting that they submit a letter by a specific date. Faculty can be notoriously bad about getting recommendations in on time. It is your job to ensure that individuals who are writing your recommendations actually send them in. Double-check this and if the letters have not arrived by close to the due date, contact the faculty member and request politely that they send the letter ASAP. Identify the general area of research you would like to pursue. It should be more specific than just ecology or plant ecology, but it should not be so narrow as to exclude relevant options. Seek advice from your mentors. Although it may be difficult, it is important to try to narrow your interests. This underscores the benefit of gaining exposure to different research areas as an undergraduate so that you can begin to narrow your interests. One way to gain further insight is to explore job opportunities in research after you graduate but before starting grad school. This may help you hone in on an area of research that you really enjoy (see the job boards mentioned above). Select a range of institutions from major research universities to smaller Ph.D. granting institutions. You should choose at least one university where you are fairly certain of being admitted. It is sometimes the case that large research universities may be less likely to accept master’s students, or that these applicants are given lower priority or less funding than students applying to doctoral programs. This varies by department and discipline, so do your research or ask to be sure. Attending graduate school while caregiving is challenging. Graduate school is demanding, requires a high level of commitment, and does not pay well, but it can be tremendously rewarding. You can succeed in graduate school while caregiving, especially when you find a department with supportive and well-established programs that are already in place. Here, we offer insight and advice to those who have children or plan to have children during graduate school. Some of our recommendations are likely specific to graduate schools in the United States, where federal and institutional support for families is limited relative to many other countries. We divided considerations into four categories: Location, Institutional Support, Departmental Culture, and Laboratory Culture. We note that similar challenges may apply to students with adult dependents. Recognize the difference between “accommodation” and “support.” An advisor insisting that they are “flexible” is accommodation; a department offering paid parental leave is support. When deciding on a program, be careful not to make too many compromises that could put an unmanageable burden on yourself. It may be a cliché, but graduate school is a marathon, not a sprint. When you have children, that marathon maybe even longer and steeper, but it can also be enormously rewarding. For the sake of yourself and your family, choose a program that provides tangible support for parents at multiple levels. Do not worry about asking too many questions. Asking questions will allow you to identify programs that are a good fit for you; showing initiative will also reflect well on you to prospective recruiters. Being assertive demonstrates that you understand the challenges and that you are ready to take on the challenging but rewarding path ahead of you. For quite some time, universities and departments have been promoting diversity initiatives to support minoritized students and to combat tokenization of students from underrepresented groups; these initiatives gained significant additional momentum in 2020. While it is great to see this goal on institutional websites, you should evaluate the degree to which these initiatives are primarily publicity statements rather than actual commitments to diversity. Ask yourself: Are universities truly changing current policies and what actions have institutions and departments taken to make their programs more inclusive and welcoming? Here, we offer insight and advice to assist in identifying programs that are working toward solving the many serious inequities in ecology and evolutionary biology (Graves 2019, Tseng et al. 2020). There are many dimensions of diversity including race, gender, religion, sexual orientation, among others. To help minoritized individuals identify an outstanding program, we divide this box into four considerations: Location, Institutional Support, Departmental Culture, and Laboratory Culture. You should strive to find a supportive program where you will thrive and be accepted for who you are. Seek out programs that genuinely invest in inclusion and equity and provide real support rather than mere accommodation. Find an institution that welcomes you, one that invests in programs that provide marginalized students with critical social, cultural, and mental health support. Departments that provide a one-time fee waiver for your application are accommodating, but a department that provides a livable wage to live in a safe and welcoming neighborhood is supportive. Choose well and wisely. I was used to being an outlier, smoothing my rough edges, moderating my responses to trend to the mean. Attending schools where I was often one of two black kids in a class of 25 was the norm. Being a black girl who thought animals were fascinating required another layer of adjustment as well. My history in ecology began as an undergrad. I was at the University of Pennsylvania and all of my profs in biology were white men, the grad TA’s were mostly white men, with 1 or 2 white females for leavening. I took several courses that fascinated me, particularly Ecology with Dan Janzen and a behavioral ecology field course with W. John Smith. Although I was deeply interested in behavioral ecology, I had no model for how to pursue it. I grew up in the inner city, raised by parents whose career aspirations had been stifled by the lack of opportunities for smart, young black people. Upon graduating, I worked a few years in medical research and then decided to pursue a master’s degree, mostly in hopes of getting a better job. Because my transcript best matched the Ecology and Evolution program at the University of Pittsburgh, that is where I landed. It was like being in a foreign country. None of my family had ever held a non-clinical degree; I was not raised to think about graduate school, unlike nearly all of my fellow students and mentors. I still remember taking my comprehensive examination at the end of my first year. I spent 2 hours sequestered in a conference room with half a dozen white middle-aged men, desperately trying to figure out what they expected of me. I knew that they were (mostly) only interested in helping me, but I could not comprehend the nuances of their questions. I finally excused myself to get a drink and let the cool water from the fountain bathe my eyes as I cried. They took pity on me and I continued in the program. Fieldwork was traumatic; aside from a few weeks at summer camp, I had not spent time in the field and had certainly never contemplated standing alone in a marsh in hip waders on a cold April day while hunters unleashed shotgun blasts nearby! My academic career as a graduate student in ecology was like my undergrad experience. I became so accustomed to being the only female and only person of color in the room that I stopped noticing. I could not even let myself imagine that being denied a fellowship, a cushy TA assignment, funding, etc. could have anything to do with my color or gender; that way lay despair. Although falling in love with teaching was my impetus to finish my Ph.D., I never intended to join the tenure stream. When I look back on it now, I am sure that the alienation I felt was a major (if silent) contributor to that choice. How far have we come since those days nearly 40 years ago? In my department, we have ~60 faculty (tenure and appointment stream). For the last 25 years, I have been the only self-identified minority faculty member, but now we have two. Our department, like our university and academia in general, is starting to shift. Like the first hints of dawn after a long, cold night, hope is on the horizon. But, we still have a long way to go. Another important consideration in choosing a program is whether there are opportunities to acquire career training beyond research and traditional coursework. Not all students want to pursue the prototypical track of obtaining their doctorate, moving on to a postdoctoral position, and then accepting a professorship at a Ph.D. granting institution. These tenure-stream professorships tend to be scarce and fairly competitive. Fortunately, there are a wide array of other employment opportunities once you complete your degree, including science communication, working in the private sector (e.g., ecological consulting, natural history museums), teaching at a primarily undergraduate institution, and state and federal positions (e.g., U.S. Forest Service, policy advising). Does the university or the department you are considering offer courses, certificates, or training in pedagogy, statistics, GIS, or science communication that will allow you to expand your skill set in a way that would be attractive to employers? Does the department have affiliations with researchers or adjunct or affiliated faculty who work in these other sectors that could provide career advice? The ESA journal Frontiers in Ecology and Environment has an Exploring Ecological Careers section where more than a dozen articles explore career options outside of academia (https://esajournals.onlinelibrary.wiley.com/doi/toc/10.1002/(ISSN)1540-9309.Exploring-Ecological-Careers). Your selection of an advisor is the most important decision you will make with regard to your graduate degree. Identify 5–10 professors who might serve as your potential advisor in graduate school. Find their laboratory webpage or Google Scholar page to understand more about their research. These professors should be conducting research in an area you are interested in, and at universities, you are interested in attending. Some universities that you are interested in attending may not have faculty that work in your area of interest, while some faculty with strong research overlaps may not be at an institution you would consider attending. The faculty member is almost always more important than the specific university (Fox 2020) and narrowing your scope of institutions may also limit your chances of finding the right faculty advisor. Do not go into this without careful thought. Ask mentors for advice on appropriate advisors. Although it may be possible to switch advisors once you enroll, switching advisors can sometimes become awkward and politically difficult, and there may not be another professor who has an opening for a student or one who matches your research interests. Thus, choose your advisor as wisely as possible in the first place (for some advice, see The Interview below). Finally, the degree that a potential advisor is progressive in terms of diversity and inclusion or is family-friendly or both may be of central importance for many applicants (Boxes 1, 2 offer advice). You

For decades, outbreaks of insect herbivores in tropical forests were considered unusual or rare events primarily because of high plant diversity and the top-down impact of enemies. An alternative explanation is that these outbreaks are common but occur on sparsely distributed hosts high in the canopy and at scales of one or a few individual trees. Here, we report an outbreak of a saturniid in the genus Citioica Travassos & Noronha near the Amazon Basin of Ecuador on a single tree of Inga edulis Mart. The outbreak caused near complete defoliation (>90% leaf loss) and did not occur on nearby conspecifics. This is only the twenty-third documented case of a saturniid outbreak, of which more than 60% occurred in tropical habitats. This is the first report of an outbreak on a single tree. Members of the local indigenous communities are well aware of these Citioica outbreaks and collect these caterpillars for food whenever outbreaks are detected, suggesting that these isolated outbreaks are fairly common. Further research is required to explore the possibility that insect outbreaks in tropical forests may be more common than previously suspected but occur over very small spatial scales undetected high in the forest canopy.

During foraging, moose break stems of numerous saplings in order to lower shoot tips to a height they can reach. Both stem-breaking and bedding occur throughout the range of moose and these behaviors together create disturbed patches, which increase habitat heterogeneity forest-wide. Here, we explore this phenomenon in two ways: a case study in which we quantify the size of moose beds and evaluate the physical and taxonomic extent of stem breaking for more than 300 broken trees at seven sites in western Massachusetts and Connecticut, and a literature review on stem-breaking. In our case study, moose broke the stems of 17 deciduous woody species and were twice as likely to snap main stems rather than lateral stems. After breaking a stem, moose browsed shoot tips 96% of the time. Moose snapped primarily shade-intolerant or intermediately tolerant species (92%); the frequency of broken shade-intolerant trees was 2-4 times greater than the frequency of their occurrence in the stand, and the only shade-tolerant species moose broke were shrubs or understory tree species. We measured the 10 moose beds that we encountered; they ranged in size from 1.5-3.4 m(2) (mean = 2.5 m(2)). We performed a systematic review of stem breaking by moose throughout their range and found 14 publications that characterized stem-breaking by moose in Europe and North America. These studies typically focused on a few tree species of economic importance, on how stem-breaking increased forage access to the tips of broken trees, or how moose damage in general redirected or delayed succession. The one report of stem-breaking across all species found 1% of all trees were broken, while as many as 60% of individuals of preferred species were broken at sites in both North America and Europe. Stem-breaking contributes to keeping some forest stands in an earlier stage of succession, and bedding may create small patches of crushed vegetation that may enhance local plant diversity at small spatial scales. Thus, our findings and review support previous research that characterize moose as ecosystem engineers, keystone species, or both.

Monospecific stands of invasive plants can dramatically restructure habitat for fauna, thereby elevating population densities or promoting foraging of consumer species who benefit in the altered habitat. For example, dense stands of invasive plants may protect small mammals from predators, which in turn could increase foraging pressure on seeds that small mammals feed upon. We used a before-after, control-impact experimental design to test whether small mammal capture rates were higher and giving-up densities (GUDs) lower beneath dense stands of Berberis thunbergii, an invasive shrub with a rapidly expanding range throughout eastern North America. Our experimental design included three plot categories: 1) plots heavily invaded by B. thunbergii, 2) control plots lacking invasive shrub cover, and 3) invaded plots where we eradicated B. thunbergii midway through the study. Although our overall small mammal capture rate was low, small mammal captures were 65% higher in B. thunbergii invaded habitat relative to control plots and eradication lowered capture rates by 77%. GUDs were also 26% higher within B. thunbergii relative to control plots and eradication decreased GUDs by 65%. Our findings suggest that small mammals perceive dense stands of B. thunbergii as relatively safe foraging habitat. Prior surveys within our study locations revealed dramatically depressed tree seedling densities under B. thunbergii, thus invasive plants may promote intensive foraging by small mammals and reduce recruitment for species with foraged seeds or seedlings.

Forests are the most diverse and productive terrestrial ecosystems on Earth, so sustainably managing them for the future is a major global challenge. Yet, our understanding of forest diversity relies almost exclusively on the study of trees. Here, we demonstrate unequivocally that other growth forms (shrubs, lianas, herbs, epiphytes) make up the majority of vascular plant species in both tropical and temperate forests. By comparing the relative distribution of species richness among plant growth forms for over 3,400 species in 18 forests in the Americas, we construct the first high-resolution quantification of plant growth form diversity across two ecologically important regions at a near-continental scale. We also quantify the physical distribution of plant species among forest layers, that is, where among the vertical strata plants ultimately live their adult lives, and show that plants are strongly downshifted in temperate forests vs. tropical forests. Our data illustrate a previously unquantified fundamental difference between tropical and temperate forests: what plant growth forms are most speciose, and where they ultimately live in the forest. Recognizing these differences requires that we re-focus ecological research and forest management plans to encompass a broader suite of plant growth forms. This more holistic perspective is essential to conserve global biodiversity.

Salvage logging after forest disturbances is a controversial but common practice, yet it remains relatively poorly studied, particularly after windstorms within deciduous forests. Here, we present results from the first growing season following salvage logging on four gaps (3.5–6 ha) created by a 2012 tornado. While salvage logging shifted the distribution of stem heights to smaller sizes, it increased woody species seedling diversity by 33% and total plant richness and diversity by 20%. Salvaging did not alter or homogenize community composition. These short-term differences in richness and diversity were partly driven by a more even distribution of seedlings among species and increased species recruitment from the seed bank after salvaging. Overall, the initial impacts of salvage operations in a mixed deciduous forest redirected forest regeneration towards a more diverse community with a wider pool of species.