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Article Contents

Introduction, what is hbe, a systematic overview of current research, hbe: strengths, weaknesses, opportunities, and open questions, supplementary material, human behavioral ecology: current research and future prospects.

Forum editor: Sue Healy

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Daniel Nettle, Mhairi A. Gibson, David W. Lawson, Rebecca Sear, Human behavioral ecology: current research and future prospects, Behavioral Ecology , Volume 24, Issue 5, September-October 2013, Pages 1031–1040, https://doi.org/10.1093/beheco/ars222

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Human behavioral ecology (HBE) is the study of human behavior from an adaptive perspective. It focuses in particular on how human behavior varies with ecological context. Although HBE is a thriving research area, there has not been a major review published in a journal for over a decade, and much has changed in that time. Here, we describe the main features of HBE as a paradigm and review HBE research published since the millennium. We find that the volume of HBE research is growing rapidly, and its composition is changing in terms of topics, study populations, methodology, and disciplinary affiliations of authors. We identify the major strengths of HBE research as its vitality, clear predictions, empirical fruitfulness, broad scope, conceptual coherence, ecological validity, increasing methodological rigor, and topical innovation. Its weaknesses include a relative isolation from the rest of behavioral ecology and evolutionary biology and a somewhat limited current topic base. As HBE continues to grow, there is a major opportunity for it to serve as a bridge between the natural and social sciences and help unify disparate disciplinary approaches to human behavior. HBE also faces a number of open questions, such as how understanding of proximate mechanisms is to be integrated with behavioral ecology’s traditional focus on optimal behavioral strategies, and the causes and extent of maladaptive behavior in humans.

Very soon after behavioral ecology (henceforth BE) emerged as a paradigm in the late 1960s and early 1970s, a tradition of applying behavioral ecological models to human behavior developed. This tradition, henceforth human behavioral ecology (HBE), quickly became an important voice in the human-related sciences, just as BE itself was becoming an established and recognized approach in biology more generally. HBE continues to be an active and innovative area of research. However, it tends not to receive the attention it might, perhaps in part because its adherents are dispersed across a number of different academic disciplines, spanning the life and social sciences. Although there were a number of influential earlier reviews, particularly by Cronk (1991) and Winterhalder and Smith (2000) , there has not been a major review of the HBE literature published in a journal for more than a decade. In this paper, we undertake such a review, with the aim of briefly but systematically characterizing current research activity in HBE, and drawing attention to prospects and issues for the future. The structure of our paper is as follows. In the section “What is HBE?”, we provide a brief overview of the HBE approach to human behavior. The section “A systematic overview of current research” presents our review methodology and briefly describes what we found. We argue that the HBE research published in the period since 2000 represents a distinct phase in the paradigm’s development, with a number of novel trends that require comment. Finally, the section “HBE: strengths, weaknesses, opportunities, and open questions” presents our reflections on the current state and future prospects of HBE, which we structure in terms of strengths, weaknesses, opportunities, and open questions.

BE is the investigation of how behavior evolves in relation to ecological conditions ( Davies et al. 2012 ). Empirically, there are 2 arms to this endeavor. One arm is the study of how measurable variation in ecological conditions predicts variation in the behavioral strategies that individuals display, be it at the between-species, between-population, between-individual, or even within-individual level. (Throughout this paper, “ecological conditions” is to be interpreted in its broadest sense, to include the physical and social aspects of the environment, as well as the state of the individual within that environment.). The other arm concerns the fitness consequences of the behavioral strategies that individuals adopt. Because fitness—the number of descendants left by individuals following a strategy at a point many generations in the future—cannot usually be measured within a study, this generally means measuring the consequences of behavioral strategies in some more immediate proxy currency related to fitness, such as survival, mating success, or energetic return. The 2 arms of BE are tightly linked to one another; the fitness consequences of some behavioral strategy will differ according to the prevailing ecological conditions. Moreover, central to BE is the adaptationist stance. That is, we expect to see, in the natural world, organisms whose behavior is close to optimal in terms of maximizing their fitness given the ecological conditions that they face. This expectation is used as a hypothesis-generating engine about which behaviors we will see under which ecological conditions. The justification for the adaptationist stance is the power of natural selection. Selection, other things being equal, favors genes that contribute to the development of individuals who are prone to behaving optimally across the kinds of environments in which they have to live ( Grafen 2006 ). Note that this does not imply that behavioral strategies are under direct genetic control. On the contrary, selection favors various mechanisms for plasticity, such as individual and social learning, exactly because they allow individuals to acquire locally adaptive behavioral strategies over a range of environments ( Scheiner 1993 ; Pigliucci 2005 ), and it is these plastic mechanisms that are often in immediate control of behavioral decisions. However, the capacity for plasticity is ultimately dependent on genotype, and plasticity is deployed in the service of genetic fitness maximization.

BE is also characterized by a typical approach, to which actual exemplars of research projects conform to varying degrees. This approach is to formulate simple a priori models of what the individual would gain, in fitness terms, by doing A rather than B, and using these models to make predictions either about how variation in ecological conditions will affect the prevalence of behaviors A and B, or about what the payoffs to individuals doing A and B will be, in some currency related to fitness. These models are usually characterized by the assumption that there are no important phylogenetic or developmental constraints on the range of strategies that individuals are able to adopt and also by a relative agnosticism about exactly how individuals arrive at particular behavioral strategies (i.e., about questions of proximate mechanism as opposed to ultimate function; Mayr 1961 ; Tinbergen 1963 ). The assumptions of no mechanistic constraints coming from the genetic architecture or the neural mechanisms are known, respectively, as the phenotypic gambit ( Grafen 1984 ) and the behavioral gambit ( Fawcett et al. 2012 ). To paraphrase Krebs and Davies (1981 ), “think of the strategies and let the mechanisms look after themselves.” We return to the issue of the validity of the behavioral gambit in particular in section “Open questions.” However, one of the remarkable features of early research in BE (what Owens 2006 calls “the romantic period of BE”) was just how well the observed behavior of animals of many different species was explained by very simple optimality models based on the gambits.

HBE is the study of human behavior from an adaptive perspective. Humans are remarkable for their ability to adapt to new niches much faster than the time required for genetic change ( Laland and Brown 2006 ; Wells and Stock 2007 ; Nettle 2009b ). HBE has been particularly concerned with explaining this rapid adaptation and diversity, and thus, the concept of adaptive phenotypic plasticity has been even more central to HBE than it is to BE in general. HBE represents a rejection of the notion that fundamentally different explanatory approaches are necessary for the study of human behavior as opposed to that of any other animal. Note that this does not imply that humans have no unique cognitive and behavioral mechanisms. On the contrary, they clearly do. Rather, it implies that the general scientific strategy for explaining behavior instantiated in BE remains similar for the human case: understand the fitness costs and benefits given the ecological context, make predictions based on the hypothesis of fitness maximization, and test them. There is a pleasing cyclicity to the development of HBE. BE showed that microeconomic models based on maximization, which had come from the human discipline of economics, could be used at least as a first approximation to predict the behavior of nonhuman animals. HBE imported these principles, enriched from their sojourn in biology by a focus on fitness as the relevant currency, back to humans again.

The first recognizably HBE papers appeared in the 1970s (e.g., Wilmsen 1973 ; Dyson-Hudson and Smith 1978 ). The pioneers were anthropologists, and to a lesser extent archaeologists. A major focus was on explaining foraging patterns in hunting and gathering populations ( Smith 1983 ), though other topics were also represented from the outset ( Cronk 1991 ). The focus on foragers was due to the evolutionary antiquity of this mode of subsistence, as well as these being the populations in which optimal foraging theory was most straightforwardly applicable. However, there is no reason in principle for HBE research to be restricted to such populations. The emphasis in HBE is on human adaptability; humans have mechanisms of adaptive learning and plasticity by virtue of which they can rapidly find adaptive solutions to living in many kinds of environments. Thus, we might expect their behavior to be adaptively patterned in societies of all kinds, not just the types of human society, which have existed for many millennia.

The first phase of HBE lasted through the 1980s ( Borgerhoff Mulder 1988 ). In the second phase, the 1990s, HBE grew rapidly, with Winterhalder and Smith (2000) estimating that there were nearly 300 studies published during the decade. Its focus broadened to encompass more studies from nonforaging subsistence populations, such as horticulturalists and pastoralists (e.g., Borgerhoff Mulder 1990 ), and the use of historical demographic data (e.g., Voland 2000 ; Clarke and Low 2001 ). There were also some pioneering forays into the BE of industrialized populations ( Kaplan 1996 ; Wilson and Daly 1997 ). The 1990s were characterized by an increasing emphasis on topics which fall under the general headings of distribution (cooperation and social structure) and particularly reproduction (mate choice, mating systems, reproductive decisions, parental investment), rather than production (foraging). Anthropologists continued to dominate HBE, and the methodologies of the studies reflect this: many of the studies represented the field observations of a single field researcher from a single population, usually a single site. Having briefly outlined what HBE is and where it came from, we now turn to reviewing the HBE research that has appeared in the years since the publication of Winterhalder and Smith (2000) .

Our objective was to ascertain what empirical research has been done within the HBE paradigm since 2000, and characterize its key features, quantitatively where possible. We thus conducted a systematic search of 17 key journals for papers published between the beginning of 2000 and late 2011, which clearly belong in the HBE tradition (see Supplementary material for full methodology). This involved some contentious decisions about how to draw the boundaries of HBE and in the end, we drew it narrowly, including only papers containing quantitative data on naturally occurring behavior in human populations and employing a clearly adaptive perspective. This excludes a large number of studies that take an adaptive perspective but measure hypothetical preferences or decisions in experimental scenarios. It also excludes many studies that focus on nonbehavioral traits such as stature or physical maturation. The sample is not exhaustive even of our chosen subset of HBE, given that some HBE research is published in edited volumes, books, or journals other than those we searched. However, we feel that our strategy provides a good transect through current research, which is prototypically HBE, and the sampling method is at least repeatable and self-consistent over time.

We used the full text of the papers identified to code a number of key variables relevant to our review, including year of publication, journal, first author country of affiliation, and first author academic discipline. We also adopted Winterhalder and Smith’s (2000) ternary classification of topics into production (foraging and other productive activity), distribution (resource sharing, cooperation, social structure), and reproduction (mate choice decisions, sexual selection, life-history decisions, parental and alloparental investment). Finally, we coded the presence of some key features we wished to examine: the presence of any data from foraging populations, the presence of any data from industrialized populations, the use of secondary data, and the use of comparative data from more than one population.

The search resulted in a database of 369 papers (see Supplementary material for reference list and formal statistical analysis; an endnote library of the references of the papers in the database is also available from the corresponding author). The distribution of papers across journals is shown in Table 1 , which also shows the median year of publication of a paper in that journal. The overall median year of publication for the full sample was 2007; thus, the table can be used to identify those journals that carried HBE papers disproportionately earlier in the study interval (e.g., American Anthropologist , median 2004), and those which carried them disproportionately more recently (e.g., American Journal of Human Biology , median 2009). The total number of papers found per year increased significantly over the 12 years sampled, from around 20 at the beginning to nearly 50 in 2011 ( Figure 1a ; regression analysis suggests an average increase of 2.4 papers per year). In the Supplementary material , we show that HBE papers also increased as a proportion of all papers published in our target journals. First authors were affiliated with institutions in 28 different countries, with 57.5% based in the United States and 20.1% in the United Kingdom. In terms of discipline, anthropology (including archaeology) was strongly represented (49.9% of papers), followed by psychology (19.5%) and biology (12.7%). The remaining papers came from demography (3.3%), medicine and public health (3.0%), sociology and social policy (2.4%), economics and political science (2.2%), or were for various reasons unclassifiable (7.0%). However, the growth in number of papers over time was due to increasing HBE activity outside anthropology ( Figure 1a ). In 2000–2003, 64.0% of papers were from anthropology departments, whereas by 2009–2011, this figure was 47.4%. Our search strategy may, if anything, have underestimated the growth in HBE research from outside anthropology, because our search strategy was based on the journals that had carried important BE or HBE research prior to 2000 and did not include any specialist journals from disciplines such as demography or public health.

Numbers and percentages of papers in the database by journal. Also shown is the median year of publication of an HBE paper in the sample in that journal

a Formerly Journal of Cultural and Evolutionary Psychology .

b Targeted search only; for all other journals, all abstracts read.

Number of published papers identified by year over the study period (a) by disciplinary affiliation of first author; (b) by type of study population (other = agriculturalist, pastoralist, horticulturalist, or multiple types); (c) by tripartite classification of topic.

In terms of type of population studied, 80 papers (21.7%) contained some data from foragers, broadly defined to include any subsistence population for whom foraging forms a substantial part of the diet. One hundred and forty-five papers (39.3%) contained data from industrialized populations. The remainder of papers studied either contemporary or historical agricultural, horticultural, and pastoral populations. As Figure 1b shows, the amount of work on industrialized populations has tended to increase over time, with 22 such papers in 2000–2002 (29.3% of total) and 58 in 2009–2011 (43.0%). By contrast, the amount of work on forager populations is much more stable (20 papers [26.7%] in 2000–2002, 27 papers [20.0%] in 2009–2011). As for topic, we classified 64.8% of our papers as concerning reproduction, with 9.5% concerning production and 13.3% distribution. The remaining 12.5% either spanned several topics or fit none of the 3 categories. Table 2 gives some examples of popular research questions addressed in each of the 3 topic areas. The preponderance of reproduction has increased over time ( Figure 1c ); in 2000–2002, 53.3% of the papers fell into this category, whereas by 2009–2011, it was 68.9%. In fact, the growth of HBE papers during the study period has been completely driven by an increase in papers on reproductive topics (see Supplementary material ). We classified papers according to whether they involved analysis of secondary data sets gathered for other purposes. The number of papers involving such secondary analysis increased sharply through the study period, whereas those involving primary data did not (see Supplementary material ). Comparative analyses also increased significantly over time, but not faster than the overall growth in paper numbers.

Some examples of popular research questions in our database of recent HBE papers

To summarize, the data suggest that HBE has changed measurably in the period since 2000. Some of the changes in this period represent continuations of trends already incipient before, such as the expansion away from foraging and foragers toward reproduction and other types of population ( Winterhalder and Smith 2000 ). Our analysis suggests that it is primarily research into the BE of industrialized societies, which has expanded in the subsequent years, such that over 40% of HBE research published in the most recent 3-year period was conducted on such populations. More “traditional” HBE studies of foraging and small-scale food producing societies have continued, but only at a modestly increased rate compared with the 1990s. An unexpected feature of HBE post-2000 is the expansion of HBE in disciplines outside anthropology. Much of the growth has come from the adoption of HBE ideas by researchers based in departments of psychology, and, to a modest extent, other social sciences such as demography, public health, economics, and sociology. This is concomitant with the increasing focus on large-scale industrialized societies, as well as changes in methodology. Anthropologists often work alone or in small teams to gather special-purpose, opportunistic data sets from a particular field site, and many of the pioneering HBE studies were done in this way. In demography, public health, and sociology, by contrast, research tends to be based on very large, systematically collected, representative data sets, such as censuses, cohort, and panel studies, which are designed with multiple purposes in mind. Particular researchers can then interrogate them secondarily to address their particular questions. As HBE has welcomed more researchers from these other social sciences, it has also adopted these secondary methods more strongly (see section “Strengths” for further discussion). We also note the increase in the number of comparative studies. Comparative methods (albeit usually comparing related species rather than populations of the same species) have been a strong feature of BE since the outset (or before, Cullen 1957 ), and thus this is a natural development for HBE. HBE comparative studies use existing cross-cultural databases ( Quinlan 2007 ), integrate multiple ethnographic or historical sources ( Brown et al. 2009 ), or, increasingly, coordinate researchers to collect or derive standardized measures across multiple populations ( Walker et al. 2006 ; Borgerhoff Mulder et al. 2009 ). Comparative studies have become more powerful in their analytical strategies (see section “Strengths”).

The literature review in section “A systematic overview of current research” allowed us to characterize current HBE research and show some of the ways it has changed in the last decade. In this section, we discuss what we see as the strengths, weaknesses, opportunities, and open questions for HBE as a paradigm. This is inevitably more of a personal assessment than the preceding sections, and we appreciate that not everyone in the field will share our views.

The first obvious strength of HBE is vitality . As Darwinians, it comes naturally to us to assume that something that is increasing in frequency has some beneficial features. Thus, the fact that the number of recognizably HBE papers per year found by our search strategy has doubled in a decade, and that there are more and more adopters outside of anthropology, indicates that a range of people find an HBE approach useful. Where does this utility spring from? In part, it is that HBE models tend to make very clear, a priori predictions motivated by theory. The same cannot be said of all other approaches in the human sciences, and, arguably, the more we complicate behavioral ecological models by including details about how proximate mechanisms work, the more this clarity tends to disappear. We return in section “Open questions” to the issue of whether agnosticism about mechanism can be justified, but we note here that a great strength of (and defense for) simple HBE models is that they so often turn out to be empirically fruitful, despite their simplicity. Whether we are considering when to have a first baby ( Nettle 2011 ), what the effects of having an extra child will be in different ecologies ( Lawson and Mace 2011 ), whether to marry polygynously, polyandrously, or monogamously ( Fortunato and Archetti 2010 ; Starkweather and Hames 2012 ), or which relatives to invest time and resources in ( Fox et al. 2010 ), predictions using simple behavioral ecological principles turn out to be useful in making sense of empirically observed diversity in behavior. HBE has also demonstrated the generality of certain principles, such as the fact that male culturally defined social success is positively associated with reproductive success in many different types of society, albeit that the slope of the relationship differs according to features of the social system ( Irons 1979 ; Kaplan and Hill 1985 ; Borgerhoff Mulder 1987 ; Hopcroft 2006 ; Fieder and Huber 2007 ; Nettle and Pollet 2008 ).

A related strength of HBE is its broad scope . HBE models can apply to many kinds of behavioral decision (in principle, all kinds) and in all kinds of society. It is relatively rare in the human sciences for the same set of predictive principles to apply to variation both within and between societies and to societies ranging from small-scale subsistence populations to large-scale industrial states, but HBE thinking about, for example, reproductive decisions has exactly this scope ( Nettle 2011 ; Sear and Coall 2011 ). This would be a strength indeed, even without the crucial additional feature that the explanatory principles invoked are closely related to those that can be applied to species other than our own. Thus, HBE brings a relative conceptual coherence to the study of human behavior, a study that has traditionally been spread across a number of different disciplines each with different conceptual starting points.

Another strength of HBE as we have defined it here is its relatively high ecological validity . Much psychological research into human behavior relies on hypothetical self-reports and self-descriptions, or contrived experimental situations ( Baumeister et al. 2007 ), and much of behavioral economics consists of artificial games whose relevance to actual allocation decisions outwith the laboratory has been questioned ( Levitt and List 2007 ; Bardsley 2008 ; Gurven and Winking 2008 ). Although human behavioral ecologists use such techniques as their purposes require, at the heart of HBE is still a commitment to looking at what people really do, in the environments in which they really live, as a central component of the endeavor. Furthermore, HBE’s focus on behavioral diversity means that it has studied a much wider range of populations than other approaches in the human sciences (see Henrich et al. 2010 ), and this has led to a healthy skepticism of simple generalizations about human universal preferences or motivations ( Brown et al. 2009 ). Measuring relationships between behavior and fitness-relevant outcomes across a broad range of environments, HBE has now amassed considerable evidence in favor of its core assumptions that context matters when studying the adaptive consequences of human behavior and that behavioral diversity arises because the payoffs to alternative behavioral strategies are ecologically contingent.

HBE is also characterized by increasing methodological rigor. The early phases of HBE were defined by exciting theoretical developments, as evolutionary hypotheses for human behavioral variation were first formulated and presented in the literature. However, conducting empirical studies capable of rigorously testing hypotheses derived from HBE theory presents a number of methodological challenges, not least because the human species is relatively long lived and rarely amenable to experimental manipulation. These challenges are now being increasingly overcome, as HBE expands its tool kit to include new sources of data, statistical methods, and study designs. As noted in the section “A systematic overview of current research,” recent years have witnessed an increased use of secondary demographic and social survey data sets, which often provide larger, more representative samples and a broader range of variables than afforded by field research. Some sources of secondary data have also enabled lineages to be tracked beyond the life span of any individual researcher, providing valuable new data on the correlates of long-term fitness (e.g., Lahdenpera et al. 2004 ; Goodman and Koupil 2009 ).

Statistical methods have also become more advanced. Multilevel analyses are now routinely used in HBE research to deal with hierarchically structured data and accurately partition sources of behavioral variance at different levels (e.g., within and between villages; Lamba and Mace 2011 ). Phylogenetic comparative methods, which utilize information on historical relationships between populations, have become popular for testing coevolutionary hypotheses since they were first applied to human populations in the early 1990s ( Mace and Pagel 1994 ; Mace and Holden 2005 ), though debate remains about their suitability for modeling behavioral transmission in humans ( Borgerhoff Mulder et al. 2006 ). Issues of causal inference are also being addressed with more sophisticated analytical techniques. For example, structural equation modeling and longitudinal methods such as event history analysis have enabled researchers to achieve greater confidence when controlling for potential cofounding relationships (e.g., Sear et al. 2002 ; Lawson and Mace 2009 ; Nettle et al. 2011 ). HBE researchers are also following wider trends in the social and natural sciences by exploring alternatives to classic significance testing, such as information-theoretic and Bayesian approaches for considering competing hypotheses ( Towner and Luttbeg 2007 ). Some researchers have also been able to harness “natural experiments” in situations where comparable populations or individuals are selectively exposed to socioecological change. For example, Gibson and Gurmu (2011) examined the effect of changes in land tenure (from family inheritance to government redistribution) on a population in rural Ethiopia, demonstrating that competition between siblings for marital and reproductive success only occurs when land is inherited across generations. These advancements represent an exciting and necessary step forward, as empirical methods “catch up” with the powerful theoretical framework set out in the early days of HBE.

Finally, HBE has shown itself capable of topical innovation. A pertinent recent example is cooperative breeding (typically loosely defined in HBE as the system whereby women receive help from other individuals in raising their offspring). The idea that human females might breed cooperatively had been around for several decades ( Williams 1957 ), and began to be tested empirically in the late 1980s and 1990s (e.g., Hill and Hurtado 1991 ), but it was the 21st century that saw a real upsurge in interest in this topic, leading to a revitalization of the study of kinship in humans ( Shenk and Mattison 2011 ). HBE has now mined many of the rich demographic databases available for our species to test empirically the hypothesis that the presence of other kin members is associated with reproductive outcomes such as child survival rates and fertility rates. These analyses typically find support for the hypothesis that women adopt a flexible cooperative breeding strategy where they corral help variously from the fathers of their children, other men, and pre- and postreproductive women ( Hrdy 2009 ).

Though we see HBE as a strong paradigm, there are some important weaknesses of its current research to be noted. The first is HBE’s relative isolation from the rest of BE. The core journals of BE are Behavioral Ecology and Behavioral Ecology and Sociobiology . Our search revealed only 8 HBE papers in these journals (2.2% of the sample). The vast majority of papers in our sample appeared in journals which never carry studies of species other than humans, and we know of rather few human behavioral ecologists who also work on other systems. West et al. (2011) have recently argued that evolutionary concepts are widely misapplied (or outdated understandings are applied, a phenomenon colloquially dubbed “the disco problem”) in human research, due to insufficient active integration between HBE and the rest of evolutionary biology.

HBE is clearly not completely decoupled from the rest of BE (see Machery and Cohen 2012 for quantitative evidence on this point). For example, within BE, there has been a decline in interest in foraging theory and a rise in interest in sexual selection ( Owens 2006 ), which are mirrored in the changes in HBE described in section “A systematic overview of current research.” Behavioral ecologists have also become less concerned with simply showing that animals make adaptive decisions, and more concerned with the nature of the neurobiological and genetic mechanisms underlying this ( Owens 2006 ). Parallel developments have occurred in the human literature, with the rise of adaptive studies of psychological mechanisms (see e.g., Buss 1995 ). Our search strategy did not include these studies, because their methodologies are different from those of “classical” HBE, but there is no doubt that they have increased in number. Finally, we note that there has been a recent increase in interest in measuring natural selection directly in contemporary human populations ( Nettle and Pollet 2008 ; Byars et al. 2010 ; Stearns et al. 2010 ; Milot et al. 2011 ; Courtiol et al. 2012 ). This anchors HBE much more strongly to evolutionary biology in general. Despite these developments, we see the isolation of HBE from the rest of biology as a potential risk. We hope to see more behavioral ecologists start to work on humans, and more projects across taxonomic boundaries, in the future.

Finally, we note the rather restricted topic base. HBE has had a great deal to say recently about mating strategies, reproductive decisions, fertility, and reproductive success, but much less about diet, resource extraction, resource storage, navigation, spatial patterns of habitat use, hygiene, social coordination, or the many other elements involved in staying alive. In part, this is because, as HBE expands to focus more on large-scale populations, it discovers that there are already disciplines (economics, sociology, human geography, public health) that deal extensively with these topics. It is in the general area of reproduction that it is easiest to come up with predictions that are obviously Darwinian and differentiate HBE from existing social science approaches. Nonetheless, the explanatory strategy of HBE is of potential use for any topic where behavioral effort has to be allocated in one way rather than another, and thus we would hope to see a broadening of the range of questions addressed as HBE continues to grow.

Opportunities

As HBE continues to expand, we see a major opportunity for HBE to build bridges to the social sciences. At the moment, most HBE papers are published in journals that only carry papers that take an adaptive evolutionary perspective, not general social science journals. Thus, HBE is possibly as separated from other approaches to human behavior as it is from parallel approaches to the behavior of other species. This may be because early proponents of HBE saw it as radically different from existing social science approaches to the same problems, by virtue of its generalizing hypothetico-deductive framework and commitment to quantitative hypothesis testing ( Winterhalder and Smith 2000 ). However, the social science those authors came into closest contact with was sociocultural anthropology, which is perhaps not a very typical social science (see Irons 2000 for an account of the hostile reception of HBE within sociocultural anthropology). As HBE’s expansion brings it into closer proximity with disciplines like economics, sociology, demography, public health, development studies, and political science, there may be more common ground than was previously thought. Social scientists are united in the notion that human behavior is very variable and that context is extremely important in giving rise to this variation. These are commitments that HBE obviously shares. Indeed, although it is still common in the human sciences for authors to rhetorically oppose “evolutionary” to “nonevolutionary” (or “social” and “biological”) explanations of the same problem as if these were mutually exclusive endeavors ( Nettle 2009a ), HBE defies such dichotomies adeptly.

Much of social science is highly quantitative and, generally lacking the ability to perform true experiments, relies on multivariate statistical approaches applied to observational data sets to test between competing explanations for behavior patterns. HBE is just the same, and indeed, since the millennium, has become much more closely allied to other social sciences, adopting the large-scale data resources they provide, as well as methodological tools like multilevel modeling, which they have developed to deal with these. HBE employs a priori models based on the individual as maximizer, a position not shared explicitly by all social sciences. However, this approach is widespread in economics and political science. Indeed, it was economics that gave it to BE. The big difference between HBE and much of social science is the explicit invocation of inclusive fitness (or its proxies) as the end to which behavior is deployed. This does not necessarily make it a competing endeavor, especially because what is measured in HBE is not usually fitness itself, but more immediate proxies. Rather, HBE models can often be seen as adding an explicitly ultimate layer of explanation, giving rise to new predictions and unifying diverse empirical observations, without being incompatible with existing, more proximate theories.

Indeed, our perception is that a number of social science theories make assumptions about the ends of behavior, which are quite similar to those of HBE, just not explicitly expressed in Darwinian terms; basically, people’s sets of choices are constrained by the environment in which they have to live, and they make the best choices they can given these constraints, often with knock-on effects that behavioral ecologists would describe as trade-offs. Examples include the work of Geronimus on how African American women adjust their patterns of childbearing to the prevailing rates of mortality and morbidity in their neighborhoods ( Geronimus et al. 1999 ), the work of Drewnowski and colleagues on how people adjust the type of foodstuffs they consume to the budgets they have to spend ( Drewnowski and Specter 2004 ; Drewnowski et al. 2007 ), or Downey’s work on the effects of increasing family size on socioeconomic outcomes of the children ( Downey 2001 ). If the introductory sections of any of these papers were written from a more explicitly Darwinian perspective, they would look perfectly at home in a BE journal. The breaking down of the social science–natural science divide has long been held as desirable, but is not easy to achieve in practice. HBE’s boundary with the social sciences may be one frontier where some progress can occur. Social scientists have long lamented the fragmentation of their field into multiple disciplinary areas with little common ground (e.g., Davis 1994 ). Given HBE’s broad scope and general principles, it has the potential to serve as something of a lingua franca across social scientists working on different kinds of problems.

A related opportunity for HBE is the potential for applied impact . HBE models have the potential to provide new and practical insights into contemporary world issues, from natural resource management ( Tucker 2007 ) to the consequences of inequality within developed populations ( Nettle 2010 ). The causes and consequences of recent human behavioral and environmental changes (including urbanization, economic development, and population growth) are recurring themes in recent studies in HBE. The utility of an ecological approach is clearly demonstrated in studies exploring the effectiveness of public policies or intervention schemes seeking to change human behavior or environments. HBE models clarify that human behavior tends to be deployed in the service of reproductive success, not financial prudence, health, personal or societal wellbeing ( Hill 1993 ), an important insight that differs from some economic or psychological theories. By providing insights into ultimate motivations and proximate pathways to human behavioral change, HBE studies can sometimes offer direct recommendations for the design and implementation of future initiatives ( Gibson and Mace 2006 ; Shenk 2007 ; Gibson and Gurmu 2011 ). Addressing contemporary world issues does, however, present methodological and theoretical challenges for HBE, requiring more explicit consideration of how research insights may be translated into interventions and communicated to policymakers and users ( Tucker and Taylor 2007 ).

Open questions

An open question for HBE is how the study of mechanism can be integrated into functional enquiry. This is an issue for BE generally, not just the human case. As mentioned in the section “What is HBE?”, BE has tended to proceed by the behavioral gambit—the assumption that the nature of the proximate mechanisms underlying behavioral decisions is not important in theorizing about the functions of behavior. It is important to understand the status of the behavioral gambit because it has sometimes been unfairly criticized (see Parker and Maynard Smith 1990 ). In the natural world, individuals do not always behave optimally with respect to any particular decision because there are phylogenetic or mechanistic constraints on their ability to reach adaptive solutions. However, in general terms, the only way to discover the existence of such departures from optimality is to have a theoretical model that shows what the optimal behavior would be and to test empirically whether individual behavior shows the predicted pattern. Where it does not, this may point to unappreciated constraints or trade-offs and thus shed light on the biology of the organism under study. Thus, the use of the term gambit is entirely apt; the behavioral gambit is a way of opening the enquiry designed to gain some advantage in the quest to understand. It is not the end game.

Where there is no sizable departure from predicted optimality, the ultimate adaptive explanation does not depend critically on understanding the mechanisms. This does not mean the question of mechanism is unimportant, of course; mechanistic explanations must still be sought and integrated with functional ones. This is beginning to occur in some cases. In the field of human reproductive ecology, the physiological mechanisms involved in adaptive strategies are beginning to be understood ( Kuzawa et al. 2009 ; Flinn et al. 2011 ), and there is also increasing interchange between HBE researchers and experimentalists studying psychological mechanisms ( Sear et al. 2007 ), which is clearly a development to be welcomed.

Where there is a patterned departure from optimality, understanding the mechanism becomes more critical. Aspects of mechanism can then be modeled as additional constraints, which may explain the strategies individuals pursue. For example, Kacelnik and Bateson (1996) showed that the pattern of risk aversion for variability in food amount and risk proneness for variability in food delay is not predicted by optimal foraging theory, except when Weber’s law (the principle that perceptions of stimulus magnitude are logarithmically, not linearly, related to actual stimulus magnitude) is incorporated into models as a mechanistic constraint. At a deeper level, though, this just raises further questions. Why should Weber’s law have evolved, and once it has evolved, can selection relax it for any particular task? These are what McNamara and Houston call “evo-mecho” questions ( McNamara and Houston 2009 ). Departures from optimality in one particular context raise such questions pervasively. Issues such as the robustness, neural instantiability, efficiency, and developmental cost of different kinds of mechanisms become salient here, and many apparently irrational quirks of behavior become interpretable as side effects of evolved mechanisms whose overall benefits have exceeded their costs over evolutionary time ( Fawcett et al. 2012 ). However, we would still argue that the best first approximation in understanding a question is to employ the behavioral gambit to generate and test simple optimality predictions, even though an understanding of mechanism will be essential for explaining why these may fail.

Although the issue of how incorporation of mechanism changes the predictions of BE models is a general one, in the human case, it has been discussed in particular with reference to transmitted culture because this is a class of mechanism on which humans are reliant to a unique extent ( Richerson and Boyd 2005 ). Transmitted culture refers to the behavioral traditions that arise from repeated social learning. Social learning can be an evolutionarily adaptive strategy, and the equilibrium solutions reached by it will often be the fitness-maximizing ones under reasonable assumptions ( Henrich and McElreath 2003 ). After all, if reliance on culture on average led to maladaptive outcomes, there would be strong selection on humans to rely on it less. Indeed, there is evidence that humans tend to forage efficiently for socially acquired information, using it when it is adaptive to do so ( Morgan et al. 2012 ). Thus, we would argue that culture can be treated, to a first approximation, just like any other proximate mechanism: that is, it can be set aside in the initial formulation of functional explanations ( Scott-Phillips et al. 2011 , though see Laland et al. 2011 for a different view). As an example, we could take Henrich and Henrich’s (2010) data on food taboos for pregnant and lactating women in Fiji. These authors show that the taboos reduce women’s chances of fish poisoning by 30% during pregnancy and 60% during breastfeeding and thus are plausibly adaptive. The fact that in this case it is culture by which women acquire them, rather than genes or individual learning, does not affect this conclusion or the data needed to test it. However, the quirks of how human social learning works may well explain some nonadaptive taboos that are found alongside the adaptive ones, which are in effect carried along by the generally adaptive reliance on social learning. Thus, although the behavioral gambit can be used to explain the major adaptive features of these taboos, an understanding of the cultural mechanisms is required to explain the details of how the observed behavior departs in subtle ways from the optimal pattern. Culture may often lead to maladaptive side effects in this way ( Richerson and Boyd 2005 ). Although its general effect is to allow humans to rapidly reach adaptive equilibria, nonadaptive traits can be carried along by it, and, compared with other proximate mechanisms, it produces very different dynamics of adaptive change.

A final open question is the extent of human maladaptation. Humans have increased their absolute numbers by orders of magnitude and colonized all major habitats of the planet, so they are clearly adept at finding adaptive solutions to the problem of living. However, there are also some clear cases of quite systematic departures from adaptive behavior. Perhaps most pertinently, the low fertility rate typical of industrial populations still defies a convincing adaptive explanation, despite being a longstanding topic for HBE research (see Borgerhoff Mulder 1998 ; Kaplan et al. 2002 ; Shenk 2009 ). There are patterns in the fertility of modernizing populations, which can be readily understood from an HBE perspective: parents in industrialized populations who have large families suffer a cost to the quality of their offspring, particularly with regard to educational achievement and adult socioeconomic success, so there is a quality–quantity trade-off ( Lawson and Mace 2011 ). Moreover, the reduction in fertility rate is closely associated with improvement in the survival of offspring to breed themselves, so that, as the transition to small families proceeds, the probability of having at least one grandchild may remain roughly constant ( Liu and Lummaa 2011 ). However, despite all this, it remains the case that people in affluent societies could still have many more grandchildren and great-grandchildren by having more children, and yet they do not ( Goodman et al. 2012 ). Any explanation of the demographic transition must, therefore, invoke some kind of maladaptation or mismatch between the conditions under which decision-making mechanisms evolved and those under which they are now operating.

Our review has shown that HBE is a growing and rapidly developing research area. The weaknesses of HBE mostly amount to a need for more research activity, and the unresolved questions, though important, do not in our view undermine HBE’s core strengths of theoretical coherence and empirical utility. HBE is being applied to more questions in more human populations with better methods than ever before. Our hope is that HBE will inspire more behavioral biologists to work on humans, for whom a wealth of data is available, and more social scientists to adopt an adaptive, ecological perspective on their behavioral questions, thus adding a layer of deeper explanations, as well as generating new insights.

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• Adam Driscoll, University of Wisconsin, La Crosse
• Alan Ewert, Indiana University
• Angela Franz-Balsen, German Society for Human Ecology
• Luc Hens, Flemish Institute for Technological Research
• Cassandra Johnson Gaither, USDA Forest Service
• Andrew K. Jorgenson, Boston College
• Stefano Longo, North Carolina State University
• Priscila Lopes, Universidade Federal do Rio Grande do Norte
• Andrew MacKenzie, The Australian National University
• Thom Meredith, McGill University
• Angela Mertig, Middle Tennessee State University
• Iva Pires, Universidade Nova de Lisboa
• Liam Phelan, University of Newcastle
• Melinda Storie, Northeastern Illinois University
• Mihnea Tanasescu, Free University of Brussels
• Joanne Vining, University of Illinios, Urbana
• Rachael Wakefield-Rann, University of Technology Sydney
• Cory Whitney, University of Bonn

Manuscripts should be submitted online via http://mstracker.com/submit1.php?jc=her . Authors should follow the guidelines on the Society for Human Ecology (SHE) webpage https://www.societyforhumanecology.org/instructions-for-authors . Attributions should only be made where authors have had a direct intellectual contribution in writing the manuscript.

Human Ecology Review: Volume 27, Number 2 »

Human Ecology Review: Volume 27, Number 1 »

Human Ecology Review: Volume 26, Number 2 »

Human Ecology Review: Volume 26, Number 1 »

Human Ecology Review: Volume 25, Number 2 »

Human Ecology Review: Volume 25, Number 1 »

Human Ecology Review: Volume 24, Number 2 »

Special issue: addressing the great indoors — a transdisciplinary conversation.

Human Ecology Review: Volume 24, Number 1 »

Human Ecology Review: Volume 23, Number 2 »

Special issue: human ecology—a gathering of perspectives: portraits from the past—prospects for the future.

Human Ecology Review: Volume 23, Number 1 »

Human Ecology Review: Volume 22, Number 2 »

Human Ecology Review: Volume 22, Number 1 »

Human Ecology Review: Volume 21, Number 2 »

Human Ecology Review: Volume 21, Number 1 »

Human Ecology Review: Volume 20, Number 2 »

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Human Ecology (IEG, Second edition, 2018)

2018. Human Ecology, International Encyclopedia of Geography, edited by D. Richardson, N. Castree, M. F. Goodchild, A. Kobayashi, W. Liu and R. A. Marston. doi:10.1002/9781118786352.wbieg0477.pub2. Second edition. https://doi.org/10.1002/9781118786352.wbieg0477.pub2

Related Papers

Gregory Knapp

2017.Human Ecology, in The International Encyclopedia of Geography: People, the Earth, Environment, and Technology, edited by Douglas Richardson et al., pp. 3392-3400. Wiley.

2007. “Human Ecology,” Encyclopedia of Environment and Society, edited by Paul Robbins, Vol. 3, pp. 880-884. Sage Publications, Thousand Oaks, California.

Landscape and Urban Planning

Roderick J . Lawrence

Clive Hamilton

Ecological Economics

Manfred Max-neef

Todd Lindley

This partner text with the University of North Georgia Press was created under a Round Eleven Mini-Grant for Ancillary Materials Creation and Revision. New supporting materials for the book were created under a Round Fifteen Mini-Grant and include exercises, lecture slides, and sample questions. The UNG Press also hosts a version of the text on their site: https://ung.edu/university-press/books/introduction-to-human-geography.php Authors\u27 Description: Geography is a diverse discipline that has some sort of connection to most every other academic discipline. This connection is the spatial perspective, which essentially means if a phenomenon can be mapped, it has some kind of relationship to geography. Studying the entire world is a fascinating subject, and geographical knowledge is fundamental to a competent understanding of our world. In this chapter, you will learn what geography is as well as some of the fundamental concepts that underpin the discipline. These fundamental terms...

Human Ecology

Ludomir Lozny

Edward Elgar Publishing eBooks

David Demeritt

Ben Fitzhugh

A B S T R A C T Human ecodynamics (H.E.) refers to processes of stability, resilience, and change in socio-ecological relationships or systems. H.E. research involves interdisciplinary study of the human condition as it affects and is affected by the rest of the non-human world. In this paper, we review the intellectual history of the human ecodynamics concept over the past several decades, as it has emerged out of classical ecology, anthropology, behavioral ecology, resilience theory, historical ecology, and related fields, especially with respect to the study of long-term socioecological change. Those who study human ecodynamics reject the notion that humans should be considered external to the environments in which they live and have lived for millennia. Many are interested in the resilience and sustainability of past human-natural configurations, often striving to extract lessons from the past that can benefit society today. H.E. research, involving the study of paleoenvironments and archaeology, has taken shape around a series of methodological advances that facilitate the study of past chronology, pa-leoecology, paleodemography, mobility, trade, and social networks. It is only through integrated study of 'coupled human-natural systems'—'socio-ecosystems'—that we can hope to understand dynamic human-environmental interactions and begin to manage them for sustainable goals. Local and traditional or Indigenous knowledge provides another important influence to human ecodynamics research, and we explore how such knowledge can provide both expert witness into the operation of socioecological systems and insight into the human/cultural dimensions of those systems. Ultimately, we conclude that human ecodynamics is more encompassing than a number of related approaches and can provide a nexus for productive research. Through its interdisciplinary breadth, the framework unites scholarship that tends to be more isolated to address complex problems that are best tackled with diverse perspectives.

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An Interdisciplinary Journal

Volume 50, Issue 2

Alternative conservation paradigms and ecological knowledge of small-scale artisanal fishers in a changing marine scenario in argentina.

• Daniela M. Truchet
• Belén M. Noceti
• Rocío M. Truchet

The Disconnect Between Knowledge and Perceptions: A Study of Fishermen’s Local Ecological Knowledge and Their Perception of the State of Fisheries and How These Are Managed in the Dominican Republic

• Elizabeth L. Mclean
• Graham E. Forrester
• Carlos G. García-Quijano

The Politics of Indigenous Environmental Knowledge in Vietnam

• Pamela McElwee

A Natural Disaster Framed Common Pool Resource Game Yields No Framing Effects Among Mongolian Pastoralists

• Thomas Conte

How the Enga Cope with Frost in the 21st Century: Food Insecurity, Migration, and Development in the Papua New Guinea Highlands

• Jerry K. Jacka
• Sarah Posner

Divergence in Nutritional Intake and Physical Activity Patterns Among Households in a Village of Ethnic Minorities in Northern Laos at the Initial Stage of Health Transition

• Mihoko Kibe
• Shinsuke Tomita
• Masahiro Umezaki

Eco-Symbiotic Complementarity and Trading Networks of Natural Resources in Nahua Communities in Mountain Regions of Mexico

• Elisa Lotero-Velásquez
• Eduardo García-Frapolli
• Andrea Martínez-Ballesté

A Combination of Cultural Values and Economic Benefits Promote Tolerance Towards Large Mammals in a Hotspot of Human-Wildlife Conflicts in Eastern India

• Akashdeep Roy
• Suraj Kumar Dash
• Sambandam Sathyakumar

Divergence and Convergence in Traditional Plant-Based Medicinal Practices of Haitian Migrants in Montreal, Miami and Cayenne

• M. A. Tareau
• A. Cuerrier

A Qualitative Study on How Perceptions of Environmental Changes are Linked to Migration in Morocco, Senegal, and DR Congo

• Lore Van Praag
• Samuel Lietaer
• Caroline Michellier

The Role of Religiosity on Deontological Climate Message Acceptance: A U.S. Survey

• Victoria DePalma

Ambient Air Temperature, Air Quality and the Timing of Excess Mortality Among Young Men in the United States

• Robert Bozick

Correction to: Ambient Air Temperature, Air Quality and the Timing of Excess Mortality Among Young Men in the United States

A new but gloomy picture: the first photographic evidence of grey wolf in nowshera district, khyber pakhtunkhwa, pakistan.

• Romaan Hayat Khattak
• Ejaz Ur Rehman

The Dawn of Everything

• Lauren Harding

Correction to: Land Use Changes Assessment using a triangulated framework: Perception Interviews, Land-Use/Land Cover Observation, and Spatial Planning Analysis in Tanjung Batu and Derawan Island, Indonesia

• Kevin Muhamad Lukman
• Yuta Uchiyama
• Ryo Kohsaka
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Human Ecology: A Guide to Research

Research process, pick a topic.

• Find Research Sources
• Write & Cite
• Human Ecology Department This link opens in a new window
• Get Library Help

Ask a Librarian

Librarian Liaison

Attribution.

This guide was created by a former Milne Librarian.

• In 2021, it was adapted by Jennifer Jensen with ideas from guides by Michelle Hendley, Heather Statler, and Sarah Rhodes. 

Welcome to the Human Ecology Research Guide! Use the tabs at left to navigate. This guide contains information about library resources (and how to use them) for Human Ecology students. (Image from SUNY Oneonta CANTO collection )

The Research Process

This library guide is designed to help you along each step of the research process as you prepare and write papers for your classes. For general  Research Paper Help, try this guide , too. 

Each section of the guide connects with one step of the research process:

• Pick a topic with ideas from the news, library databases, and more.
• Find Research Sources by searching for reference items, articles, books, and other library resources to help you learn about your topic.
• When you are ready to get writing, the Write & Cite page has helpful links to citation and writing/revising resources.

Pick a Topic

Looking for ideas? Here are a few ways to generate ideas for a research paper topic: 

• Consider adapting a topic you are studying for another class -- get a new perspective on it, and use your research twice
• Browse chapter titles in your textbook or unit titles in your class syllabus
• Scan newspaper headlines (see "News and Topic Databases" below) 
• Check out what's trending in social media sites you like
• Use the "Browse Issues" section in the library's Opposing Viewpoints database (see link below)
• Watch a film or documentary   and explore the film's ideas further with your research paper

News & Topic Databases

• Newspaper Source Plus Includes over 850 full-text U.S. and international newspapers as well as over 800,000 television and radio news transcripts.
• Google Newspapers Archive This link opens in a new window Provides free access to scanned archives of newspapers and links to other newspaper archives on the web. It contains more than 2,000 scanned newspapers, including The Afro American (1902-1992), The Pittsburgh Press (1819-1992), and The Village Voice (1955-2004).
• Opposing Viewpoints in Context This link opens in a new window Contains a variety of media rich content, including over 20,000 pro/con viewpoints, approximately 19,000 reference articles, interactive maps, infographics, and more. Cross-searchable with Gale In Context: Global Issues. National Debate Topics category provides quick and easy access to content on frequently studied and discussed issues. Educator Resources include a research toolkit with tips on analyzing issues with opposing viewpoints, distinguishing fact from opinion, evaluating information sources, and recognizing deceptive arguments.
• Next: Find Research Sources >>
• Last Updated: May 8, 2024 3:20 PM
• URL: https://libguides.oneonta.edu/humec

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Collection  10 March 2022

Top 100 in Ecology

This collection highlights our most downloaded* ecology papers published in 2021. Featuring authors from aroud the world, these papers showcase valuable research from an international community.

*Data obtained from SN Insights which is based on Digital Science's Dimensions.

The first true millipede—1306 legs long

• Paul E. Marek
• Bruno A. Buzatto
• Juanita Rodriguez

Lethal coalitionary attacks of chimpanzees ( Pan troglodytes troglodytes ) on gorillas ( Gorilla gorilla gorilla ) in the wild

• Lara M. Southern
• Tobias Deschner
• Simone Pika

Defining priority areas for blue whale conservation and investigating overlap with vessel traffic in Chilean Patagonia, using a fast-fitting movement model

• Luis Bedriñana-Romano
• Rodrigo Hucke-Gaete
• Daniel M. Palacios

Earliest evidence of herd-living and age segregation amongst dinosaurs

• Adriana C. Mancuso
• Vincent Fernandez

Extreme miniaturization of a new amniote vertebrate and insights into the evolution of genital size in chameleons

• Jörn Köhler
• Miguel Vences

Excess protein enabled dog domestication during severe Ice Age winters

• Maria Lahtinen
• David Clinnick
• Suvi Viranta

Honey bee hives decrease wild bee abundance, species richness, and fruit count on farms regardless of wildflower strips

• G. M. Angelella
• C. T. McCullough
• M. E. O’Rourke

Tracking late Pleistocene Neandertals on the Iberian coast

• Eduardo Mayoral
• Ignacio Díaz-Martínez
• Ricardo Díaz-Delgado

Exposure to foreign gut microbiota can facilitate rapid dietary shifts

• A. M. Fisher

Self-care tooling innovation in a disabled kea ( Nestor notabilis )

• Amalia P. M. Bastos
• Kata Horváth
• Alex H. Taylor

Discovery of a colossal slickhead (Alepocephaliformes: Alepocephalidae): an active-swimming top predator in the deep waters of Suruga Bay, Japan

• Yoshihiro Fujiwara
• Masaru Kawato
• Shinji Tsuchida

Microplastics in fish and fishmeal: an emerging environmental challenge?

• Christina J. Thiele
• Malcolm D. Hudson
• Giovanna Sidaoui-Haddad

An Italian dinosaur Lagerstätte reveals the tempo and mode of hadrosauriform body size evolution

• Alfio Alessandro Chiarenza
• Matteo Fabbri
• Federico Fanti

Seagrasses provide a novel ecosystem service by trapping marine plastics

• Anna Sanchez-Vidal
• Miquel Canals

Mistaken identity may explain why male sea snakes ( Aipysurus laevis , Elapidae, Hydrophiinae) “attack” scuba divers

• Tim P. Lynch
• Ross A. Alford
• Richard Shine

Explainable identification and mapping of trees using UAV RGB image and deep learning

• Masanori Onishi
• Takeshi Ise

The giant panda is cryptic

• Ossi Nokelainen
• Nicholas E. Scott-Samuel

Pivot burrowing of scarab beetle ( Trypoxylus dichotomus ) larva

• Haruhiko Adachi
• Makoto Ozawa
• Shigeru Kondo

Redefining the oceanic distribution of Atlantic salmon

• Audun H. Rikardsen
• David Righton
• Kim Aarestrup

Chest beats as an honest signal of body size in male mountain gorillas ( Gorilla beringei beringei )

• Edward Wright
• Sven Grawunder
• Martha M. Robbins

Powered flight in hatchling pterosaurs: evidence from wing form and bone strength

• Darren Naish
• Mark P. Witton
• Elizabeth Martin-Silverstone

Two wild female bonobos adopted infants from a different social group at Wamba

• Nahoko Tokuyama
• Kazuya Toda
• Shintaro Ishizuka

Multiple pygmy blue whale acoustic populations in the Indian Ocean: whale song identifies a possible new population

• Emmanuelle C. Leroy
• Jean-Yves Royer
• Tracey L. Rogers

Action potentials induce biomagnetic fields in carnivorous Venus flytrap plants

• Anne Fabricant
• Geoffrey Z. Iwata
• Dmitry Budker

North Pacific warming shifts the juvenile range of a marine apex predator

• Kisei R. Tanaka
• Kyle S. Van Houtan
• Salvador J. Jorgensen

Spatial and temporal pattern of wildfires in California from 2000 to 2019

• Tirtha Banerjee

Effects of climate change and land cover on the distributions of a critical tree family in the Philippines

• Sean E. H. Pang
• Jose Don T. De Alban
• Edward L. Webb

African soil properties and nutrients mapped at 30 m spatial resolution using two-scale ensemble machine learning

• Tomislav Hengl
• Matthew A. E. Miller
• Jonathan Crouch

Russian forest sequesters substantially more carbon than previously reported

• Dmitry Schepaschenko
• Elena Moltchanova
• Florian Kraxner

Isotope data from amino acids indicate Darwin’s ground sloth was not an herbivore

• Julia V. Tejada
• John J. Flynn
• Brian N. Popp

Late Pleistocene human paleoecology in the highland savanna ecosystem of mainland Southeast Asia

• Kantapon Suraprasit
• Rasmi Shoocongdej
• Hervé Bocherens

Early Pleistocene faunivorous hominins were not kleptoparasitic, and this impacted the evolution of human anatomy and socio-ecology

• Manuel Domínguez-Rodrigo
• Enrique Baquedano
• Diego González-Aguilera

An ecological niche shift for Neanderthal populations in Western Europe 70,000 years ago

• William E. Banks
• Marie-Hélène Moncel
• Frédéric Santos

Surface slicks are pelagic nurseries for diverse ocean fauna

• Jonathan L. Whitney
• Jamison M. Gove
• Gregory P. Asner

Field measurements of a massive Porites coral at Goolboodi (Orpheus Island), Great Barrier Reef

• Nathan Cook
• Vicki Saylor

Ongoing ecological and evolutionary consequences by the presence of transgenes in a wild cotton population

• Valeria Vázquez-Barrios
• Karina Boege

Novel and disappearing climates in the global surface ocean from 1800 to 2100

• Katie E. Lotterhos
• Áki J. Láruson
• Li-Qing Jiang

Partial shading by solar panels delays bloom, increases floral abundance during the late-season for pollinators in a dryland, agrivoltaic ecosystem

• Maggie Graham
• Serkan Ates
• Chad W. Higgins

The early hunting dog from Dmanisi with comments on the social behaviour in Canidae and hominins

• Saverio Bartolini-Lucenti
• Joan Madurell-Malapeira
• Lorenzo Rook

Comprehensive comparative morphology and developmental staging of final instar larvae toward metamorphosis in the insect order Odonata

• Genta Okude
• Takema Fukatsu
• Ryo Futahashi

G enome-wide data implicate terminal fusion automixis in king cobra facultative parthenogenesis

• Daren C. Card
• Freek J. Vonk
• Warren Booth

New evidence from exceptionally “well-preserved” specimens sheds light on the structure of the ammonite brachial crown

• C. P. A. Smith
• N. H. Landman

Population decline in a ground-nesting solitary squash bee ( Eucera pruinosa ) following exposure to a neonicotinoid insecticide treated crop ( Cucurbita pepo )

• D. Susan Willis Chan
• Nigel E. Raine

Tropical deforestation induces thresholds of reproductive viability and habitat suitability in Earth’s largest eagles

• Everton B. P. Miranda
• Carlos A. Peres
• Colleen T. Downs

Biodiversity and ecosystem functions depend on environmental conditions and resources rather than the geodiversity of a tropical biodiversity hotspot

• Christine I. B. Wallis
• Yvonne C. Tiede
• Jörg Bendix

Summer weather conditions influence winter survival of honey bees ( Apis mellifera ) in the northeastern United States

• Martina Calovi
• Christina M. Grozinger
• Sarah C. Goslee

Evidence that spillover from Marine Protected Areas benefits the spiny lobster ( Panulirus interruptus ) fishery in southern California

• Hunter S. Lenihan
• Jordan P. Gallagher
• Daniel C. Reed

Metal concentrations in coastal sharks from The Bahamas with a focus on the Caribbean Reef shark

• Oliver N. Shipley
• Cheng-Shiuan Lee
• Austin J. Gallagher

First tracks of newborn straight-tusked elephants ( Palaeoloxodon antiquus )

• Carlos Neto de Carvalho
• Zain Belaústegui
• Clive Finlayson

An integrative analysis uncovers a new, pseudo-cryptic species of Amazonian marmoset (Primates: Callitrichidae: Mico ) from the arc of deforestation

• Rodrigo Costa-Araújo
• José S. Silva-Jr.
• Tomas Hrbek

eDNA sampled from stream networks correlates with camera trap detection rates of terrestrial mammals

• Arnaud Lyet
• Loïc Pellissier
• Robin Naidoo

Large-scale variations in the dynamics of Amazon forest canopy gaps from airborne lidar data and opportunities for tree mortality estimates

• Ricardo Dalagnol
• Fabien H. Wagner
• Luiz E. O. C. Aragão

Tipping point realized in cod fishery

• Christian Möllmann
• Xochitl Cormon
• Martin Quaas

Temporal activity patterns suggesting niche partitioning of sympatric carnivores in Borneo, Malaysia

• Miyabi Nakabayashi
• Tomoko Kanamori

No evidence of physiological declines with age in an extremely long-lived fish

• Derek J. Sauer
• Britt J. Heidinger
• Mark E. Clark

International fisheries threaten globally endangered sharks in the Eastern Tropical Pacific Ocean: the case of the Fu Yuan Yu Leng 999 reefer vessel seized within the Galápagos Marine Reserve

• Elisa Bonaccorso
• Nicté Ordóñez-Garza
• Juan M. Guayasamin

Indirect effects of invasive rat removal result in recovery of island rocky intertidal community structure

• Carolyn M. Kurle
• Kelly M. Zilliacus
• Donald A. Croll

Assessing the carbon capture potential of a reforestation project

• David Lefebvre
• Adrian G. Williams

The evolutionary history of manatees told by their mitogenomes

• Érica Martinha Silva de Souza
• Lucas Freitas
• Mariana Freitas Nery

Microplastics accumulate fungal pathogens in terrestrial ecosystems

• Gerasimos Gkoutselis
• Stephan Rohrbach
• Gerhard Rambold

Advances in automatic identification of flying insects using optical sensors and machine learning

• Carsten Kirkeby
• Klas Rydhmer
• Kaare Græsbøll

Aboveground plant-to-plant communication reduces root nodule symbiosis and soil nutrient concentrations

• Yuta Takahashi
• Kaori Shiojiri
• Akira Yamawo

The rhizosphere microbiome plays a role in the resistance to soil-borne pathogens and nutrient uptake of strawberry cultivars under field conditions

• Cristina Lazcano
• Kelly Ivors

Highest risk abandoned, lost and discarded fishing gear

• Eric Gilman
• Michael Musyl
• Brandon Kuczenski

Future thermal regimes for epaulette sharks ( Hemiscyllium ocellatum ): growth and metabolic performance cease to be optimal

• Carolyn R. Wheeler
• Jodie L. Rummer
• John W. Mandelman

Earliest evidence of marine habitat use by mammals

• Anton F.-J. Wroblewski
• Bonnie E. Gulas-Wroblewski

Constraining the chronology and ecology of Late Acheulean and Middle Palaeolithic occupations at the margins of the monsoon

• James Blinkhorn
• Hema Achyuthan
• Jana Ilgner

Neonicotinoids disrupt memory, circadian behaviour and sleep

• Kiah Tasman
• Sergio Hidalgo
• James J. L. Hodge

Evaluation metrics and validation of presence-only species distribution models based on distributional maps with varying coverage

• Kamil Konowalik
• Agata Nosol

Ecosystem response persists after a prolonged marine heatwave

• Robert M. Suryan
• Mayumi L. Arimitsu
• Stephani G. Zador

Rhizobia use a pathogenic-like effector to hijack leguminous nodulation signalling

• Safirah Tasa Nerves Ratu
• Albin Teulet
• Shin Okazaki

Camera trap placement for evaluating species richness, abundance, and activity

• Kamakshi S. Tanwar
• Yadvendradev V. Jhala

Assessing biophysical and socio-economic impacts of climate change on regional avian biodiversity

• Simon Kapitza
• Pham Van Ha
• Brendan A. Wintle

Smell of green leaf volatiles attracts white storks to freshly cut meadows

• Martin Wikelski
• Michael Quetting
• Jonathan Williams

Dinosaur senescence: a hadrosauroid with age-related diseases brings a new perspective of “old” dinosaurs

• Justyna Słowiak
• Tomasz Szczygielski
• Dawid Surmik

Positive effects of COVID-19 lockdown on river water quality: evidence from River Damodar, India

• Baisakhi Chakraborty
• Biswajit Bera
• Pravat Kumar Shit

Snow algae blooms are beneficial for microinvertebrates assemblages (Tardigrada and Rotifera) on seasonal snow patches in Japan

• Nozomu Takeuchi
• Krzysztof Zawierucha

Coastal wetlands can be saved from sea level rise by recreating past tidal regimes

• Mahmood Sadat-Noori
• Caleb Rankin
• William Glamore

Identifying unknown Indian wolves by their distinctive howls: its potential as a non-invasive survey method

• Sougata Sadhukhan
• Holly Root-Gutteridge
• Bilal Habib

Optimal fishing effort benefits fisheries and conservation

• Emma V. Sheehan
• Martin J. Attrill

The spreading of the invasive sacred ibis in Italy

• Marco Cucco
• Gianfranco Alessandria
• Irene Pellegrino

Exceptional fossil assemblages confirm the existence of complex Early Triassic ecosystems during the early Spathian

• Christopher P. A. Smith
• Thomas Laville
• Arnaud Brayard

Allelopathic effect of Artemisia argyi on the germination and growth of various weeds

Organophosphate poisoning of Hyacinth Macaws in the Southern Pantanal, Brazil

• Eliane C. Vicente
• Neiva M. R. Guedes

The critical role of natural history museums in advancing eDNA for biodiversity studies: a case study with Amazonian fishes

• C. David de Santana
• Lynne R. Parenti

Novel approach to enhance coastal habitat and biotope mapping with drone aerial imagery analysis

• João Gama Monteiro
• Jesús L. Jiménez
• João Canning-Clode

High resolution biologging of breaching by the world’s second largest shark species

• Jessica L. Rudd
• Owen M. Exeter
• Lucy A. Hawkes

The largest hoplophonine and a complex new hypothesis of nimravid evolution

• Paul Zachary Barrett

Dangerous demographics in post-bleach corals reveal boom-bust versus protracted declines

• Juliano Morais
• Renato A. Morais
• David R. Bellwood

Welwitschia : Phylogeography of a living fossil, diversified within a desert refuge

• Norbert Jürgens
• Imke Oncken
• Barbara Rudolph

Response of bacterial and fungal communities to high petroleum pollution in different soils

• Polina Galitskaya
• Liliya Biktasheva
• Svetlana Selivanovskaya

Integrating multiple chemical tracers to elucidate the diet and habitat of Cookiecutter Sharks

• Aaron B. Carlisle
• Elizabeth Andruszkiewicz Allan
• John O’Sullivan

Environmental stressors, complex interactions and marine benthic communities’ responses

• Charlotte Carrier-Belleau
• David Drolet
• Philippe Archambault

eDNA metabarcoding for biodiversity assessment, generalist predators as sampling assistants

• Louise Nørgaard
• Carsten Riis Olesen
• Laura Iacolina

Distance sampling surveys reveal 17 million vertebrates directly killed by the 2020’s wildfires in the Pantanal, Brazil

• Walfrido Moraes Tomas
• Christian Niel Berlinck
• Ronaldo Morato

Description of five new species of the Madagascan flagship plant genus Ravenala (Strelitziaceae)

• Thomas Haevermans
• Annette Hladik
• Patrick Blanc

A symbiotic aphid selfishly manipulates attending ants via dopamine in honeydew

• Tatsumi Kudo
• Hitoshi Aonuma
• Eisuke Hasegawa

Ixodiphagus hookeri wasps (Hymenoptera: Encyrtidae) in two sympatric tick species Ixodes ricinus and Haemaphysalis concinna (Ixodida: Ixodidae) in the Slovak Karst (Slovakia): ecological and biological considerations

• Alicja Buczek
• Weronika Buczek
• Michał Stanko

Genomic signatures of the evolution of defence against its natural enemies in the poisonous and medicinal plant Datura stramonium (Solanaceae)

• I. M. De-la-Cruz
• J. Núñez-Farfán

Timely poacher detection and localization using sentinel animal movement

• Henrik J. de Knegt
• Jasper A. J. Eikelboom
• Herbert H. T. Prins

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• Philos Trans R Soc Lond B Biol Sci
• v.378(1873); March 27, 2023

Infectious disease ecology and evolution in a changing world

Kayla c. king.

1 Department of Biology, University of Oxford, Oxford OX1 3SZ, UK

Matthew D. Hall

2 School of Biological Sciences, Monash University, Melbourne 3800, Australia

Justyna Wolinska

3 Department of Evolutionary and Integrative Ecology, Leibniz Institute of Freshwater Ecology and Inland Fisheries (IGB), 12587 Berlin, Germany

4 Department of Biology, Chemistry, and Pharmacy, Institute of Biology, Freie Universität Berlin (FU), 14195 Berlin, Germany

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This article has no additional data.

1.  Introduction

Managing the consequences of climate change and human activity is one of the greatest challenges of the twenty-first century. Extreme weather events, such as heatwaves and droughts, are becoming more common and more severe [ 1 ], and urban and agricultural expansion is contributing to the loss of biodiversity and the degradation of ecosystems [ 2 ]. This new reality challenges the capacity of host species to persist and forces infectious diseases to rapidly evolve. Indeed, the COVID-19 pandemic has emphasized how quickly infectious diseases can evolve and spread—with consequences for transmission, virulence and evasion of host defences [ 3 ]—and that disease dynamics will play out differently across regions of the globe [ 4 , 5 ].

Addressing these challenges requires multi-faceted approaches that explore the effects of human-induced change on host–pathogen (including parasites) interactions across space and time (i.e. ecological and evolutionary timescales). Developing measures to enhance the resilience of natural and agricultural communities to these changes is also critical. Yet, these concepts have traditionally been tackled from disparate viewpoints with little empirical overlap. Ecological and epidemiological research, for example, has linked the spread of disease to environmental factors (e.g. [ 6 – 8 ]). But there is a pressing need for evolutionary research to capture how hosts and their pathogens may evolve under the sweeping environmental changes that populations now face. There is evidence that abiotic environmental factors can impact selection in host–pathogen interactions (e.g. [ 9 ]), but often empirical work examines these questions in abstract ways (hot versus cold, low versus high food). In turn, suggestions to use ecological and evolutionary principles in the management of agricultural pests and pathogens do not often consider the social and economic factors that underlie any long-term intervention [ 10 ].

This special issue aims to bring together research from different fields (ecological, evolutionary, epidemiological and applied) and approaches to better understand and address the impact of human-driven environmental change on infectious disease. It addresses a lack of comprehensive discussion on key issues that arise because different types of environmental change often form their own fields of research, rather than being studied as interconnected symptoms of human activity. To address this gap, the contributions herein focus on three key themes: climate change and infection outcomes ; understanding host–pathogen interactions in dynamic environments ; and outbreaks and pathogen evolution in human-altered ecosystems . By comparing different forms of global change, integrating across multiple fields, and identifying empirical and theoretical research gaps, this issue's goal is to showcase and spark new thinking on infectious disease evolution in a rapidly changing world.

2.  Theme 1: climate change and infection outcomes

Thermal conditions can strongly impact host- and pathogen infection-related traits. Temperature has been shown to alter the encounter rates between host and parasites, host susceptibility and tolerance to infection and, finally, the infectivity and virulence of parasites (reviewed in [ 11 – 14 ]). There is a common prediction of increased pathogen development and replication rates as well as enhanced parasite transmission under elevated temperatures (i.e. ‘warmer hence sicker world’ scenario, e.g. [ 12 ]). However, the empirical evidence is inconclusive—certain studies support but others contradict (e.g. [ 15 – 17 ]) this hypothesis.

The first group of papers focus on the most widely considered facet of global change—climate change and rising temperatures—but address the pressing need to consider these in the light of host and pathogen traits as well as their spread and evolutionary potential. Empirical studies explore the direct impacts of warming on key pathogen infectivity and host resistance traits, with implications for disease outbreaks. A field study extends exploration of trait variation in the larger host community to disease risk as temperature increases. An empirical study and an opinion piece further consider the roles host and pathogen adaptation play in climate-driven disease emergence and severity. Whether hosts can shift their thermal tolerance, or pathogens evolve to become more virulent, has major implications for species persistence in a warming world.

To better understand the impact of environmental temperatures on disease outbreaks, two papers focus on changes in mechanisms underpinning the ability of parasites to infect [ 18 ] and of hosts to resist [ 19 ]. In effect, these papers take care to start from the beginning, where the host and parasite first meet, using the planktonic crustacean Daphnia and its bacterial parasites. Marcus et al . [ 18 ] found that parasite spores exposed to higher temperatures were impaired in their ability to attach to hosts and subsequently establish infection. The degree of impact on these traits also varied depending on whether spores were desiccated or kept wet during heat spells. It is not the case that a warmer world is sicker if parasite survival is at risk between hosts. Like most hosts, waterfleas have physical barriers to infection and also have cellular immune responses. In detailed experiments, Sun et al . [ 19 ] found that warming affected these defence traits differently, with consequences for infection outcomes. Disease spread can also depend on how temperature interacts with traits in host communities [ 20 , 21 ]. In a thorough field study set in the Swiss Alps, Halliday et al . [ 22 ] found– complex relationships between trait variation in plant communities driven by temperature gradients and prevalence of infection in those communities. Together, these three studies highlight that to predict disease dynamics under climate change, we should consider multiple traits at the interface of host–pathogen interactions and their variation within species and communities.

The final two papers in this theme address the evolutionary potential of hosts [ 23 ] and pathogens [ 24 ] in the face of rising temperatures. The geographical ranges of hosts, and the pathogens they carry, are expanding owing to changing climates [ 25 ]. Of particular concern is the future impact of climate change on the potential for mosquitoes—vectors for many human disease-causing viruses—to adapt to shifts in their thermal environment and move into new areas. Ware-Gilmore et al . [ 23 ] thoughtfully address this issue by studying how the heritable genetic variation and physiological responses in the mosquito Aedes aegypti may affect the upper thermal limits in populations over evolutionary time. In addition to driving vector evolution, will climate change select for more virulent pathogens? In the light of the classic virulence–transmission trade-off [ 26 ], Hector et al . [ 24 ] discuss what happens to pathogen virulence, burdens, and replication in host populations suffering heat stress. The authors find predicting the evolution of pathogen virulence amidst climate change might require a better understanding of transmission strategies and covariation among pathogen traits.

3.  Theme 2: understanding host–pathogen interactions in dynamic environments

The study of abrupt environmental changes, such as hot versus cold or pristine versus polluted (e.g. see [ 9 , 27 , 28 ]) is often used to interpret the ecological or evolutionary impact of global change. However, the varied and dynamic nature of human-induced change is highly realistic. The simplification of environmental change is understandable given the added sample sizes that studying multiple host or pathogen genotypes or species necessitates. Yet, this situation is rapidly changing. For the study of thermal change, for example, a range of temperatures are increasingly being used to study thermal performance curves [ 8 , 29 ] or even the variation in daily thermal fluctuations or heatwaves [ 30 ]. New theoretical and empirical approaches, however, are now needed to expand our capacity to quantify the dynamics of environmental change and predict host and pathogen evolution in nature.

The first two papers in this theme evaluate how well current modelling approaches perform in capturing the rapidly changing environments that host and pathogens face. Best & Ashby [ 31 ] review the main approaches used to model host–pathogen evolution when ecological dynamics fluctuate owing to either extrinsic (seasonality, food availability) or intrinsic (time lags) factors. They then provide an in-depth guide on how to implement one main method and apply this approach to fluctuations arising from seasonally varying resources, among others. By contrast, Jiranek et al. [ 32 ] review the use of mechanistic models to study host–pathogen interactions under different scenarios of climate change, with a focus on plant systems. They outline the challenge of linking disease outbreaks with weather variables when climate change will likely affect many aspects of host and pathogen physiology, host demography, and pathogen life cycles, and these effects may frequently be nonlinear. The authors then discuss how mechanistic models overcome this limitation. These models can leverage data from wild and agricultural plant–pathogen systems to understand the complex feedback loops arising among physiological, demographic and evolutionary processes.

Complementing the modelling-focused perspectives are two empirical papers exploring how local environmental conditions predict disease prevalence, severity and evolution. Graham et al. [ 33 ] highlight the utility of using high-throughput phenotyping to make disease comparisons across large environmental, spatial and temporal gradients. By surveying seagrass wasting disease in eelgrass meadows throughout their northern range (covering eight degrees of latitude) they show that disease prevalence and severity was lower in cooler sites, colder years, and higher latitudes. The authors provide several suggestions for how this new information can improve eelgrass management. The final paper in this theme, by Melero et al. [ 34 ], considers how climate change and human activity might induce changes in plant development that can shape the evolution of host–virus interactions. Using Arabidopsis thaliana as a model system, the authors experimentally evolved a turnip mosaic virus at three different host developmental stages. They found hosts in later developmental stages were prone to faster and more severe infections, but the virus nonetheless evolved more rapidly in younger hosts.

4.  Theme 3: outbreaks and pathogen evolution in human-altered ecosystems

The third group of papers considers the broader extent of direct human activity on host–pathogen interactions. Synthesis and empirical papers explore the socio-economic challenges of implementing evolutionarily responsive practices in agriculture, and the consequences for pathogen evolution when an intervention, such as well-meaning habitat restoration, goes wrong. Geffersa et al . [ 35 ] summarize research efforts on crop disease management based on deployment of resistance genes. The goal is to disrupt pathogen adaptation and prevent the breakdown of resistance. However, practical uptake of such strategies is limited, and applied evolutionary research to control pathogen adaptation can have socio-economic challenges. Geffersa et al . develop a conceptual framework for the economic valuation of engineering of genes conferring resistance, emphasizing the value of these strategies beyond economic benefits. Feau et al . [ 36 ] argue that the introduction of new host species can accelerate pathogen evolution and affect long-established host–pathogen coevolutionary dynamics. Specifically, the emergence of a new pathogen lineage with the intensification of poplar tree cultivation causes stem infections in a new host. This represents a serious threat to poplars and could affect both natural and planted forests. Finally, Manley et al . [ 37 ] explore how conservation measures to protect pollinators—planting wildflowers along fields—affect disease prevalence in pollinator communities. They found wildflower patches did not act as transmission hubs, but reduced the prevalence of some viral infections, playing an unintended but additional role in pollinator conservation.

Complementing these papers is a novel empirical exploration of the consequences of emerging pollutants, namely nanoplastics [ 38 ] and pharmaceuticals [ 39 ]. Unlike other well-studied pollutants, wastewater treatment is often inadequate in removing nanoplastics and pharmaceuticals. These pollutants will thus remain a problem in the coming decades. Plastic production is estimated to reach 33 billion tons by 2050 [ 40 ] and is particularly insidious because plastics break down into smaller particles called micro- or nanoplastics (size less than 5 mm and less than 100 nm, respectively) that can cross cell membranes, penetrate organs and bioaccumulate in organisms. In the current issue, Manzi et al . [ 38 ] and Aulsebrook et al . [ 39 ] both used the planktonic crustacean Daphnia as experimental host. Manzi et al . found that it depends on the type of parasite whether and to what extent nanoplastics affect the infection. Low plastic load had no direct negative consequences for the host, but infection rates either greatly increased ( Metschnikowia bicuspidata ) or were impeded ( Ordospora colligata ). These results indicate that distinct parasite species can show contradictory responses to a contaminant and that nanoplastics can favour co-infections.

Thousands of pharmaceuticals are used for health management in humans, pets and agricultural animals, and over 600 products have now been detected in the wild [ 41 ]. These pollutants remain bioactive when excreted, are often resistant to degradation, and target receptors conserved in many species [ 42 – 44 ]. Aulsebrook et al . [ 39 ] showed that the non-monotonic effects of fluoxetine were only expressed once a host was infected, demonstrating that the full impact of pharmaceuticals may only be experienced in the presence of other stressors. Parameterizing an epidemiological model, they further explore how fluoxetine can shape the likelihood of an infectious disease outbreak. Their result reiterates the findings of Manzi et al . [ 38 ] that the effects of pollutants are likely to be pathogen species or genotype specific. Both pollutants exemplify the complexity of modern human activity on disease dynamics by acting in a nonlinear (non-monotonic) manner and causing an unexpected exaggeration of infection outcomes at trace amounts.

5.  Future directions and conclusion

The consequences of global change to infectious disease ecology and evolution are relevant for the health of humans, animals, plants and the environment. In this issue, climate change, environmental pollution and the increasing movement of people, animals or cultivars, are presented as examples of human-induced change that can affect the emergence and evolution of hosts, pathogens and vectors. Beyond public or ecosystem health concerns, pathogen spread and evolution due to global change have also been presented as intimately linked to issues of food security—crop production and other aspects of agriculture (including aquaculture and apiculture). Arising from the series of papers across the three themes of this special issue are these takeaways:

• 1. We need to know more about the potential and realized evolutionary paths of hosts and pathogens in a human-altered world. Both host and pathogen responses to climate change, emerging pollutants, or even interventions, are likely to be species or genotype specific. Pathogens and many pathogen-carrying invertebrates can also evolve on short timescales that are relevant for predicting disease outbreaks, as well as the likelihood and impact of zoonoses.
• 2. A warmer or heavily modified world is not always sicker. Temperature affects each component of host–pathogen interactions in unique ways, from host demography to within-host pathogen burden, making simple generalizations difficult. While traditional pollution at a greater dose is usually more damaging for hosts or pathogens, for emerging pollutants (i.e. pharmaceuticals), the greatest effects can often occur at the smallest doses, owing to the way these chemicals target conserved pathways.
• 3. Not all populations will be equally impacted by change. Host and pathogen populations should be expected to be adapted to their local environment, and disease outcomes will likely vary with latitude, altitude, water depth (in the case of aquatic organisms), or prior exposure to human activity. Local adaptation is key both to understanding how host and pathogen responses to human activity might vary over space and time, and for making predictions on the future distribution of vectors and pathogens under different change scenarios.
• 4. Opportunities to expand the scope with which human-induced change is studied are sorely needed. Directly incorporating fluctuating ecological dynamics into our studies or using empirical data to build mechanistic models of different types of change offer some solutions for predicting evolutionary change in response to human modification.
• 5. Parasitism is one of the most common lifestyles on Earth [ 45 ]. There is a need to incorporate interactions between host species and their pathogens as new ecotoxicological endpoints to better assess the ecological consequences of novel pollutants. Assessing the effects of any pollutant in isolation and, in particular, dismissing infection may lead to a severe underestimation of their real impact on individual host physiology, with upscaling effects on overall populations and ecosystems.
• 6. Adapting evolutionary or ecological principles into the management of human activity, pests or pathogens is not without its costs. The longer timescales with which these implementations operate, particularly when compared with traditional agricultural approaches, for example, create additional social and economic challenges that are often not appreciated when eco–evo ideas are first suggested.

Overall, this issue summarizes current progress and identifies remaining gaps in our understanding of infectious disease ecology and evolution in a global change framework. We hope the special issue will help drive new research on host–pathogen interactions, integrating traditionally isolated fields of study.

Acknowledgements

We thank all lead authors and their collaborators who agreed to be part of this special issue. We are also grateful to Helen Eaton and the editorial team at Phil. Trans. R. Soc. B , who patiently and efficiently managed the special issue, and the inevitable delays, over the last year.

Data accessibility

Authors' contributions.

K.C.K.: writing—original draft, writing—review and editing; M.D.H.: writing—original draft, writing—review and editing; J.W.: writing—original draft, writing—review and editing.

All authors gave final approval for publication and agreed to be held accountable for the work performed herein.

Conflict of interest declaration

We declare we have no competing interests.

We received no funding for this study.

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Human Ecology Research Paper

View sample sociology research paper on human ecology. Browse other research paper examples for more inspiration. If you need a thorough research paper written according to all the academic standards, you can always turn to our experienced writers for help. This is how your paper can get an A! Feel free to contact our custom writing service for professional assistance. We offer high-quality assignments for reasonable rates.

Ecology is defined as “the science of the economy of animals and plants; that branch of biology which deals with the relations of living organisms to their surroundings, their habits and modes of life, etc.” (Oxford English Dictionary 2000). The first half of that definition implies an impulse toward efficient interdependence. Applied to humans, the second half of that definition implies much more than what is usually meant by the term human ecology, however. A consideration of the relations of individuals to their surroundings, their habits, and modes of life would include almost all aspects of what are now seen as “contextual effects.” It would include some aspects of developmental psychology (e.g., Barker and Wright 1954) and much of environmental psychology (Bechtel 1997). These are part of what is sometimes called autoecology (treating organisms individually as opposed to synecology, the study of collectivities), a field of inquiry that is perhaps part of ecology but not what is usually included under the term. That definition would also include studies, such as Gerth and Mills’s (1953) Social Structure and Character, among many others, that examine how a changing social structure affects the mode of life of a particular population. In contrast, sociological human ecology has been most concerned with investigating what are now termed aggregation effects, that is, how people, living together, create social organization. A reciprocal relationship between an individual or collectivity and its environment is posited. Human ecology is, therefore, concerned with the nature and process of community development (Hawley 1950).

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Get 10% off with 24start discount code, a brief outline of the early development of human ecology.

The term ecology was coined by the German botanist Ernst Haeckel in 1868, reviving much older concerns. The term human ecology may have been first used by J. Paul Goode, a University of Chicago geographer, at the 1907 meeting of the Association of American Geographers in describing an introductory university course. By 1920, Barrington Moore (1920), president of the Ecological Society of America, characterized geography as “human ecology” in his presidential address, and the journal Ecology reviewed Ellsworth Huntington and Sumner W. Cushing’s Principles of Human Geography under the title “Human Ecology” the following year. In 1922, Harlan Barrows, concerned about the continuing relevance of the field as the era of discovery neared its end, accepted the attribution as the new and future identity for geography in his presidential address to the Association of American Geographers. Anthropologists showed an interest in the relation of their field to ecology as early as 1903, but the term human ecology does not appear to have been used by them until the 1930s. Today, human ecological theory may be most intensively discussed by anthropologists.

Robert Park’s 1915 essay “The City: Suggestions for the Investigation of Human Behavior in the City Environment” is sometimes thought to be the origin of ecological thought in sociology, but neither does Park mentions the term nor are the outlines of ecological theory visible. (The 1925 republication of that essay includes a substantially rewritten introduction including the term.) Ecology does not play a large role in his 1921 essay on “Sociology and the Social Sciences” either. That three-part essay roots sociology firmly in history, links collective phenomena to interaction, and outlines a program for research. Park did discuss human ecology in his 1925 presidential address to the American Sociological Society, but it was not until 1936 that Park addressed ecological theory systematically. Nevertheless, Hawley (1950:8) attributed the origin of human ecology in sociology to pages 161 to 216 in Park and Burgess’s ([1921] 1969) Introduction to the Science of Sociology.

Park and Burgess’s ([1921] 1969) monumental 1,000page text is a comprehensive attempt at understanding social phenomena. Beginning with a discussion of the place of sociology among the social sciences (including the 1921 journal essay) and ending with a treatise on the nature of progress, the argument builds from human nature to society and the group. Connections are drawn between social contact and isolation, interaction and social forces, and four types of social relationships— competition, conflict, accommodation, and assimilation—to arrive at the forms of social control and collective action. Competition for ultimately limited resources and the connections engendered played a central role in the argument laid out in the text. These were classified as presocial because competition does not necessarily entail contact. Conflict implies a mutual awareness of the competing parties. Accommodation implies patterned adjustments of behavior, while assimilation implies a despecialization in behavioral patterns and a dissolution of social demarcation.

The range of issues and thought considered by Park and Burgess is impressive. A total of more than 180 collected edited contributions representing the work of more than 100 authors were introduced, reworked, and discussed by the authors. Park authored 14 of the included readings. Simmel was responsible for 10. Charles Darwin, Albion Small, and William Graham Sumner each had four readings included. Five authors, including John Dewey and W. I. Thomas, had three. Durkheim, Smith, Spencer, Le Bon, and 11 others each had two pieces, and 97 authors, including Comte, Cooley, Galton, Giddings, Gras, William James, Rousseau, Santayana, Schopenhauer, Sombart, Toennies, and Lester Ward, each made one contribution to the book. The large number of topics and breadth of authors suggests the diversity of intellectual sources that early American sociology drew on.

Despite the discussion of ecological concepts in one of the chapters of that work, a systematic discussion of human ecology was left to Roderick McKenzie in his 1924 essay “The Ecological Approach to the Study of the Human Community” and his 1927 article on “The Concept of Dominance and World-Organization.” The outlines of contemporary human ecological theory become visible in these two works. In the first work, McKenzie stated the basic claim that competition, selection, and accommodation determine the size and organization of human communities before identifying four types of communities, enumerating the basic factors behind community growth and decline, specifying the relationship between resources, population, and organization, and discussing the relationship between the basic ecological mechanisms and the internal structure of communities. In the second work, McKenzie provided a broad road map to the development of Asia over the seven or eight decades since the work appeared. He discussed the relationship between transportation technology and European colonial expansion, pointing out that the separation of communication from transportation has permitted the concentration of control as it allowed the decentralization of operations, and he outlined the changes in patterns of production and trade brought about by the expansion process. The application of ecological theory to the world system was refined and expanded in a subsequent book chapter (McKenzie 1933). The outlines of contemporary human ecological methodology become visible in his 1933 book, The Metropolitan Community, where social demography, geographic mapping, and summary analysis of spatial movements were used to support arguments about social organization (McKenzie [1933] 1997). The spatial implications of the process of ecological succession on intra-urban residential patterns, in the form of the concentric zone model, was laid out by Burgess in 1925. This work, so well known to students of urban development today, did not figure prominently in the then contemporary theoretical debates.

Each of the three main founders of sociological human ecology was a part-time theorist. Park, who came to Chicago as a part-time, practice-track lecturer in 1914 from the Tuskegee Institute and left in 1932 for Fisk University devoted much of his time to studying the role of newspapers in what is now sometimes called the “public sphere.” Burgess showed more sustained interest in the family than in the city. McKenzie, who apparently developed an interest in sociology after first becoming a classicist of some accomplishment, had his productive life cut short by illness. There was remarkably little systematic theoretical work advancing sociological human ecology before 1950. Human ecology was perhaps more extensively defined by its critics (e.g., Alihan [1938] 1964; Firey 1947) than by its proponents.

Ecological ideas were “in the air,” however. E. Warming’s Oecology of Plants appeared in 1909, Frederic Clements’s Plant Succession in 1916. Both extended the work of Charles Darwin. Frederic Le Play’s research on the relationship between family structure and the social environment in the mid-nineteenth century was known (Zimmerman and Frampton 1935). Galpin (1915) performed research on rural areas that had an impact on ecologically minded sociologists. J. Arthur Thomson, a biologist, and Patrick Geddes, one of the founders of modern city planning (Geddes 1915), collaborated on a booklength work of human ecology (1931) that extended earlier work. Herbert G. Wells, Julian Huxley, and George P. Wells published a three-volume comprehensive overview of ecology, with much of the last volume being devoted to human ecology, in 1934, which followed H. G. Wells’s major survey of history (1921). J. W. Bews (1935), a botanist, offered a statement of human ecology that was extensively reviewed. The work of N. S. B. Gras (1922) and Adna Weber (1899) had an impact on the theorizing in human ecology. This was an era of sweeping generalization and the “big work,” but there were many big ideas current at the time, and comparisons between humans and other species was only one of them.

Amos Hawley produced the first book-length statement of human ecology in 1950 and the first one that provided a systematic treatment of the theory, integrating social mechanisms and morphological outcomes. This has become the central statement of ecological theory. Hawley gave new prominence to the intricacies of human-commensal relations (discussed by Park and Burgess ([1921] 1969) but subsequently downplayed). That statement gave human ecology its contemporary form by codifying and extending the ecological thought of McKenzie and illustrating the resulting theory with the type of data and application in McKenzie’s Metropolitan Community. Shortly before the appearance of Hawley’s book, Donald Bogue published a study of metropolitan dominance (1949). Otis Dudley Duncan (1951) examined the question of optimal city size. Duncan (1959, 1964) later codified the thinking about the ecological complex and developed ecological theory and methods to study inter-urban dependences and patterns of development (Duncan et al. 1960; see also Vance and Smith 1954). Until about 1950, the illustrations used in ecological theorizing were not particularly urban in orientation even if all the protagonists had long-standing interests in cities as a social formation. Countering the urban preoccupation, Gibbs and Martin (1959) argued that the proper focus of human ecology was sustenance (subsistence) organization. Hawley’s (1986) refined statement of ecological theory de-emphasized spatially delimited communities. Much of the recent theoretical advance has been in applying the ecological approach to studies of formal organizations (Bidwell and Kasarda 1985), industry structure and entrepreneurship (Carroll and Hannan 2000), cultural institutions (Blau 1989; Wuthnow 1987), and voluntary organizations (McPherson 1983).

This review is too narrow, however. While human ecology is not always well integrated into the theory, since the 1920s and continuing into the present, it has provided theoretical guidance for researchers investigating many types of behaviors that are not coordinated by a centralized authority. Park’s contention that social distance would be reflected in spatial distance provides a holistic framework for understanding the patterned nature of noninstitutionalized behavior from crime and vice to voting and religious affiliation. Ecology’s population perspective on social phenomena guides much research on stratification processes, including residential sorting and labor markets, family formation, and marriage. Despite the relative paucity of theoretical work, Quinn (1940) was able to find 347 pieces of ecological literature in 1940, and, a decade later, Quinn (1950) had found enough human ecology to fill 23 chapters in a 500-page review. The contributions in Micklin and Poston (1998) provide an extensive survey of human ecological theory and research. Berry and Kasarda’s (1977) book includes a review of the history of sociological human ecology in addition to an application of the theory to urban issues. Given the placement and purpose of this research paper, the discussion will focus on territorially delimited communities.

Understanding the Development of Human Ecology

Some of the early theoretical works in human ecology (Hollingshead 1940; Wirth 1945) and other disciplines were part of a boundary-making activity as factions of academics competed for recognized expertise over particular areas, but they were also symptomatic of a search for direction. Much of the early work in ecological theory must be seen as tentative first approximations of a theory—and more recent thought perhaps only a second approximation. In sociology’s exploratory context, a theory with predictive power may be beyond reach. A theory capable of generating post hoc interpretations is useful, and one that is incorrect in informative ways can lead to successively more powerful formulations.

Our coordinated knowledge which in the general sense of the term is Science, is formed by the meeting of two orders of experience. One order is constituted by the direct, immediate discriminations of particular observations. The other order is constituted by our general way of conceiving the Universe. They will be called, the Observational Order, and the Conceptual Order. The first point to remember is that the observational order is invariably interpreted in terms of the concepts supplied by the conceptual order. . . . We inherit an observational order, namely types of things which we do in fact discriminate; and we inherit a conceptual order, namely a rough system of ideas in terms of which we do in fact, interpret. (Whitehead 1933:198)

The quote, repeated by different authors who developed biological and sociological ecological theory, apparently captures the situation of sociologists in the first half of the twentieth century. They were searching for a methodology that would allow sociologists to adequately characterize the ways people lived together, and at the same time they were searching for ways of conceptualizing and understanding how that was accomplished. They were also searching for ways of bringing the observational and conceptual orders together, adjusting each in the process.

The following quote exemplifies the observational order that sociologists felt they needed to understand a century ago. Because Chicago played such a central role in the development of American sociology, an observation of that city is cited.

Chicago is one of the most incredible cities. By the lake there are a few comfortable residential districts, mostly with stone houses, and right behind them there are little old wooden houses. Then come the ‘‘ tenements ” of the workingmen and absurdly dirty streets which are unpaved, or there is miserable macadamization. In the “ city, ” among the “ skyscrapers, ’’ the condition of the streets is utterly hair-raising. And they burn soft coal. In broad daylight one can see only three blocks ahead—everything is haze and smoke, the whole lake is covered by a huge pall of smoke in which the sails of the ships putting to sea quickly disappear. It is an endless human desert. From the city one travels into the endless distance, past blocks with Greek inscriptions and then past others with Chinese taverns, Polish advertisements, German beer parlors, until one gets to the “ stockyards. ” For as far as one can see from the clock tower of the firm Armour & Co. there is nothing but herds of cattle, lowing, bleating, endless filth. But on the horizon all around—for the city continues for miles and miles, until it melts into the multitude of suburbs—there are churches and chapels, grain elevators, smoking chimneys, and houses of every size. All hell had broken loose in the “ stockyards ”: an unsuccessful strike, masses of strikebreakers; daily shootings with dozens of dead; a streetcar was overturned and a dozen women were squashed because a “ non-union man ” had sat in it; dynamite threats against the “ Elevated Railway, ” and one of its cars was actually derailed and plunged into the river. Right near our hotel a cigar dealer was murdered in broad daylight—all in all, a strange flowering of culture. There is a mad pell-mell of nationalities: Up and down the streets the Greeks shine the Yankees’ shoes for 5 cents. The Germans are their waiters, the Irish take care of their politics, and the Italians of their dirtiest ditch digging. With the exception of the better residential districts, the whole tremendous city is like a man whose skin has been peeled off and whose intestines are seen at work. One can see everything—in the evening the prostitutes are placed in a show window with electric light and the prices are displayed! A characteristic thing is the maintenance of a specific Jewish-German culture. Theaters present in Yiddish The Merchant of Venice (with Shylock prevailing, however) and their own Jewish plays. Everywhere one is struck by the tremendous intensity of work—most of all in the “ stockyards ” with their “ocean of blood.” From the moment when the unsuspecting bovine enters the slaughtering area, is hit by a hammer and collapses, it is in constant motion but [is] always (in the rhythm of work) tied to the machine that pulls the animal. One sees an absolutely incredible output in this atmosphere of steam, muck, blood, and hides. When they finish work, people often must travel for hours to get home. The streetcar company is bankrupt and hence does not purchase any new cars. The old ones break down every few moments. Around 400 people are killed or crippled in accidents every year. The company has calculated that those 400 indemnities cost it less than the required precautions, so it does not bother to introduce them. (Excerpted from Max Weber’s (1975:285–87) letters, written during his 1904 visit to the United States)

Weber is known for, among his other accomplishments, theoretical writings on cities. His main work on the topic (1958) was, appropriately enough, originally titled “Illegitimate Domination.” Yet his own work focused not on these observations but the power relationships and social contracts among the burghers of Medieval European cities in the absence of charisma or tradition as a legitimating force for social hierarchy.

With only minor revision, this was the situation a few years later, when McKenzie, Park, and later, Burgess arrived in Chicago. An American sociologist in Chicago observed what Weber saw, unfiltered by the interpretation of historians. If there was order here, it was not obviously one born out of social contract, much less of moral consensus. A social theory based on such agreement would be out of place. If the state had a role in creating order, its reach was seriously limited, and whatever the factors guiding behavior, they were not written into law. A theory stressing the importance of institutionalized authority would lack explanatory power. Individual and class interests probably motivated behavior but may not have determined the observed outcomes. An overly “organicist” conception of order would just as obviously have been out of place. In spite of the high level of apparent disorder, with individuals working at cross-purposes, somehow this social formation managed to survive. People continued arriving, and prosperity, at least most of the time, increased.

Searching for a way of conceptualizing what they were observing, Park (1936) and several others looked to the ecologist’s “web of life” and cited Darwin’s example of the cats and the clover to illustrate the binding factor in social arrangements.

[Darwin] found . . . that humblebees were almost indispensable to the fertilization of the heartsease, since other bees do not visit this flower. The same thing is true with some kinds of clover. Humblebees alone visit red clover, as other bees cannot reach the nectar. The inference is that if the humblebees became extinct or very rare in England, the heartsease and red clover would become very rare, or wholly disappear. However, the number of humblebees in any district depends in a great measure on the number of field mice, which destroy their combs and nests. It is estimated that more than twothirds of them are thus destroyed all over England. Near villages and small towns the nests of humblebees are more numerous than elsewhere and this is attributed to the number of cats that destroy the mice. Thus next year’s crop of purple clover in certain parts of England depends on the number of humblebees in the district; the number of humblebees depends upon the number of field mice, the number of field mice upon the number and the enterprise of the cats, and the number of cats as someone has added—depends on the number of old maids and others in neighboring villages who keep cats. (P. 2)

The image of cooperation without direct communication with its many indirect and surprising connections has captured the imagination of many sociologists. Competition was evident in cities a century ago but not a war of all against all. “Competitive cooperation,” a sociological idea that was pressed into biological service, seemed to capture the basis of human community. According to Park and Burgess ([1921] 1969:507), “competition invariably tends to create an impersonal social order in which each individual, being free to pursue his own profit, and, in a sense, is compelled to do so, makes every other individual a means to that end.” In other words, competition pushes people to cooperate. It has been difficult to keep both aspects of the concept simultaneously in focus, however, with some sociologists perhaps overemphasizing just one aspect of the concept. The ecological model allowed for a very populist image of human society wherein each person had an effect on others (even if unequal) and allowed for the possibility of a civil society and public sphere that did not exist in the shadow of the state—ideas that run throughout Park’s work (e.g., Park [1904] 1972).

The biology was, to some extent, used as a loose analogy or trope, and the early human ecologists made no attempt to press the point. Human ecologists’ efforts were frequently directed toward understanding exactly how the plant and animal analogy fell short, leading to much of the future development of the theory. Two points of divergence stand out. First, humans are better able to communicate and plan than are plants or animals. Relationships among them are thus more complex because coalitions can emerge from the competition, leading to conflict and possibly accommodation and assimilation in addition to specialization and segregation. Second, the different types of humans produced, to use Mills’s ([1959] 1967) phrase, are not species. They are capable of geographic and social movement, leading to a concern for patterns of social mobility. Despite the biological connotations of the term, human ecology is essentially interactionist sociology. Ecology, as Hawley (1944:399–400) pointed out, is a social science whether applied to plants, animals, or humans (with some then contemporary books carrying titles such as Plant Sociology ). Human ecology, with its observation that patterns of behavior are more variable than biological structure across species and its stress on adaptation to the social and physical environment, has often served as an antidote to explanations for social and cultural differences based on population genetics.

Park’s 1921 essay on sociology and social science suggested that following periods of sociology as a philosophy of history and as schools attempting to define a point of view, a period of investigation and research was beginning, and some of the subsequent debate was over the correct way of observing social phenomena. Much of the effort at the University of Chicago during the first third of the twentieth century went to community studies. McKenzie’s dissertation on Columbus, Ohio, neighborhoods was a multimethod attempt to create an adequate “observational order” characterizing social organization. Early human ecologists referred quite positively to the work of the “social survey movement,” including Booth (1920) in his landmark study of London. Several of the community studies performed during that era, such as Lynd and Lynd’s (1929) Middletown, had selected relatively small, stable communities for study. Larger cities, having differentiated into multiple social worlds, were obviously too extensive for any one participant or observer to adequately observe directly, leading to a crisis in methodology. Homans (1949) and Coleman (1990), among others, rued the disappearance of community studies with an encompassing view of social organization. The use of quantitative analysis and secondary sources emerged in the 1930s and 1940s as a method of performing a naturalistic study while maintaining a holistic view. While some commentators have suggested that quantitative analysis was a strategy for gaining scientific legitimacy, by 1950 the claim to that legitimacy was more strongly made by sample surveys and small-group laboratory research, both of which sacrificed holism.

Human Ecology Distilled

The fundamental contention of human ecology is that community (social organization) arises through the interaction of a population and its physical and social environment. Ecology assumes a collective tendency to maximize life that is, however, limited by the repertoire of tools, techniques, and information (technology) available. That simple statement carries many implications. Because every living organism requires access to the biophysical environment for food and water, environmental dependence, however indirect, is an ever-present fact of life. The ultimate dependence of all humans on the biophysical environment implies that social organization is not purely a social construction emerging out of interaction. The social construction is thereby anchored. Because individuals are time bound, economizing on time is a significant factor in the maximization of life. Because interdependence is a means of maximizing life by allowing environmental access while economizing on time, it forms the basis for social relations. Because the intrinsic limitation on the behavioral variability of humans is indeterminate, interdependence can take many—sometimes simple, sometimes complex—forms.

Competition for resources among individuals plays a critical role in social organization. Competition can have at least three general types of effects on community (Schnore 1958), however, with the path of least resistance possibly prevailing. Demographic responses, a decline in births or increase in mortality, are one possible response to competition for resources (Hannan and Freeman 1977). Technological responses, the more efficient use of resources through conservation or the enlargement of the resources base, are a second collective response to competition (Boserup 1966; Hawley 1950:203; Schumpeter 1950). Organizational responses are a third type of response to competition. As Cooley (1930a:164) has suggested, “If ‘all the world’s a stage’ [competition] is a process that distributes the parts among the players” (quoted in Hawley [1950:201]). It even helps create those roles. Competition leads to specialization and, therefore, differentiation. Differentiation results in a hierarchy of activities when the most productive activity is in most direct contact with the environment, resulting in its becoming a dominant or key function defining the conditions under which others will operate, creating sometimes intricate patterns of cooperation in which the participants might not even experience competition per se.

Activities are performed by people, leading to a process of matching individuals and activities and creating social stratification. The competition for resources that creates differentiation encourages the formation of coalitions to gain and protect such resources. The possible bases for coalitions are nearly endless, including even eye color. The salience of particular characteristics for such purposes depends on the ease and effectiveness with which they can be mobilized as a basis for action (Nielson 1985; Olzak 1986). Competition creates the impetus to form coalitions, and the success of the various efforts creates social categories that become the basis for commensal relations. While in some cases the categoric basis for recruitment is easily measured, in other cases difficult-to-measure criteria apply, such as the nature of a personnel director’s university education (Useem 1989) or the neighborhood in which a person resides. The sometimes excessive credentialing requirements for particular occupations, selecting on parental income, could also be included among the commensal mechanisms of resource allocation. None of the bases are primordial.

The resulting social organization is composed of two interweaving, but distinct, types of interdependence relations: symbiotic and commensal. Symbiotic relations imply a degree of mutual interdependence among complementary and, therefore, functionally differentiated and specialized units, such as the different species in the “web of life” example above. Commensal relations, arising among those who engage in similar activities in a symbiotic web, imply common, parallel actions. Being built on the basis of similarity, they imply both a degree of competition for resources and cooperation to protect access to them. This mix of interests leads to restrictions on individual actions, providing a basis for understanding the structural bases of action. Despite focusing on the morphology of social organization, human agency in all its varied manifestations is essential to the ecological theoretical framework. The human “balance of nature” can be attained only when life expands to the maximum attainable under prevailing conditions. The disparate aims, attempts, and projects of many individuals imply that the collective response to an environment is not simply a result of an individual’s action writ large nor merely a cultural schema made real.

The “balance of nature” may be elusive, existing mainly in the minds of researchers, however (Park 1936:5, quoting Charles Enid). That equilibrium may be affected by any event that upsets the relationship between population (a set of individuals), organization (a set of direct and indirect social relations created by interrelated efforts at sustaining life), environment (a set of direct and indirect resources), and technology (a set of tools, techniques, and information), resulting in a population problem that becomes an impetus for social change that can be resolved in the three general ways outlined.

Changes in Social Organization

If the human “balance of nature” is essentially a matter of matching persons with activities, anything that disturbs that balance leads to social change. Changes in the productivity of sustenance technology have had a major impact on social organization, increasing the aggregate population that could be supported and the degree of hierarchical placement. Over the past century or so, communication and transportation technologies have clearly had substantial implications, increasing the scope of social integration. Each of these has directly or indirectly affected the size of the population compared with the set of activities. Technology, in all its various forms, is central to theories of large-scale social organization because it is a mechanism that stabilizes social relations despite changing actors, beliefs, and interests (Latour 1991). Human ecologists have explored some implications of the basic model more extensively compared with others, however.

Technological Change

Advances in subsistence technology are a necessary precondition for any significant increase in the size or complexity of any social formation (Nolan and Lenski 1999:65). Those technologies have evolved from hunting and gathering to horticultural to agricultural to industrial. Over the last century or so, that has resulted in a shift from farm to factory to office in the United States and other countries. The Industrial Revolution was due, in part, to the increasing efficiency of agricultural production, and the growth of service employment rests on increasing manufacturing efficiency (Baumol, Blackman, and Wolff 1989). The spatial distribution of population has changed as particular industries have expanded or contracted as new cities grew on the basis of specific products (Duncan and Lieberson 1970).

Subsistence technology influences social stratification by affecting the volume of surplus wealth and by influencing the bargaining power of all involved. Rewards (the product of labor) will be shared to the extent required to insure the survival and continued productivity of those others whose actions are necessary or beneficial to themselves (Lenski 1966). Recall that competition tends to create a social order wherein individuals are compelled to use others as means to their own ends (Park and Burgess 1921:507). But since each technology requires a different set of human resources for maximum output, the outcomes of the process of bargaining vary with the methods of production available. In the simplest societies, which generate small surpluses, such as hunter-gatherer bands, goods will be distributed largely on the basis of need. Across types of society, from those based on hunting and gathering to those based on agriculture, inequality grew as power differentials based on the degree of centrality in a network of interdependence increased. That historical trend reversed with the appearance of mature industrial societies because the greater complexity of functional interdependences gave those with specializations that supported the key function increased bargaining power. In agricultural societies, for example, 90 to 95 percent of the population was involved in the same basic function, leaving many individuals with little bargaining power and a few with central coordinating roles. The division of labor is substantially more complex in industrial societies, and individual functions are significantly more tightly integrated—with the automobile assembly line being the ideal typical example of each participant having veto power over production at any one moment. An increased ability to communicate, sometimes aided by proximity, gave those performing similar functions greater ability to solidify commensal relations, with those being often institutionalized in the form of labor unions, professional organizations, and industry groups.

Communication Technology and Ecological Expansion

A concern for communication and transportation technologies stems from human ecology’s roots in interactionist sociology (Park 1921:13, 17). The unity of social formations is built on interaction. Thus, any development that extends the reach of communication, whether writing and newspapers (Park 1923) or railroads and automobiles (Ogburn 1946; Ullman 1980), theoretically affects the size and structure of human communities. Over the past century or so, improvements in transportation and communication technologies have changed unusually rapidly, and their impact on social organization has been pervasive. Both material and nonmaterial aspects of these technologies have been important, and their development is closely intertwined. Large firms, for example, are communication technologies facilitating and channeling interaction. These technologies have allowed the size and geographic scope of organizational units to expand, giving rise to, at the largest scale, globalization.

Face-to-face exchange is the basic form of social interdependence. Urban planners sometimes use a radius of a fiveminute walk in defining and planning neighborhoods (Leccese and McCormick 1999). The maximum distance people are willing to commute daily over long periods is approximately one hour each way, and the reach of daily delivery trips appears to delimit regions (Hawley 1950). Temporal rhythms combined with existing transportation technology, by affecting the frequency of interaction, helps form the texture of social organization. The extent of interaction is limited by the costs of carrying out interactions and the anticipated benefits to completing the interaction. Therefore, social innovations enhancing the ability to interact expanded the maximum possible size, differentiation, and hierarchy of populations. Improvements in the technology of transportation have expanded the range of interaction.

Each mode of interaction has its own characteristics, having to do with the relative costs of movement and of loading and unloading and with the democracy of destination. Navigation is limited to water, and river transportation is constrained by the course of water flow, leading to a long, thin settlement pattern, such as that along the ancient Nile. Sea navigation is less constrained but is focused by the location of natural harbors. Singapore, for example, owes its status as a busy port (and its prosperity) to its favorable location, Chicago to a river link between the Great Lakes and the Mississippi Valley. The development of railroads allowed the growth of large inland cities, with the railroad yards often within sight of the central business district. Atlanta and Columbus rose at railroad intersections. Both ships and railroads have relatively high terminal costs but low transportation costs, privileging distant relative to moderately near locations for interaction while concentrating activities at particular points of transfer. These seemingly trivial factors have had large effects on large-scale social organization.

Three inventions, all emerging within a short period of history, the wireless radio (1895), the automobile (1986), and the airplane (1903), revolutionized communication and travel because of the greater distances that could be covered and because of the greater democracy in their movement. The telegraph had already begun divorcing communication from transportation by 1840. Before that time, all communication required travel by a person (Pred 1966). The radio freed the transmission of information from the constraints of wires. More recently, airports have developed as centers for interaction (Irwin and Kasarda 1991). These centers may have been based on earlier settlements, but air transport further opened internal sites for further population growth. In the United States, air transport helped Denver grow, and the emerging freight transfer belt reaching from Louisville through Memphis and beyond may follow.

There is a tendency for population and wealth to collect at breaks or interruptions in routes of transportation (Cooley [1894] 1930b). Mechanical breaks in movement resulting from a change in mode of transport have led to commercial breaks resulting from a change of ownership. Freight must be unloaded from one carrier and placed aboard another, and often it must be stored until a second carrier is ready or until an exchange is completed. Terminal facilities must, therefore, include warehousing and space for support functions such as repair and the administration of trade. That implies workers to handle the cargo and its administration, which, in turn, calls for their own support in the form of housing, food supply, and even entertainment. The accretion of personnel and physical structure at the point of interruption is cumulative; each new addition or elaboration entails others, and ultimately a large and complexly organized settlement unit takes form (Hawley 1950:243).

Some of the costs of interaction are purely social in nature, and as the frequency of various types of exchange increased, the attendant negotiations were regularized and the extent of interaction grew. The development of writing and, in stages, printing were milestones in the development of the technology of communication, each having a major social impact through preserving information over time and diffusing it through space. The gradual institutionalization of interaction in associational units, such as business firms and other bureaucracies, in the form of behavioral templates for negotiation also allowed the extent of interaction to increase. Large business firms are, in part, communication technologies (Beniger 1986) and, in part, templates or algorithms for social activity (Berlinski 2000). Internal economies of scale not being a sufficient condition, the growth of large firms was closely related to the need to coordinate activities over long distances, with the railroad companies themselves being among the earliest large firms and those that coordinated buying, production, and sales over long distances rapidly following (Chandler 1977). Appearing only in the second half of the nineteenth century in industry, by the 1920s, they were the most influential nongovernmental organizations. Such formal organizations emerged when the volume of exchange was sufficiently large and steady to routinize the transactions, lowering their costs, allowing them to schedule the flow of goods more closely than markets could.

Mismatches and Surplus Populations

The railroad, the automobile, the airplane, and the large organization have continued to radically, but not completely, shrink space. Near the end of the nineteenth century, streetcars and electric trains extended the reach of daily movement, particularly for those who could afford the transportation, allowing homes to be further separated from workplaces. Such separation allowed some workplaces to expand, facilitating the functional integration of a larger population. The combination of physically and socially expanding social systems created a physical and social path for new in-migrants: As the established population gained a more elevated position in the functional hierarchy and moved to more desirable residences more distant from the city center, a physical and social place was created for the new arrivals. Ecological expansion set in motion a process of physical and social succession that allowed for upward mobility. Put prosaically, as some people got better jobs and moved to the suburbs, occupational and residential space was created for others. Ecological expansion facilitated residential succession. Maintaining a balance between the number and type of people and the number and type of activities is an important ongoing social process, and the mechanisms of allocation are a central concern for human ecologists (Duncan 1965; Logan 1996; Stewman and Konda 1983). The processes of adjustment between population and activities is rarely smooth or efficient, leading to “surplus populations” that are themselves impetuses for social change.

The surprises induced by the many mismatches in social and geographic location, such as the slum just outside the “Gold Coast” (Zorbaugh 1929), the hobo’s “main stem” near the business district (Anderson 1923), or the artist colony in a city of commerce, add a sense of dynamism to community life. Artists and literary writers do not necessarily require an urban location, however. They have little need to interact directly with the city around them. Indeed, many artistic communities have emerged in out-of-the-way locations such as Black Mountain, North Carolina. Nevertheless, the process of ecological expansion, outlined above, has often created underutilized space in the older centers of many cities, and artists have sometimes gathered in such urban zonesin-transition. The abandonment of Greenwich Village by the well-to-do at the beginning of the twentieth century for more efficient, uptown residences created a convenient, low-cost location for literary people, who were not closely integrated with the financial, commercial, and manufacturing activities surrounding them (Coser 1965). Deviance has been interesting to sociologists both as a social problem and, because of its theoretical relationship to social change, as a sociological issue. Certain types of crime and “vice” once collected in urban zones-in-transition, not because of any direct relationship between population density and crime but because such locations were the sites of least resistance.

The ecological expansion that allowed many people to move to higher-quality suburban housing and artists to congregate in central cities also created spaces for those whose proximity was also not required by nearby productive activities but who did not have the social power to be elsewhere. Ecological expansion, combined perhaps with the operation of commensal relations in the allocation of resources, has led to the segregation of population by firm, occupation, and residential area. As an extreme case, the diffusion of automobile transportation plus a “declining significance of race” in determining residence (Wilson 1978) has ironically led to residential hypersegregation along racial, educational, and income lines in the United States (Massey and Denton 1993). Employment in occupations and firms remains racially and gender segregated to a degree that cannot be explained by the qualifications of the respective groups (Jacobsen 1997; Petersen and Morgan 1995; Reskin and Cassirer 1996). Such hypersegregation may have led to the creation of an underclass—a set of people who are not just poor but effectively cut off from economic opportunities for themselves and their children. This situation can be seen in the spatial employment mismatches that still plague many U.S. cities. These mismatches are not merely spatial phenomena. They are intensified by commensal relations in the form of local governments that, under some conditions, may effectively undermine the educational opportunities for central-city residents (Kasarda 1989).

Sometimes, whole cities and regions have been made redundant by the industrial restructuring brought about by ecological expansion. Even so, the expansion process was not purely a matter of technological advance. Surplus population in rural areas and the U.S. South helped attract employment as transportation improvements allowed dispersion. Similarly, the declining mortality rates in what used to be called the Third World created a shift in the global locus of population. In 1950, two-thirds of the nonagricultural labor force in the world was found in the then industrialized countries. By 2000, fully two-thirds of a much larger nonagricultural labor force was to be found in the previously less-industrialized countries. (It is difficult to remember that a few short decades ago, labor shortages were so acute in Europe and North America that migrants were actively recruited and full employment was seen as a realistic policy goal.) As the technical and organizational aspects of transportation and communication technologies developed to the extent that close integration was possible, a surplus population was waiting for opportunities (Frobel, Heinrichs, and Kreye 1980).

Human Ecology in Perspective

Human ecology could be classified along with a larger family of theories that seek the explanation for patterns of social organization in the attempts of people to solve everyday problems. Several of the ecological postulates are, therefore, shared by other theoretical approaches in sociology. The assumption that social relations are rooted in biological needs is fundamental to the theories developed by Mead (1934), for example. The view of social organization as a set of exchanges of resources can be found in Blau (1964), among many others. The idea that social organization can be characterized as a complex pattern of relationships is developed by Nadel (1957) and others. Similarly, a division of social relationships into those that are directly concerned with productive work and those that are in reaction to the conditions of productive work is maintained by Homans (1950) and others. Human ecology differs from other approaches to social organization perhaps in its focus on the holistic implications of these postulates, examining social organization as a population characteristic, rather than individual behavior or cultural meaning.

Human ecology’s holistic stance is justified on the basis of the equifinality of micromotives. Becker (1976), for example, provided a dramatic demonstration to show that three very different decision rules—utility maximization, tradition-bound behavior, and random decisions—all produce the same aggregate patterns of behavior. Accordingly, Coleman (1990), White (1992), and Collins (2004) have each recently published comprehensive theoretical works examining how individual motivations and behavior aggregate into large-scale social organization. Each of those authors relied on very different assumptions and mechanisms to explain similar outcomes. By providing a broad overview of social process, macrosociology has much to contribute to the understanding of contemporary social organization.

One important aspect of that overview would be human ecology’s perspective on power, disconnecting outcomes from actions and intentions. The ecological conception of dominance is somewhat broader than the concept of interpersonal power. Interpersonal power is usually defined in terms of person A compelling person B to act in accord with person A’s will against B’s own will for A’s benefit. An ecological sense of power includes all that affects daily rhythms, the course of lives, occupational and residential distribution, and so on—that is, dominance is defined in terms of effect—the influencing of behavior—rather than in terms of intention or benefit. Often political institutions lack the means to overcome diffuse forces (Mann 1986). Ecology does not make assumptions about the stability of a set of actors either. In any form of large-scale social organization, there is likely to be a changing set of people and a changing set of role relations, and the connections between actions and outcomes is tenuous (March and Olsen 1976; Suttles 1990). By sampling on cities that were in some way successful, some of the literature on growth coalitions misinterprets the presence of particular types of actors for the causes of outcomes, overestimating the power of a small elite to determine events (Logan and Crowder 2002).

Becausecommunityisdefined“not[by]like-mindedness, but [by] corporate action” (Park and Burgess 1921:42), the ecological approach to social organization informs the psychological, cultural, and moral aspects of human experience. The sociology of knowledge, for example, attempts to relate beliefs, attitudes, and values to social position (Mannheim 1936). Consequently, David Riesman (1950) based his division of tradition-directed, innerdirected, other-directed character types on the social positions and demographic rates linked to ecological organization. Important aspects of contemporary culture can be understood in the light of ecological expansion and the attendant population mismatches (Harvey 1989). More important, examining social relations from the perspective of functional integration allows for an understanding of moral ambiguity and cultural conflict in ways that theories based on a consensus normative order cannot (Alexander 1982:108). Such theories do not account for how values and norms are formed or for the reasons for conflict. A simple model positing that (1) in equilibrium “rule and action are one and the same” (Hawley 1986), (2) an occupational and demographic shift was induced by ecological expansion, and (3) a cultural lag exists (Ogburn 1964) appears to be quite useful in understanding many contemporary moral dilemmas (Hochschild 1989).

Today, we are increasingly tightly linked in a web of interdependences, yet many feel a loss of community. Browsing through the architecture and planning sections of a library or bookstore turns up many books suggesting that making our streets a little narrower, our settlements a bit more dense, our land uses somewhat more integrated, and our parking lots smaller would somehow rebuild community. Others suggest that living in central cities, perhaps in apartments instead of split-levels, traveling by foot or bus instead of automobile, and spending more time in coffee shops or bars instead of in living rooms would somehow heal our psyches. These books are full of drawings and photographs of buildings and streetscapes. They are largely empty of information on how the residents and users of the depicted spaces use their time and interact with others. Ecological theory suggests that such physical arrangements would produce, at best, simulacra of community. Human ecology shifts attention from the layout of buildings to the symbiotic and commensal relations that constitute community.

The Future of Human Ecology

Human ecology posits that individuals adapt to their environments by forming symbiotic and commensal relationships of varying degrees of strength and pattern. It further postulates a collective tendency for the emergent pattern to maximize life to environmental limits, given the available technologies for subsistence and interaction. The heart of human ecology lies in understanding the web of life created by those relationships and how it is affected by particular changes. The process of ecological expansion has intertwined with population-activity mismatches to produce social change. Drawn by public interest and driven by intellectual comparative advantage, sociologists have explored human ecology in some directions and ignored others. Today, sociological human ecology could be characterized as a set of loose congeries of researchers, some of whom have independently rediscovered the basic ecological insights with no common research agenda (Freese 2001). It may be time for a new systematic synthesis. This research paper will close with just three questions for urban research—each, so far, only partially explored—that ecological theory may help address.

What effect do multilocational firms have on urban development? The present level of spatial integration probably owes as much to the consequent efforts to standardize products and procedures (as exemplified by the diverse activities of the International Standards Organization, which now goes so far as to specify standards for management methods) as to the jet plane and Internet themselves. Such possibilities suggest that cross-cutting forms of social organization—functionally integrated formal organizations and spatially delimited regions—have substantially changed the nature of spatial development. Physical break-in-bulk points were once also change-in-ownership points, requiring inspection, administration, and storage. To the extent that the former no longer implies the latter, the number of personnel required at break-in-bulk points diminishes (relative to what it otherwise would have been). Administration and goods movement may become progressively further decoupled from each other. Moreover, to the extent that the movement of goods is closely coordinated, the need for warehousing and other support facilities may be diminished. It is, for example, possible for a truck to arrive at a container port or airport just in time to whisk away or deliver a cargo, diminishing the functional importance of agglomeration. Cities are a collective method of economizing on time—but only one of several.

What will be the role of the so-called world cities as the global economy continues to expand? A handful of cities appear to be powerful command and control centers, based on finance and innovation in an expanding global economy. Power in the global system has proved to be ambiguous, however. Structural equivalence in trade and financial networks reduces the benefits of centrality. Trade surpluses in manufacturing goods have given “peripheral” countries considerable financial power. New York City has long ceased to be the primary center of scientific and technical innovation envisioned by Vernon (1960, 1966). Technological innovation occurs within a broad and expanding zone of indifference within wealthy societies. Research and development activities and other higherorder services appear to be relatively free of locational constraints. Attracting such activities has become an area of much tension for localities, indicating that research and development are not necessarily bound to particular places and that the locational factors are not well understood. Major multinational corporations are now headquartered in relatively small, seemingly remote towns with no apparent attenuation in accessibility. Much as some U.S. cities have become empty cores surrounded by prospering suburbs, the world system may develop in much the same way, with no clear role for the “world cities.”

What is driving gentrification? Over the past several decades, there has been a small but perceptible flow of highly educated, mostly childless adults to a few select cities. Some have suggested that this is an expression of a “new class” or evidence of an exogenous culture change. Another possible explanation for this small migratory counterflow is based on the effects of the difficulties of matching people to jobs, which has led to progressively longer periods of education. In many countries, the surplus of educated labor has resulted in the externalization of employment (“boundaryless careers”), weak wages, and flat career trajectories, much as an oversupply of manufacturing capability in some sectors has led to the externalization of many activities (Piore and Sabel 1984). In some countries, a surplus of population has led to a pattern of inequality where the income variation within education, experience, and gender categories accounts for an increasing degree of total income equality. Some of those relatively underpaid service workers, unable to afford increasingly expensive suburban housing or to start families, may be congregating in a subset of American cities. From the point of view of the cities, this appears as gentrification, but from a larger point of view, it may be an expression of a “fear of falling” (Ehrenreich 1989).

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