The concrete tangle: Tolerance of anthropogenically disturbed habitats influences cophylogenetic congruence between migratory birds and their haemosporidian parasites

Urbanization usually decreases biodiversity, but select species can tolerate or thrive in human-modified environments. Broad environmental tolerance typically characterizes species that persist in urban habitats. Heterogeneous environmental conditions used by urban-adapted species could influence the specialization that is adaptive for their parasites. Parasites subjected to diverse environments may experience selective pressures favoring host generalism, which could promote host-switching in parasites of urban-tolerant hosts. We test the hypothesis that a host species’ tolerance of urban habitats corresponds with decreased cophylogenetic congruence with their parasites using avian haemosporidians (genera Plasmodium and Haemoproteus). Bird species vary in their response to urbanization, ranging from exclusion to achieving high densities. Avian haemosporidia are diverse intra-erythrocytic parasites with specificity ranging from specialists with strong co-evolutionary relationships with their hosts to generalists more prone to host-switching. Employing a global database of associations between haemosporidian lineages and avian host species, we use the Procrustean Approach to Cophylogeny and phylogenetic generalized linear mixed models to test if host–parasite links associated with urban-adapted bird species have lower contributions to overall cophylogenetic congruence. Our results show that the integration between urban tolerance and migratory strategy affect co-evolutionary relationships between hosts and parasites. Specifically, fully migratory bird species known to occur in disturbed habitats on average have co-evolutionary histories with their haemosporidian parasites that are more strongly characterized by host-switching. Understanding the ecological drivers of host-switching is key to predicting and managing parasite spread mediated by cross-species transmission, especially as an urbanizing world promotes higher abundances of urban-adapted species.

Urbanization is among the most permanent and transformative types of anthropogenic land use (McKinney, 2009). The loss, degradation, and conversion of natural habitats with urbanization poses a mounting threat to the planet’s biodiversity, especially as the pace of urban expansion continues to accelerate to accommodate human population growth (Angel et al., 2011). However, urban expansion does not affect all species equally, nor does it affect all species negatively. Wildlife responses to urbanization rely on a suite of species-specific ecological, behavioral, physiological, and life history characteristics, which can sometimes be linked to shared evolutionary history (Betke et al., 2023; Iglesias-Carrasco et al., 2022; Rodewald & Gehrt, 2014). In other words, although the net effect of urbanization on biodiversity is negative, any given species occupies a continuum of urban tolerance based on how well its functional traits suit it to urban life (the urban tolerance hypothesis; (Sol et al., 2014). At one end of this continuum, “urban avoiders” are sensitive to anthropogenic disturbance and tend to be excluded from urban environments (Blair, 1996). At the other extreme, species termed “urban exploiters”, “urban adapters”, or “synanthropes” are highly tolerant of modified habitats and can effectively use, occupy, and achieve high population densities in urban environments (Blair, 1996; Francis & Chadwick, 2012; Santini et al., 2019). The ecological traits typifying synanthropes are relatively well-characterized in wild birds. Globally, urban-tolerant bird species tend to be smaller-bodied, gregarious, more dispersal-prone, and have broader niche breadths in terms of both dietary and habitat generalism (Bonier et al., 2007; Møller, 2009; Neate-Clegg et al., 2023). The contribution of these functional traits to urban tolerance can vary with geography (Neate-Clegg et al., 2023) but is relatively consistent. While urban-tolerant mammals have adopted diverse and taxonomically stratified strategies for urban life (Betke et al., 2023; Santini et al., 2019), urban bird communities are generally defined by this set of “winning” traits irrespective of phylogeny (Evans et al., 2011), (Santini et al., 2019). As the species-specific traits associated with urban tolerance filter the rapidly homogenizing bird diversity present in anthropogenic habitats, these traits should become increasingly represented in urban bird communities (Luck & Smallbone, 2011). Depauperate diversity in urban communities and increased densities of urban-adapted species can have consequences for infectious diseases. Urban-adapted species often harbor greater parasite richness than species unassociated with urban landscapes and can serve as reservoir hosts, increasing infection prevalence in their communities (Albery et al., 2022; Ecke et al., 2022; Marm Kilpatrick et al., 2006). While most comparative work to date has focused on how urban tolerance shapes variation in parasite diversity and abundance, this characteristic could also favor parasites with certain traits. For example, urbanization may favor transmission of pathogens spread through direct contact owing to density-dependent transmission and close contact around clumped anthropogenic resources (Murray et al., 2019). Beyond transmission mode, an equally fundamental feature of parasites is their host specificity, a measure of ecological specialization that can reflect both the number and phylogenetic breadth of hosts a parasite can exploit (Poulin, 2007). Parasites with high host specificity are associated with a single or few closely related hosts (i.e., “host specialists”), whereas parasites with low specificity may use numerous, phylogenetically distant hosts (i.e., “host generalists”). Host specialists are often assumed to have diversified via cospeciation, resulting in evolutionary histories that mirror that of their hosts’. In contrast, host generalists are expected to have their co-evolutionary congruence disrupted by relatively frequent host-switching events. Host specificity and emergent macro-evolutionary patterns (e.g., evolutionary histories shaped predominantly by host-switching or by cospeciation) are influenced by both host and vector ecology, particularly in parasites that are not free-living (Huyse et al. 2005). For example, solitary social systems in hosts can reinforce high specificity and cospeciation with their parasites (Hafner et al., 1994), while host migratory propensity or dispersal ability can instead lead to host-switching opportunities and lower co-evolutionary congruence (Jenkins et al., 2012; Boyd et al., 2022); but see (de Angeli Dutra et al., 2022). However, whether co-phylogenetic congruence and host switching of parasites is modified by the unique ecological traits associated with urban tolerance remains poorly understood. Here, we test the contribution of urban tolerance in avian hosts to cophylogenetic relationships with their haemosporidian parasites. Haemosporidian parasites of birds (in the genera Plasmodium, Haemoproteus, and Leucocytozoon ) are cosmopolitan, intra-erythrocytic protozoan parasites vectored by mosquitoes, biting midges, hippoboscid flies, and black flies. Haemosporidian infections are highly prevalent in wild birds and are often associated with sparse and sub-lethal effects on their hosts, such as decreases in body condition and reproductive success (Valkiūnas 2005); however, their pathogenicity is both lineage- and context-dependent, and haemosporidian infections have been implicated in the decline and extirpation of multiple bird populations (Dadam et al., 2019; Woodworth et al., 2005). Avian haemosporidia demonstrate variable degrees of host specificity, ranging from host specialists recovered consistently from a single species to generalist lineages associated with a global range of hosts (Ricklefs et al., 2005). The prevalence, diversity, and varying host specialization strategies used by haemosporidians underpin their status as a model system for studying host–parasite interactions. For example, bird–haemosporidian associations are a valuable empirical system for testing hypotheses at the interface of host migration and parasitism (de Angeli Dutra et al., 2021; Emmenegger et al., 2018). Avian haemosporidia are also especially useful for testing hypotheses related to the contributions of host–parasite links to overall cophylogenetic congruence, because they have patterns of diversification driven both by host-switching and cospeciation (Ricklefs et al., 2004; Santiago-Alarcon et al., 2014). We predict the broader niche breadth and higher dispersal ability of urban-tolerant bird species could expose haemosporidian parasites of these hosts to greater environmental heterogeneity, favoring selection for host generalism and diversification via host-switching events (Kassen 2002). Alternatively, bird species more prone to using disturbed habitats may expose their parasites to less avian host diversity, which could curb selection for host generalism (Moens & Pérez-Tris, 2016). Migration strategy may also influence cophylogenetic congruence either independently or in tandem with urban tolerance, given the widespread taxonomic and geographic distribution of migratory behavior among birds (Dufour et al., 2020; Tobias et al., 2022). A migratory host could promote host-switching events by increasing encounters with parasites and periodically suppressing host immunity (Arriero & Møller, 2008; Eikenaar & Hegemann, 2016); alternatively, migration could instead favor co-evolution of parasites to better exploit a host with seasonally variable internal and external environments (Clayton et al., 2003). An urban-tolerant, generalist host species may subject its parasites to similar pressures at a smaller ecological scale, which could result in convergent cophylogenetic patterns between parasites and birds that are resident but urban-tolerant and birds that are migratory but urban-intolerant. To test these predictions, we employed a global database of associations between haemosporidian lineages and avian host species and then used the Procrustean Approach to Cophylogeny (PACo) and phylogenetic generalized linear mixed models (PGLMMs) to test if host–parasite links associated with urban-adapted and migratory bird species have lower contributions to overall cophylogenetic congruence. Host–parasite data We used the malaviR package in R to programmatically download all available haemosporidian lineages from the MalAvi database (accessed 4/4/2023), a comprehensive, standardized, and dynamically updated record of haemosporidian cytochrome b sequences from avian hosts (Bensch et al., 2009). We conservatively pruned the dataset to include only hosts identified to species level and limited our data to those host–parasite links recorded at least twice to reduce including erroneous associations while attempting to preserve signal from rare and/or specialist haemosporidian lineages. The trimmed MalAvi alignment included a total of 739 haemosporidian lineages (288 in the genus Plasmodium and 451 in the genus Haemoproteus ). Lineages in the genus Leucocytozoon were used to construct the haemosporidian phylogeny but excluded from downstream analyses of host–parasite associations for computational feasibility. The resulting dataset comprised 1,655 associations between birds and haemosporidian parasites. Host and parasite phylogeny We standardized host species names in MalAvi against the BirdTree taxonomy and then used BirdTree to produce a consensus phylogeny for passerine hosts using 2,000 randomly sampled trees generated from the Hackett backbone (Jetz et al., 2012). We removed bird species not present in the MalAvi database, resulting in a total of 640 host species spanning 89 avian families. We constructed a maximum likelihood phylogeny of haemosporidian parasite lineages in IQ-TREE 2.2.2.6 (Minh et al., 2020) using 1,000 bootstrap replicates and a GTR+I+G substitution model (Abadi et al., 2019). Host species traits We next assigned urban occurrence, disturbed habitat occurrence, an urban tolerance index (UTI), and an avoider/exploiter metric to each host species present in the MalAvi database using the dataset curated by González-Lagos et al., 2021. Briefly, this dataset defines species-level urban and human-disturbed occurrence using the International Union for Conservation of Nature (IUCN) habitat classification data to determine if a species’ native range overlaps with modified environments. The urban tolerance index (UTI) is a continuous value using the ratio between individuals of a species occupying an urban area and those occupying surrounding natural environments to account for the dependence of urban occurrence on natural populations bordering urban sites (Evans et al., 2011; Sol et al., 2017). Large positive UTI values are thus indicative of species very tolerant of urban habitat, whereas large negative UTI values suggest low urban tolerance (Evans et al., 2011; González-Lagos et al., 2021). Lastly, the exploiter/avoider metric uses community simulations to designate species as urban exploiters or avoiders based on whether they are significantly more or less likely to be abundant in urban environments than expected by chance dispersal from surrounding habitats (Sol et al., 2014). Because the UTI and exploiter/avoider metrics rely on population surveys of bird assemblages across urban gradients, these metrics characterize fewer species ( n =639 for urban or disturbed habitat occurrence, n =235 for urban tolerance, and n =171 for exploiter/avoider status) but represent their tolerance of urban habitats more effectively than occurrence data alone (Sol et al., 2014). We also assigned migratory status to each host species per the AVONET database (Tobias et al., 2022). We collapsed migratory status into two categories: fully migratory species, wherein most of the population are long-distance migrants, and non-fully migratory species, which comprise both partially migratory and sedentary species. Statistical analysis To first test for overall congruence between the host and parasite phylogenies and the relative contribution of host–parasite links associated with urban-tolerant hosts to this global fit, we used PACo as implemented with the paco package in R (Hutchinson et al., 2017). We constructed a host–parasite association matrix and separate phylogenetic distance matrices for hosts and parasites using the ape package (Paradis et al., 2004). Phylogenetic distance matrices were then converted to principal coordinates prior to Procrustes imposition. The sum of squares residuals yielded by the superimposition between the host and parasite principal coordinates indicates their overall congruence, and we used a goodness-of-fit test against 1,000 random permutations of the association matrices to determine if this level of congruence was greater than expected by chance. To assess the contribution of individual interactions to overall congruence, we used a jackknife procedure to calculate the corrected Procrustes residuals for each host–parasite link. A lower residual value indicates a larger contribution of a given interaction to global cophylogenetic congruence. We then analyzed these PACo residuals in a series of PGLMMs using the brms package (Bürkner, 2017). We averaged residuals per host and used their log-transformed values as a Gaussian response in eight models. These models differed in the predictor variable used to describe urban tolerance (i.e., urban occurrence, disturbed habitat occurrence, UTI, or exploiter/avoider metric) and additive or interaction effects between migration strategy and urban tolerance metric (Table 1). All models included a host phylogenetic random effect via covariance matrix from the host phylogeny as well as a fixed effect of number of citations per host species as a proxy for research effort, programmatically collected with the easyPubMed package. We ran all PGLMMs using four chains for 20,000 iterations, a burn-in of 50%, and default priors. We verified model convergence by inspecting trace plots and R-hat values. We extracted posterior means and 95% credible intervals for each model coefficient in brms . We compared models representing the same number of host species (Table 1) using the leave-one-out cross-validation information criterion (LOOIC) via the brms package, wherein the model with the lowest LOOIC score has the highest predictive performance.

The global test of cophylogenetic signal in PACo rejected the null hypothesis of random associations between bird species and their haemosporidian lineages (residual sum of squares = 1049, p < 0.001), indicating that haemosporidian evolution overall tracks that of their avian hosts (Figure 1). However, we also observed substantial heterogeneity in PACo residuals, suggesting cospeciation is not the sole driver of recorded host-parasite associations. Using our host species-level mean PACo residuals as the response variable in PGLMMs, we found that models with the interaction between migratory strategy and urban occurrence were more competitive than models with only additive effects (Table 1). The best-performing occurrence model included a species’ ability to use habitat that is anthropogenically disturbed but not urbanized (Table 1). By contrast, for models considering UTI or exploiter/avoider status, both interactive and additive PGLMMs were competitive (ΔLOOIC < 2), indicating that the interaction between urban tolerance and migration was uninformative (Arnold, 2010). Across the most parsimonious models for occurrence, UTI, and exploiter/avoider status, only the top occurrence model included a strong predictor for cophylogenetic congruence between birds and their haemosporidian parasites. The credible interval for the interaction between disturbed habitat occurrence and migratory strategy did not overlap zero, with fully migratory species that also use disturbed habitats having PACo residuals 0.026 greater than the baseline (Figure 2, Table 2). This suggests that host–parasite links associated with birds that are both long-distance migrants and are capable of occurring in anthropogenically disturbed habitats have somewhat higher residuals and contribute less to global cophylogenetic congruence.

Ecological traits are well-recognized to modify co-evolutionary relationships between host species and their parasites (Hafner et al., 1994; Page, 2003; Sweet et al., 2018), but the potential relationship between urban tolerance and cophylogenetic congruence has not been explored. By leveraging a global database of over 1,000 associations between avian hosts and their haemosporidian parasites, we demonstrate that a species’ ability to use anthropogenically modified habitats can interact with other relevant functional traits (herein migratory strategy) to influence host–parasite associations over evolutionary time. Specifically, fully migratory bird species known to occur in disturbed habitats on average have co-evolutionary histories with their haemosporidian parasites that are more strongly characterized by host-switching events (Figure 1). Past studies have yielded mixed support for migratory behavior influencing co-evolution between birds and their haemosporidian parasites (de Angeli Dutra et al., 2022; Jenkins et al., 2012). Both above studies recovered a significant signal of cospeciation between birds and their haemosporidia, which is consistent with our results. However, there is evidence that migratory behavior alone cannot predict cophylogenetic congruence for the genera Plasmodium and Haemoproteus (de Angeli Dutra et al., 2022), whereas migration can mediate host-switching in Leucocytozoon (Jenkins et al., 2012). Our results suggest this discordance could partly be explained by considering host habitat use. Migratory birds occurring in habitats classified as artificial or “disturbed” per the IUCN (e.g., land modified for agriculture, gardens, heavily degraded forests, and urban areas) had host–parasite links that contributed less to the global signal of cospeciation than birds restricted to natural habitats or birds capable of using the most disturbed habitat category (i.e, urban areas). Long-distance migration may therefore only facilitate a greater degree of host-switching (e.g., through ecological mechanisms such as migratory exposure; Figuerola & Green, 2000) if the migrant can also use habitats throughout its annual cycle that expose it to sufficient infection opportunities, such as habitats modified by humans. Tolerance for habitats which are disturbed but not fully urbanized being the most influential trait in our models could indicate that spatial heterogeneity is important for disrupting cophylogenetic congruence between birds and haemosporidian lineages. The intermediate disturbance hypothesis (IDH) posits that species diversity is highest at intermediate levels of disturbance or environmental change. Although the IDH is usually invoked in studies of plant and invertebrate communities, it has also garnered some empirical support in birds (Blair, 1996; McKinney, 2008). Low levels of anthropogenic development can increase the amount and variety of resources available to birds, increasing net species richness at the cost of native species richness (Blair, 1996). The effect of anthropogenic development on parasites, including haemosporidia, may also depend on their habitat’s position in the urban gradient (Ferraguti et al., 2023). A disturbed habitat could increase parasite prevalence by increasing host susceptibility and the proportion of competent host species in the community (Albery et al., 2022; Bichet et al., 2013; Ecke et al., 2022; Marm Kilpatrick et al., 2006), but highly urbanized habitats may suffer decreased parasite diversity because of homogenized host communities or the disrupted transmission of pathogens with complex life cycles. Habitats with an intermediate level of disturbance could instead promote parasite diversity via increased environmental heterogeneity while minimizing the characteristics of urban landscapes that are hostile to free-living parasite life stages or vectors (Calegaro-Marques & Amato, 2014). Migratory birds tolerant of habitat disturbance may therefore provide their parasites with more host-switching opportunities not only as a symptom of traversing large geographic distances during migration, but also by being exposed to a more diverse suite of host and parasite communities while breeding, overwintering, or stopping over in modified habitats. An additional consideration is how habitat disturbance could influence the abundance and competence of the arthropod vectors that transmit haemosporidia. Traits typical of anthropogenically modified habitats, such as urban heat island effects and the accumulation of stagnant water, can favor the presence of certain vectors (Araujo et al., 2015). For example, mosquito abundance, species richness, and community composition can differ across rural–urban gradients, with evidence that natural and disturbed habitats may harbor the greatest mosquito abundance and richness but that urbanized habitats attract high densities of urban-adapted species (Ferraguti et al., 2016). However, this pattern is not expected to be consistent across every haemosporidian vector. Both mosquitoes and biting midges, vectors of haemosporidia in the genera Plasmodium and Haemoproteus, have microhabitat requirements that can be accommodated by disturbed habitats (van Hoesel et al., 2019). Conversely, black fly vectors of haemosporidia in the genus Leucocytozoon are reliant on clean, running water and sufficient vegetation cover and are adapted for development and transmission in relatively cool environments (Fecchio et al., 2020; Suri et al., 2017). Differing microhabitat requirements of arthropod vectors may explain why migration alone is a substantial driver of host-switching in Leucocytozoon (Jenkins et al., 2012) but not in Plasmodium and Haemoproteus (de Angeli Dutra et al., 2022). We excluded Leucocytozoon lineages from our analyses owing to computational tractability in this study, but it is nonetheless important to investigate how these results could shift with pathogen and vector ecology. Urbanization is rapidly and drastically transforming the planet and the landscapes hosting its biodiversity, with the potential to alter and degrade host–parasite associations. We here tested the hypothesis that a bird species’ tolerance of anthropogenically modified habitats corresponds with decreased host specialization in haemosporidian parasites. We demonstrate that a species’ occurrence in habitats that are anthropogenically disturbed but not fully urbanized can influence its co-evolutionary relationship with haemosporidian parasites when the species is also a long-distance migrant. Tolerance for intermediate levels of habitat disturbance could therefore interact with host dispersal ability to increase host-switching opportunities and erode cophylogenetic congruence between birds and their haemosporidian parasites. Although the magnitude of this effect in our analyses was small (Figure 2), this finding represents a valuable step towards understanding how global change may impact co-evolution and, at finer ecological scales, patterns of host specialization in parasite communities across urban gradients. Author contributions Both authors contributed to the conception of the study, analyses, and drafting of the initial manuscript.

Support was provided from the National Science Foundation (DBI 2515340). We thank Marisa Szubryt for her assistance with phylogenetic reconstruction. We also thank Meagan Allira, Caroline Cummings, Kristin Dyer, Lauren Lock, Alicia Roistacher, Molly Simonis, and Amanda Vicente-Santos for their feedback on an earlier version of this manuscript. Data availability Raw data on host–parasite associations are available in the MalAvi database (Bensch et al., 2009), and avian trait data are available via AVONET (Tobias et al., 2022). The avian phylogeny was derived from BirdTree (Jetz et al., 2012). Mean species-level PACo residuals and associated trait data to replicate PGLMM analyses are provided in the Dryad Digital Repository: https://doi.org/10.5061/dryad.p8cz8wb20. Tables Table 1. Comparisons of PGLMMs differing in the metric of urban tolerance used (urban or disturbed habitat occurrence, urban tolerance index, or urban exploiter/avoider status). Model fit was assessed using the leave-one-out cross-validation information criterion (LOOIC), with the best-performing models represented by the lowest LOOIC. All models additionally include a phylogenetic random effect computed using the host phylogeny covariance matrix. | Model comparison | Fixed effects | Model weight | ΔLOOIC ( standard error ) | | Urban or disturbed habitat occurrence ( n=639 ) | Disturbed habitat occurrence * Migratory strategy + Research effort | 0.900 | 0 ( 48.82 ) | | Urban occurrence * Migratory strategy + Research effort | 0.018 | 5.32 ( 48.94 ) | | | Disturbed habitat occurrence + Migratory strategy + Research effort | 0.020 | 7.63 ( 48.75 ) | | | Urban occurrence + Migratory strategy + Research effort | 0.062 | 7.83 ( 48.81 ) | | | Urban tolerance index ( n=235 ) | Urban tolerance index * Migratory strategy + Research effort | 0.689 | 0 ( 28.09 ) | | Urban tolerance index + Migratory strategy + Research effort | 0.311 | 1.59 ( 28.06 ) | | | Urban exploiter/avoider status ( n=171 ) | Urban exploiter/avoider status * Migratory strategy + Research effort | 0.506 | 0 ( 20.50 ) | | Urban exploiter/avoider status + Migratory strategy + Research effort | 0.494 | 0.05 ( 20.64 ) | Table 2. Posterior means and 95% credible intervals for fixed effect coefficients in the top PGLMM for each of our three model comparisons (disturbed and urban habitat occurrence, urban tolerance index, and urban exploiter/avoider status). Bold indicates model coefficients where credible intervals do not overlap zero. | Model | Coefficient | Posterior mean | 95% credible interval | | Disturbed habitat occurrence * Migratory strategy + Research effort | Disturbed habitat occurrence | -0.003 | -0.01–0 | | Migratory strategy | -0.020 | -0.04–0 | | | Disturbed habitat occurrence * Migratory strategy | 0.026 | 0.01–0.05 | | | Number of citations | 0.001 | 0–0 | | | Urban tolerance index + Migratory strategy + Research effort | Urban tolerance index | -0.016 | -0.05–0.02 | | Migratory strategy | 0.001 | -0.02–0.02 | | | Number of citations | 0.002 | -0.01–0 | | | Urban exploiter/avoider status + Migratory strategy + Research effort | Urban exploiter/avoider status | -0.006 | -0.02–0.01 | | Migratory strategy | -0.002 | -0.02–0.02 | | | Number of citations | -0.001 | -0.01–0 | Figures and legends Figure 1. Tanglegram depicting host–parasite links between a phylogeny of birds (left) and of their haemosporidian parasites in the genera Plasmodium and Haemoproteus (right). Associations are colored to correspond with host species occurrence in modified habitats, with yellow links representing species occurring in anthropogenically disturbed habitats, red links representing species occurring in the most “severe” category of disturbance (urbanization), and green links representing species that are excluded from disturbed habitats. Illustrations are by Javier Lazaro (left) and Gediminas Valkiūnas (right). Figure 2. Posterior mean estimates and 95% credible intervals for fixed effects in the most competitive PGLMM of log-transformed mean jackknifed PACo residuals, which inform the contribution of host–parasite links to global cophylogenetic congruence (larger residual = less contribution to co-speciation). Predictors with credible intervals that do not overlap with zero are assumed to have a strong effect.

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Authors Metrics & Citations Metrics Article Usage 472views 269downloads Citations Download citation Taylor Verrett, Daniel Becker. The concrete tangle: Tolerance of anthropogenically disturbed habitats influences cophylogenetic congruence between migratory birds and their haemosporidian parasites. Authorea. 05 March 2025. DOI: https://doi.org/10.22541/au.174118532.28742495/v1 DOI: https://doi.org/10.22541/au.174118532.28742495/v1 If you have the appropriate software installed, you can download article citation data to the citation manager of your choice. Simply select your manager software from the list below and click Download. For more information or tips please see 'Downloading to a citation manager' in the Help menu.