Mark Novak

Why intraspecific trait variation matters in community ecology

Natural populations consist of phenotypically diverse individuals that exhibit variation in their demographic parameters and intra- and inter-specific interactions. Recent experimental work indicates that such variation can have significant ecological effects. However, ecological models typically disregard this variation and focus instead on trait means and total population density. Under what situations is this simplification appropriate? Why might intraspecific variation alter ecological dynamics? In this review we synthesize recent theory and identify six general mechanisms by which trait variation changes the outcome of ecological interactions. These mechanisms include several direct effects of trait variation per se and indirect effects arising from the role of genetic variation in trait evolution.

UNDERSTANDING AND PREDICTING ECOLOGICAL DYNAMICS: ARE MAJOR SURPRISES INEVITABLE

Ecological surprises, substantial and unanticipated changes in the abundance of one or more species that result from previously unsuspected processes, are a common outcome of both experiments and observations in community and population ecology. Here, we give examples of such surprises along with the results of a survey of well-established field ecologists, most of whom have encountered one or more surprises over the course of their careers. Truly surprising results are common enough to require their consideration in any reasonable effort to characterize nature and manage natural resources. We classify surprises as dynamic-, pattern-, or intervention-based, and we speculate on the common processes that cause ecological systems to so often surprise us. A long-standing and still growing concern in the ecological literature is how best to make predictions of future population and community dynamics. Although most work on this subject involves statistical aspects of data analysis and modeling, the frequency and nature of ecological surprises imply that uncertainty cannot be easily tamed through improved analytical procedures, and that prudent management of both exploited and conserved communities will require precautionary and adaptive management approaches.

Predicting community responses to perturbations in the face of imperfect knowledge and network complexity

How best to predict the effects of perturbations to ecological communities has been a long-standing goal for both applied and basic ecology. This quest has recently been revived by new empirical data, new analysis methods, and increased computing speed, with the promise that ecologically important insights may be obtainable from a limited knowledge of community interactions. We use empirically based and simulated networks of varying size and connectance to assess two limitations to predicting perturbation responses in multispecies communities: (1) the inaccuracy by which species interaction strengths are empirically quantified and (2) the indeterminacy of species responses due to indirect effects associated with network size and structure. We find that even modest levels of species richness and connectance (-25 pairwise interactions) impose high requirements for interaction strength estimates because system indeterminacy rapidly overwhelms predictive insights. Nevertheless, even poorly estimated interaction strengths provide greater average predictive certainty than an approach that uses only the sign of each interaction. Our simulations provide guidance in dealing with the trade-offs involved in maximizing the utility of network approaches for predicting dynamics in multispecies communities.

Global patterns of kelp forest change over the past half-century

Significance Kelp forests support diverse and productive ecological communities throughout temperate and arctic regions worldwide, providing numerous ecosystem services to humans. Literature suggests that kelp forests are increasingly threatened by a variety of human impacts, including climate change, overfishing, and direct harvest. We provide the first globally comprehensive analysis of kelp forest change over the past 50 y, identifying a high degree of variation in the magnitude and direction of change across the geographic range of kelps. These results suggest region-specific responses to global change, with local drivers playing an important role in driving patterns of kelp abundance. Increased monitoring aimed at understanding regional kelp forest dynamics is likely to prove most effective for the adaptive management of these important ecosystems. Kelp forests (Order Laminariales) form key biogenic habitats in coastal regions of temperate and Arctic seas worldwide, providing ecosystem services valued in the range of billions of dollars annually. Although local evidence suggests that kelp forests are increasingly threatened by a variety of stressors, no comprehensive global analysis of change in kelp abundances currently exists. Here, we build and analyze a global database of kelp time series spanning the past half-century to assess regional and global trends in kelp abundances. We detected a high degree of geographic variation in trends, with regional variability in the direction and magnitude of change far exceeding a small global average decline (instantaneous rate of change = −0.018 y−1). Our analysis identified declines in 38% of ecoregions for which there are data (−0.015 to −0.18 y−1), increases in 27% of ecoregions (0.015 to 0.11 y−1), and no detectable change in 35% of ecoregions. These spatially variable trajectories reflected regional differences in the drivers of change, uncertainty in some regions owing to poor spatial and temporal data coverage, and the dynamic nature of kelp populations. We conclude that although global drivers could be affecting kelp forests at multiple scales, local stressors and regional variation in the effects of these drivers dominate kelp dynamics, in contrast to many other marine and terrestrial foundation species.

Selection on stability across ecological scales

Much of the focus in evolutionary biology has been on the adaptive differentiation among organisms. It is equally important to understand the processes that result in similarities of structure among systems. Here, we discuss examples of similarities occurring at different ecological scales, from predator-prey relations (attack rates and handling times) through communities (food-web structures) to ecosystem properties. Selection among systemic configurations or patterns that differ in their intrinsic stability should lead generally to increased representation of relatively stable structures. Such nonadaptive, but selective processes that shape ecological communities offer an enticing mechanism for generating widely observed similarities, and have sparked new interest in stability properties. This nonadaptive systemic selection operates not in opposition to, but in parallel with, adaptive evolution.

Long-Term Studies Contribute Disproportionately to Ecology and Policy

Abstract As the contribution for long‐term ecological and environmental studies (LTEES) to our understanding of how species and ecosystems respond to a changing global climate becomes more urgent, the relative number and investment in LTEES are declining. To assess the value of LTEES to advancing the field of ecology, we evaluated relationships between citation rates and study duration, as well as the representation of LTEES with the impact factors of 15 ecological journals. We found that the proportionate representation of LTEES increases with journal impact factor and that the positive relationship between citation rate and study duration is stronger as journal impact factor increases. We also found that the representation of LTEES in reports written to inform policy was greater than their representation in the ecological literature and that their authors particularly valued LTEES. We conclude that the relative investment in LTEES by ecologists and funders should be seriously reconsidered for advancing ecology and its contribution to informing environmental policy.

Estimating nonlinear interaction strengths: an observation-based method for species-rich food webs.

Efforts to estimate the strength of species interactions in species-rich, reticulate food webs have been hampered by the multitude of direct and indirect interactions such systems exhibit and have been limited by an assumption that pairwise interactions display linear functional forms. Here we present a new method for directly measuring, on a per capita basis, the nonlinear strength of trophic species interactions within such food webs. This is an observation-based method, requiring three pieces of information: (1) species abundances, (2) predator and prey-specific handling times, and (3) data from predator-specific feeding surveys in which the number of individuals observed feeding on each of the predators prey species has been tallied. The method offers a straightforward way to assess the completeness of ones sampling effort in accurately estimating interaction strengths through the construction of predator-specific prey accumulation curves. The method should be applicable to a variety of systems in which empirical estimates of direct interaction strengths have thus far remained elusive.

Using the functional response of a consumer to predict biotic resistance to invasive prey

Predators sometimes provide biotic resistance against invasions by nonnative prey. Understanding and predicting the strength of biotic resistance remains a key challenge in invasion biology. A predators functional response to nonnative prey may predict whether a predator can provide biotic resistance against nonnative prey at different prey densities. Surprisingly, functional responses have not been used to make quantitative predictions about biotic resistance. We parameterized the functional response of signal crayfish (Pacifastacus leniusculus) to invasive New Zealand mud snails (Potamopyrgus antipodarum; NZMS) and used this functional response and a simple model of NZMS population growth to predict the probability of biotic resistance at different predator and prey densities. Signal crayfish were effective predators of NZMS, consuming more than 900 NZMS per predator in a 12-h period, and Bayesian model fitting indicated their consumption rate followed a type 3 functional response to NZMS density. Based on this functional response and associated parameter uncertainty, we predict that NZMS will be able to invade new systems at low crayfish densities (< 0.2 crayfish/m2) regardless of NZMS density. At intermediate to high crayfish densities (> 0.2 crayfish/m2), we predict that low densities of NZMS will be able to establish in new communities; however, once NZMS reach a threshold density of -2000 NZMS/m2, predation by crayfish will drive negative NZMS population growth. Further, at very high densities, NZMS overwhelm predation by crayfish and invade. Thus, interacting thresholds of propagule pressure and predator densities define the probability of biotic resistance. Quantifying the shape and uncertainty of predator functional responses to nonnative prey may help predict the outcomes of invasions.

Estimating interaction strengths in nature: experimental support for an observational approach

The complexity of food webs poses a significant hurdle for our growing understanding of the structure and dynamics of ecological communities. Empirical methods that measure the per capita strengths of trophic species interactions offer a means to identify keystone species and bridge mathematical models and data to synthesize our knowledge of population dynamics and predator feeding behaviors. Many such methods have been proposed, but few have seen independent validation of their estimates or underlying assumptions. This is particularly so with respect to the nonlinear functional responses by which predators often respond to their prey. Here I describe an empirical test of a recently proposed observational method for estimating the nonlinear strength of predator-prey interactions in the field. By applying the method to two populations of a predatory intertidal whelk, Haustrum scobina, I estimated its per capita attack rates on all nine of its observed prey species. These spanned two orders of magnitude in per capita strength. Concurrent experimental manipulations of the two predator populations provided population time series for the response of a mussel prey species, Xenostrobus pulex. I obtained independent interaction strength estimates for this focal interaction by fitting a sequence of hypothesized predator-prey models to these time series. Overall, site-specific models assuming linear functional responses performed better than all others. A direct comparison of the attack-rate estimates from the observational method with those of the best-performing nonlinear model nevertheless revealed high concordance between the two methods. The results of this study therefore support the use of the observational method in larger and more complex food webs and suggest that trophic interactions in the range of mean prey densities observed in nature are approximately linear.

Diurnal activity in a group of Gulf of Maine decapods

[The patterns of diel activity of four large decapod species in the shallow subtidal of the Isles of Shoals, Gulf of Maine, U. S. A. were investigated. During the summer of 1999 the diel abundance and size distribution of active decapod individuals were surveyed at three depth ranges at a sheltered site on Appledore Island. Densities of active American lobsters, Homarus americanus H. Milne Edwards, 1837, were, as expected, highest at night. The crabs Cancer borealis Stimpson, 1859 and Carcinus maenas (L., 1758), however, were almost exclusively active during the day. Cancer irroratus Say, 1817 were equally active during the day and the night, but the mean size of individuals was significantly larger during the day. Surveys at additional sites in 2003 confirmed that these same patterns of diel activity were present throughout the Isles of Shoals. An extensive review of the literature suggests that such diurnal activity is not only unusual for the three crab species of this study, but for the whole genus Cancer as well. Untersucht wurden die Muster der Tag-Nacht-Aktivitat vierer groser Dekapoden-Arten aus dem Flachwasser der Isles of Shoals im Golf von Maine, U. S. A. Auf der geschutzten westlichen Seite der Insel Appledore sind im Sommer 1999 Individuenzahl und Grosenverteilung aktiver Dekapoden in drei Tiefenbereichen ermittelt worden. Der Amerikanische Hummer Homarus americanus H. Milne Edwards, 1837 erwies sich erwartungsgemas als nachtaktiv, wahrend die beiden Krabben Cancer borealis Stimpson, 1859 und Carcinus maenas (L., 1758) fast ausschlieslich tagaktiv waren. Cancer irroratus Say, 1817 war tags genauso aktiv wie nachts, blos war die durchschnittliche Korpergrose der Individuen am Tage signifikant groser. Stichproben im Bereich dreier weiterer Inseln im Jahr 2003 zeigten, dass diese Arten im Gesamtbereich der Isles of Shoals dieselben Muster der TagNacht-Aktivitat aufweisen. Eine umfassende Literaturrecherche ergab, dass die hier festgestellte Tag-Nacht-Aktivitat nicht nur fur die drei untersuchten Krabben hochst ungewohnlich ist, sondern auch fur alle Arten der Gattung Cancer insgesamt., The patterns of diel activity of four large decapod species in the shallow subtidal of the Isles of Shoals, Gulf of Maine, U. S. A. were investigated. During the summer of 1999 the diel abundance and size distribution of active decapod individuals were surveyed at three depth ranges at a sheltered site on Appledore Island. Densities of active American lobsters, Homarus americanus H. Milne Edwards, 1837, were, as expected, highest at night. The crabs Cancer borealis Stimpson, 1859 and Carcinus maenas (L., 1758), however, were almost exclusively active during the day. Cancer irroratus Say, 1817 were equally active during the day and the night, but the mean size of individuals was significantly larger during the day. Surveys at additional sites in 2003 confirmed that these same patterns of diel activity were present throughout the Isles of Shoals. An extensive review of the literature suggests that such diurnal activity is not only unusual for the three crab species of this study, but for the whole genus Cancer as well. Untersucht wurden die Muster der Tag-Nacht-Aktivitat vierer groser Dekapoden-Arten aus dem Flachwasser der Isles of Shoals im Golf von Maine, U. S. A. Auf der geschutzten westlichen Seite der Insel Appledore sind im Sommer 1999 Individuenzahl und Grosenverteilung aktiver Dekapoden in drei Tiefenbereichen ermittelt worden. Der Amerikanische Hummer Homarus americanus H. Milne Edwards, 1837 erwies sich erwartungsgemas als nachtaktiv, wahrend die beiden Krabben Cancer borealis Stimpson, 1859 und Carcinus maenas (L., 1758) fast ausschlieslich tagaktiv waren. Cancer irroratus Say, 1817 war tags genauso aktiv wie nachts, blos war die durchschnittliche Korpergrose der Individuen am Tage signifikant groser. Stichproben im Bereich dreier weiterer Inseln im Jahr 2003 zeigten, dass diese Arten im Gesamtbereich der Isles of Shoals dieselben Muster der TagNacht-Aktivitat aufweisen. Eine umfassende Literaturrecherche ergab, dass die hier festgestellte Tag-Nacht-Aktivitat nicht nur fur die drei untersuchten Krabben hochst ungewohnlich ist, sondern auch fur alle Arten der Gattung Cancer insgesamt.]