Papers by Patrick Kavanagh
Proceedings of The Royal Society B: Biological Sciences, Jul 7, 2014
Several plant traits are known to evolve in predictable ways on islands. For example, herbaceous ... more Several plant traits are known to evolve in predictable ways on islands. For example, herbaceous species often evolve to become woody and species frequently evolve larger leaves, regardless of growth form. However, our understanding of how seed sizes might evolve on islands lags far behind other plant traits. Here, we conduct the first test for macroevolutionary patterns of seed size on islands. We tested for differences in seed size between 40 island-mainland taxonomic pairings from four island groups surrounding New Zealand. Seed size data were collected in the field and then augmented by published seed descriptions to produce a more comprehensive dataset. Seed sizes of insular plants were consistently larger than mainland relatives, even after accounting for differences in growth form, dispersal mode and evolutionary history. Selection may favour seed size increases on islands to reduce dispersibility, as longdistance dispersal may result in propagule mortality at sea. Alternatively, larger seeds tend to generate larger seedlings, which are more likely to establish and outcompete neighbours. Our results indicate there is a general tendency for the evolution of large seeds on islands, but the mechanisms responsible for this evolutionary pathway have yet to be fully resolved.
First, I would like to thank my supervisor KC Burns. KC has been an amazing mentor, providing me ... more First, I would like to thank my supervisor KC Burns. KC has been an amazing mentor, providing me with support and direction while giving me enough independence to grow as a researcher. KC has been a great teacher, enlightened me to the ʻtacticsʼ of publishing, and instilled confidence through his famous ʻpep-talksʼ and amazing analogies (including references to NBA legends, fishing, skateboarding and surfing). I am grateful for my colleagues and friends in SBS. Mauricio Cifuentes,
Land ownership norms play a central role in social-ecological systems, and have been studied exte... more Land ownership norms play a central role in social-ecological systems, and have been studied extensively as a component of ethnographies. Yet only recently has the distribution of land ownership norms across cultures been examined from evolutionary and ecological perspectives. Here we incorporate evolutionary and macroecological modelling to test associations between land ownership norms and environmental, subsistence, and cultural contact predictors for societies in the Bantu language family. We find that Bantu land ownership norms likely evolved on a unilinear trajectory, but not necessarily one requiring consistent increase in exclusivity as suggested by prior theory. Our macroecological analyses suggest that Bantu societies are more likely to have some form of ownership when their neighbors also do. We also find an effect of environmental productivity, supporting resource defensibility theory, which posits that land ownership is more likely where productivity is predictable. We ...
SSRN Electronic Journal, 2018
This paper was prepared under Motu's Shaping New Zealand's Low-Emission Future programme, with fu... more This paper was prepared under Motu's Shaping New Zealand's Low-Emission Future programme, with funding support from the Ministry for Primary Industries and the Aotearoa Foundation. It was informed by the Motu Researcher Workshop on Land Use and Climate Change held in Wellington on 30 April 2018, and we gratefully acknowledge the contributions by presenters and other participants in that process. We also wish to acknowledge Susan Walker from Manaaki Whenua-Landcare Research, who reviewed parts of this paper. Disclaimer Contributions to the researcher workshop and this report were made in individuals' personal capacity and their inclusion in this paper does not imply any recommendation, consensus or endorsement by all participants, their affiliated organisations or the programme funders. While every effort has been made to ensure the information in this publication is accurate, the Ministry for Primary Industries does not accept any responsibility or liability for error of fact, omission, interpretation or opinion that may be present, nor for the consequences of any decisions based on this information.
Evolutionary Human Sciences, 2020
The evolution of agriculture improved food security and enabled significant increases in the size... more The evolution of agriculture improved food security and enabled significant increases in the size and complexity of human groups. Despite these positive effects, some societies never adopted these practices, became only partially reliant on them, or even reverted to foraging after temporarily adopting them. Given the critical importance of climate and biotic interactions for modern agriculture, it seems likely that ecological conditions could have played a major role in determining the degree to which different societies adopted farming. However, this seemingly simple proposition has been surprisingly difficult to prove and is currently controversial. Here, we investigate how recent agricultural practices relate both to contemporary ecological opportunities and the suitability of local environments for the first species domesticated by humans. Leveraging a globally distributed dataset on 1,291 traditional societies, we show that after accounting for the effects of cultural transmission and more current ecological opportunities, levels of reliance on farming continue to be predicted by the opportunities local ecologies provided to the first human domesticates even after centuries of cultural evolution. Based on the details of our models, we conclude that ecology probably helped shape the geography of agriculture by biasing both human movement and the human-assisted dispersal of domesticates.
Land ownership shapes natural resource management and social-ecological resilience, but the facto... more Land ownership shapes natural resource management and social-ecological resilience, but the factors determining ownership norms in human societies remain unclear. Here we conduct a global empirical test of long-standing theories from ecology, economics, and anthropology regarding potential drivers of land ownership and territoriality. Prior theory suggests that resource defensibility, subsistence strategies, population pressure, political complexity, and cultural transmission mechanisms may all influence land ownership. We applied multi-model inference procedures based on logistic regression to cultural and environmental data from 102 societies, 71 with some form of land ownership and 31 with no land ownership. We found an increased probability of land ownership in mountainous environments, where patchy resources may be more cost effective to defend via ownership. We also uncovered support for the role of population pressure, with a greater probability of land ownership in societies...
Social inequality is now pervasive in human societies, despite the fact that humans lived in rela... more Social inequality is now pervasive in human societies, despite the fact that humans lived in relatively egalitarian, small-scale societies across most of our history. Prior literature highlights the importance of environmental conditions, economic defensibility, and wealth transmission for shaping early Holocene origins of social inequality. However, it remains untested whether the mechanisms that drive the evolution of inequality in recent human societies follow a similar trajectory. We conduct the first global analysis of pathways to inequality within modern human societies using structural equation modeling. Our analytical approach demonstrates that environmental conditions, resource intensification, and wealth transmission mechanisms impact various forms of social inequality via a complex web of causality. We further find that subsistence practices have a direct impact on some institutionalized forms of inequality. This work identifies drivers of social inequality in the modern ...
Proceedings of the Royal Society B: Biological Sciences, 2019
Although many hypotheses have been proposed to explain why humans speak so many languages and why... more Although many hypotheses have been proposed to explain why humans speak so many languages and why languages are unevenly distributed across the globe, the factors that shape geographical patterns of cultural and linguistic diversity remain poorly understood. Prior research has tended to focus on identifying universal predictors of language diversity, without accounting for how local factors and multiple predictors interact. Here, we use a unique combination of path analysis, mechanistic simulation modelling, and geographically weighted regression to investigate the broadly described, but poorly understood, spatial pattern of language diversity in North America. We show that the ecological drivers of language diversity are not universal or entirely direct. The strongest associations imply a role for previously developed hypothesized drivers such as population density, resource diversity, and carrying capacity with group size limits. The predictive power of this web of factors varies ...
Royal Society Open Science, 2018
How humans obtain food has dramatically reshaped ecosystems and altered both the trajectory of hu... more How humans obtain food has dramatically reshaped ecosystems and altered both the trajectory of human history and the characteristics of human societies. Our species' subsistence varies widely, from predominantly foraging strategies, to plant-based agriculture and animal husbandry. The extent to which environmental, social and historical factors have driven such variation is currently unclear. Prior attempts to resolve long-standing debates on this topic have been hampered by an over-reliance on narrative arguments, small and geographically narrow samples, and by contradictory findings. Here we overcome these methodological limitations by applying multi-model inference tools developed in biogeography to a global dataset (818 societies). Although some have argued that unique conditions and events determine each society's particular subsistence strategy, we find strong support for a general global pattern in which a limited set of environmental, social and historical factors pr...
Biological Journal of the Linnean Society, 2016
Recent investigations have suggested that some plants are aposematic. Our understanding of how ap... more Recent investigations have suggested that some plants are aposematic. Our understanding of how aposematism varies through plant ontogeny, however, is incomplete. Furthermore, the potential for lower leaf surfaces to signal to vertebrate herbivores that are viewing leaves from below has not been investigated. Here, we investigate ontogenetic changes in leaf colour in Pseudopanax crassifolius (Araliaceae), a tree species that is endemic to New Zealand. We demonstrate that P. crassifolius produces lateral leaf spines that peak in size during the sapling stage of development. Spots of brightly coloured tissues on the upper leaf surfaces may be warning signals. The intensity of these signals, however, peaked at the seedling stage, providing a dishonest signal of defence. Conversely, signals on lower leaf surfaces peaked in the sapling stage, providing an honest defensive signal later in ontogeny. Lateral leaf spines and all potential warning colours were absent in adults, after they grow above the reach of the largest known native megaherbivores (moa-Aves: Dinornithiformes). Overall, these results suggest that aposematism may vary predictably through plant ontogeny in response to the changing perspective of herbivores as plants grow vertically.
The American Naturalist, 2011
Rensch's rule refers to a pattern in sexual size dimorphism (SSD) in which SSD decreases with bod... more Rensch's rule refers to a pattern in sexual size dimorphism (SSD) in which SSD decreases with body size when females are the larger sex and increases with body size when males are the larger sex. Many animal taxa conform to Rensch's rule, but it has yet to be investigated in plants. Using herbarium collections from New Zealand, we characterized the size of leaves and stems of 297 individuals from 38 dioecious plant species belonging to three distantly related phylogenetic lineages. Statistical comparisons of leaf sizes between males and females showed evidence for Rensch's rule in two of the three lineages, indicating SSD decreases with leaf size when females produce larger leaves and increases with leaf size when males produce larger leaves. A similar pattern in SSD was observed for stem sizes. However, in this instance, females of small-stemmed species produced much larger stems than did males, but as stem sizes increased, SSD often disappeared. We hypothesize that sexual dimorphism in stem sizes results from selection for larger stems in females, which must provide mechanical support for seeds, fruits, and dispersal vectors, and that scaling relationships in leaf sizes result from correlated evolution with stem sizes. The overall results suggest that selection for larger female stem sizes to support the weight of offspring can give rise to Rensch's rule in dioecious plants.
Proceedings of the Royal Society B: Biological Sciences, 2014
Several plant traits are known to evolve in predictable ways on islands. For example, herbaceous ... more Several plant traits are known to evolve in predictable ways on islands. For example, herbaceous species often evolve to become woody and species frequently evolve larger leaves, regardless of growth form. However, our understanding of how seed sizes might evolve on islands lags far behind other plant traits. Here, we conduct the first test for macroevolutionary patterns of seed size on islands. We tested for differences in seed size between 40 island–mainland taxonomic pairings from four island groups surrounding New Zealand. Seed size data were collected in the field and then augmented by published seed descriptions to produce a more comprehensive dataset. Seed sizes of insular plants were consistently larger than mainland relatives, even after accounting for differences in growth form, dispersal mode and evolutionary history. Selection may favour seed size increases on islands to reduce dispersibility, as long-distance dispersal may result in propagule mortality at sea. Alternati...
Biological Journal of the Linnean Society, 2012
Mistletoes are parasitic plants, the spatial distributions of which are poorly understood on macr... more Mistletoes are parasitic plants, the spatial distributions of which are poorly understood on macroecological scales. Because of their highly unusual life history, investigating mistletoe macroecology may provide new insight into broad-scale patterns in species distributions. We collated data on the spatial distribution and host use of 65 species of Loranthaceous mistletoes across the continent of Australia, and tested two predictions. First, we predicted mistletoe diversity would be unrelated to productivity (i.e. evapotranspiration and precipitation), as the parasitic lifestyle might relax environmental constraints on their distributions. Second, we predicted that mistletoe host ranges (number of infected host species) would increase in areas with more potential host species. The basis of this prediction is that greater host generality is likely to evolve in regions with greater host diversity because of greater unpredictability in encounter rates with particular host species. Conversely, in regions with fewer potential hosts, randomly dispersing mistletoe propagules are likely to repeatedly encounter particular host species, thus favouring the evolution of host specialization. The results were generally consistent with these predictions. Mistletoe diversity across Australia was weakly associated with environmental conditions, whereas mistletoe host ranges increased significantly with total plant diversity. Macroecological patterns in mistletoes are unusual. In contrast to non-parasitic plants, mistletoe diversity is poorly correlated with productivity. Host ranges varied predictably across Australia, providing the first quantitative support for the hypothesis that mistletoes in diverse regions tend to be host generalists, whereas mistletoes in depauperate regions tend to be host specialists.
Nature Human Behaviour, 2018
Several plant traits are known to evolve in predictable ways on islands. For example, herbaceous ... more Several plant traits are known to evolve in predictable ways on islands. For example, herbaceous species often evolve to become woody and species frequently evolve larger leaves, regardless of growth form. However, our understanding of how seed sizes might evolve on islands lags far behind other plant traits. Here, we conduct the first test for macroevolutionary patterns of seed size on islands. We tested for differences in seed size between 40 island–mainland taxonomic pairings from four island groups surrounding New Zealand. Seed size data were collected in the field and then augmented by published seed descriptions to produce a more comprehensive dataset. Seed sizes of insular plants were consistently larger than mainland relatives, even after accounting for differences in growth form, dispersal mode and evolutionary history. Selection may favour seed size increases on islands to reduce dispersibility, as long-distance dispersal may result in propagule mortality at sea. Alternatively, larger seeds tend to generate larger seedlings, which are more likely to establish and outcompete neighbours. Our results indicate there is a general tendency for the evolution of large seeds on islands, but the mechanisms responsible for this evolutionary pathway have yet to be fully resolved.
Mistletoes are parasitic plants, the spatial distributions of which are poorly understood on macr... more Mistletoes are parasitic plants, the spatial distributions of which are poorly understood on macroecological scales. Because of their highly unusual life history, investigating mistletoe macroecology may provide new insight into broad-scale patterns in species distributions. We collated data on the spatial distribution and host use of 65 species of Loranthaceous mistletoes across the continent of Australia, and tested two predictions. First, we predicted mistletoe diversity would be unrelated to productivity (i.e. evapotranspiration and precipitation), as the parasitic lifestyle might relax environmental constraints on their distributions. Second, we predicted that mistletoe host ranges (number of infected host species) would increase in areas with more potential host species. The basis of this prediction is that greater host generality is likely to evolve in regions with greater host diversity because of greater unpredictability in encounter rates with particular host species. Conversely, in regions with fewer potential hosts, randomly dispersing mistletoe propagules are likely to repeatedly encounter particular host species, thus favouring the evolution of host specialization. The results were generally consistent with these predictions. Mistletoe diversity across Australia was weakly associated with environmental conditions, whereas mistletoe host ranges increased significantly with total plant diversity. Macroecological patterns in mistletoes are unusual. In contrast to non-parasitic plants, mistletoe diversity is poorly correlated with productivity. Host ranges varied predictably across Australia, providing the first quantitative support for the hypothesis that mistletoes in diverse regions tend to be host generalists, whereas mistletoes in depauperate regions tend to be host specialists.
Many island plants are characterized by unique morphology. For example, the high branching angles... more Many island plants are characterized by unique morphology. For example, the high branching angles and small leaves of divaricate plants are a common feature of the New Zealand flora. The divaricate growth form may be an adaptation to deter browsing by extinct avian herbivores (moa); alternatively aspects of the insular climate may be responsible. However, our understanding of the selective pressures responsible for the high branching angles and small leaves of divaricate plants is incomplete. Here, I tested for differences in traits associated with the divaricate growth form between plants from Chatham Island and the New Zealand mainland. Moa never reached the Chatham Islands and its flora is derived from plants on mainland New Zealand. Therefore, I predicted Chatham Island plants to have lost morphological adaptations that may have deterred moa herbivory. Traits were quantified on 316 individuals in the field, allowing for 12 island-mainland taxonomic comparisons. Chatham Island plants consistently produced smaller branching angles, larger leaves, shorter internodes and larger stems than related mainland plants. Results are therefore consistent with the hypothesis that selection for small leaves and high angled branching may be relaxed on the Chatham Islands due to an absence of moa. Smaller branching angles and larger leaves may offer a competitive advantage to Chatham Island plants.
The niche variation hypothesis predicts insular populations exhibit increased sexual size dimorph... more The niche variation hypothesis predicts insular populations exhibit increased sexual size dimorphism (SSD), to minimize intraspecific competition. Although many animal taxa conform to this prediction, insular patterns of SSD have yet to be investigated in plants. Here, we tested for differences in SSD of dioecious plants that colonised four island groups (Kermadec, Three Kings, Chatham and Auckland Islands) from New Zealand. Using herbarium collections, we quantified leaf and stem sizes of 263 individuals from 28 dioecious taxa. We developed a novel analytical technique to explore changes in the direction of SSD on islands. Lastly, we tested for evolutionary size changes of male and female plants on islands. The degree of SSD did not vary predictably between insular and mainland taxa, contrary to predictions of the niche variation hypothesis. Furthermore, the direction of SSD was not predictable on islands, while it was consistently female biased on the mainland. Our results suggest that selection favours increased size of both sexes on islands and that SSD is unpredictable for insular plants.
JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, a... more JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. abstract: Rensch's rule refers to a pattern in sexual size dimorphism (SSD) in which SSD decreases with body size when females are the larger sex and increases with body size when males are the larger sex. Many animal taxa conform to Rensch's rule, but it has yet to be investigated in plants. Using herbarium collections from New Zealand, we characterized the size of leaves and stems of 297 individuals from 38 dioecious plant species belonging to three distantly related phylogenetic lineages. Statistical comparisons of leaf sizes between males and females showed evidence for Rensch's rule in two of the three lineages, indicating SSD decreases with leaf size when females produce larger leaves and increases with leaf size when males produce larger leaves. A similar pattern in SSD was observed for stem sizes. However, in this instance, females of small-stemmed species produced much larger stems than did males, but as stem sizes increased, SSD often disappeared. We hypothesize that sexual dimorphism in stem sizes results from selection for larger stems in females, which must provide mechanical support for seeds, fruits, and dispersal vectors, and that scaling relationships in leaf sizes result from correlated evolution with stem sizes. The overall results suggest that selection for larger female stem sizes to support the weight of offspring can give rise to Rensch's rule in dioecious plants.
Mistletoes are parasitic plants, the spatial distributions of which are poorly understood on macr... more Mistletoes are parasitic plants, the spatial distributions of which are poorly understood on macroecological scales. Because of their highly unusual life history, investigating mistletoe macroecology may provide new insight into broad-scale patterns in species distributions. We collated data on the spatial distribution and host use of 65 species of Loranthaceous mistletoes across the continent of Australia, and tested two predictions. First, we predicted mistletoe diversity would be unrelated to productivity (i.e. evapotranspiration and precipitation), as the parasitic lifestyle might relax environmental constraints on their distributions. Second, we predicted that mistletoe host ranges (number of infected host species) would increase in areas with more potential host species. The basis of this prediction is that greater host generality is likely to evolve in regions with greater host diversity because of greater unpredictability in encounter rates with particular host species. Conversely, in regions with fewer potential hosts, randomly dispersing mistletoe propagules are likely to repeatedly encounter particular host species, thus favouring the evolution of host specialization. The results were generally consistent with these predictions. Mistletoe diversity across Australia was weakly associated with environmental conditions, whereas mistletoe host ranges increased significantly with total plant diversity. Macroecological patterns in mistletoes are unusual. In contrast to non-parasitic plants, mistletoe diversity is poorly correlated with productivity. Host ranges varied predictably across Australia, providing the first quantitative support for the hypothesis that mistletoes in diverse regions tend to be host generalists, whereas mistletoes in depauperate regions tend to be host specialists.
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Papers by Patrick Kavanagh