Introducing 'Ngaruroro', a New Model for Understanding Maori Wellbeing.

Indigenous peoples around the world are revitalising their ancestral beliefs, practices, and languages, including traditional understandings of health and wellbeing. In the Aotearoa (New Zealand) context, a number of ground-breaking Maori health- and wellbeing-related models have emerged, each with their own scope and applications. We sought in our qualitative studies to explore and identify several key sources of wellbeing for Maori individuals. Nine interviews were conducted with members of Maori communities to identify key themes of Maori wellbeing. We performed a Reflexive Thematic Analysis on these data and then conducted a further fifteen interviews to revise, refine, and reposition the previously generated themes. The Ngaruroro model describes wellbeing as the embodied and active process of being well in relation with one's (1) here tangata (social and familial ties), (2) te taiao (the environment), and (3) taonga tuku iho (cultural treasures) while doing what one can to make lifestyle choices that are conducive to the health of one's (4) tinana (body) and (5) wairua (spirit) while cultivating a balanced (6) ngakau (inner-system), fulfilling (7) matea (core needs) and exercising your (8) mana (authority). These themes illustrate that Maori wellbeing is dynamic, interconnected, and holistic.

Exaggerated mandibles are correlated with enhanced foraging efficacy in male Auckland tree weta.

Sexual selection has driven the evolution of weaponry for males to fight rivals to gain access to females. Although weapons are predicted to increase males' reproductive success, they are also expected to incur costs and may impair functional activities, including foraging. Using feeding assays, we tested whether the enlarged mandibles of Auckland tree weta (Hemideina thoracica) impact feeding activity (the total volume of biomass consumed, bite rate, and number of foraging visits) and foraging behaviour (time spent moving, feeding, or stationary). We predicted that increased head capsule size in male weta would hinder their foraging efficacy. However, we found that weta with longer heads fed at a faster rate and spent less time foraging than weta with smaller heads, regardless of sex. Contrary to expectations that weapons impede functional activities, our results demonstrate that exaggerated traits can improve feeding performance and may offer benefits other than increased mating success.

Behavioural differences in predator aware and predator naïve Wellington tree weta, Hemideina crassidens.

Insects have evolved a wide range of behavioural traits to avoid predation, with anti-predator behaviours emerging as important adaptive responses to the specific strategies employed by predators. These responses may become ineffective, however, when a species is introduced to a novel predator type. When individuals cannot recognise an introduced predator for instance, they may respond in ways that mean they fail to avoid, escape, or neutralize a predator encounter. New Zealand's endemic insect fauna evolved in the absence of terrestrial mammalian predators for millions of years, resulting in the evolution of unique fauna like the large, flightless Orthopteran, the weta. Here we investigate how experience with introduced mammalian predators might influence anti-predator behaviours by comparing behaviours in a group of Wellington tree weta (Hemideina crassidens) living in an ecosanctuary, Zealandia, protected from non-native mammalian predators, and a group living in adjacent sites without mammalian predator control. We used behavioural phenotyping assays with both groups to examine rates of activity and defensive aggression shortly after capture, and again after a period of acclimation. We found that weta living in protected areas were more active shortly after capture than weta in non-protected habitats where mammalian predators were present. Male weta living in non-protected areas tended to be less aggressive than any other group. These results suggest that lifetime experience with differing predator arrays may influence the expression of antipredator behaviour in tree weta. Disentangling innate and experiential drivers of these behavioural responses further will have important implications for insect populations in rapidly changing environments.

Interpreting past trophic ecology of a threatened alpine parrot, kea Nestor notabilis, from museum specimens.

When ecosystems are under severe pressure or environments change, trophic position and intraspecific niche width may decrease or narrow, signalling that conservation action is required. In New Zealand, alpine and subalpine ecosystems have been extensively modified through farming since 19th-century European settlement, with consequences for indigenous species such as the kea Nestor notabilis. We investigated feather stable isotope values in the kea and predicted a lower trophic position in modern kea populations, to reflect reduced lowland habitat and a mixed diet with more plant material. We predicted that size and sex would influence trophic values in this sexually dimorphic species, with larger birds more likely to have a high protein diet. We examined potential dietary changes in 68 museum collected kea from 1880s to 2000s, first recording accession details including provenance and sex and measuring culmen length. We used bulk carbon and nitrogen stable isotope analyses (BSIAs) of feathers and a further feather subset using compound-specific stable isotope analyses of amino acids (CSIA-AA) to obtain isotopic values and estimate trophic position. BSIA showed δ15 N values in kea feathers declined through time and could indicate that early century kea were highly omnivorous, with δ15 N values on average higher than in modern kea. Variance in δ15 N values was greater after 1950, driven by a few individuals. Few differences between males and females were evident, although females in the south region had lower δ15 N values. There was a tendency for large male birds to have higher trophic values, perhaps reflecting dominant male bird behaviour noted in historical records. Nonetheless, CSIA-AA performed on a subset of the data suggested that variation in BSIA is likely due to baseline changes rather than relative trophic position which may be more homogenous than these data indicate. Although there was more variability in modern kea, we suggest caution in interpretation. Stable isotope data, particularly CSIA-AA, from museum specimens can reveal potential change in ecological networks as well as sexually dimorphic feeding patterns within species. The data can reveal temporal and regional variation in species trophic position and changes in ecosystem integrity to inform conservation decision-making.

Weaving place-based knowledge for culturally significant species in the age of genomics: Looking to the past to navigate the future.

Relationships with place provide critical context for characterizing biocultural diversity. Yet, genetic and genomic studies are rarely informed by Indigenous or local knowledge, processes, and practices, including the movement of culturally significant species. Here, we show how place-based knowledge can better reveal the biocultural complexities of genetic or genomic data derived from culturally significant species. As a case study, we focus on culturally significant southern freshwater koura (crayfish) in Aotearoa me Te Waipounamu (New Zealand, herein Aotearoa NZ). Our results, based on genotyping-by-sequencing markers, reveal strong population genetic structure along with signatures of population admixture in 19 genetically depauperate populations across the east coast of Te Waipounamu. Environment association and differentiation analyses for local adaptation also indicate a role for hydroclimatic variables-including temperature, precipitation, and water flow regimes-in shaping local adaptation in koura. Through trusted partnerships between community and researchers, weaving genomic markers with place-based knowledge has both provided invaluable context for the interpretation of data and created opportunities to reconnect people and place. We envisage such trusted partnerships guiding future genomic research for culturally significant species in Aotearoa NZ and beyond.

Human Perceptions of Megafaunal Extinction Events Revealed by Linguistic Analysis of Indigenous Oral Traditions.

Human settlement into new regions is typically accompanied by waves of animal extinctions, yet we have limited understanding of how human communities perceived and responded to such ecological crises. The first megafaunal extinctions in New Zealand began just 700 years ago, in contrast to the deep time of continental extinctions. Consequently, indigenous Maori oral tradition includes ancestral sayings that explicitly refer to extinct species. Our linguistic analysis of these sayings shows a strong bias towards critical food species such as moa, and emphasizes that Maori closely observed the fauna and environment. Temporal changes in form and content demonstrate that Maori recognized the loss of important animal resources, and that this loss reverberated culturally centuries later. The data provide evidence that extinction of keystone fauna was important for shaping ecological and social thought in Maori society, and suggest a similar role in other early societies that lived through megafaunal extinction events.

Diet selectivity in a terrestrial forest invertebrate, the Auckland tree weta, across three habitat zones.

Insects are important but overlooked components of forest ecosystems in New Zealand. For many insect species, information on foraging patterns and trophic relationships is lacking. We examined diet composition and selectivity in a large-bodied insect, the Auckland tree weta Hemideina thoracica, in three habitat zones in a lowland New Zealand forest. We asked whether H. thoracica selectively forage from available plant food sources, and whether these choices were lipid-rich compared to nonpreferred available plants. We also identified the proportion of invertebrates in their frass as a proxy for omnivory. From reconnaissance plot sampling, together with fecal fragment analysis, we report that more than 93% of individual tree weta had eaten invertebrates before capture. Additionally, weta in the highest elevation hillslope habitat zone consumed significantly fewer species of plants on average than weta on the low-elevation terrace habitat. Upper hillslope weta also had the highest average number of invertebrate fragments in their frass, significantly more than weta in the low-elevation terrace habitat zone. Weta showed high variability in the consumption of fruit and seeds across all habitat zones. Generally, we did not observe diet differences between the sexes (although it appears that male weta in the mid-hillslope habitat ate fruits and seeds more voraciously than females), suggesting that the sexes have similar niche breadths and display similar degrees of omnivorous behavior. Extraction of leaf lipids demonstrated a range of lipid content values in available plants, and Ivlev's Electivity Index indicated that plant species which demonstrated high electivity tended to have higher concentrations of lipids in their leaves. Our findings indicate that H. thoracica forage omnivorously and selectively, and hence play multiple roles in native ecosystems and food webs.

Importance of including cultural practices in ecological restoration.

Ecosystems worldwide have a long history of use and management by indigenous cultures. However, environmental degradation can reduce the availability of culturally important resources. Ecological restoration aims to repair damage to ecosystems caused by human activity, but it is unclear how often restoration projects incorporate the return of harvesting or traditional life patterns for indigenous communities. We examined the incorporation of cultural use of natural resources into ecological restoration in the context of a culturally important but protected New Zealand bird; among award-winning restoration projects in Australasia and worldwide; and in the peer-reviewed restoration ecology literature. Among New Zealand's culturally important bird species, differences in threat status and availability for hunting were large. These differences indicate the values of a colonizing culture can inhibit harvesting by indigenous people. In Australasia among award-winning ecological restoration projects, <17% involved human use of restored areas beyond aesthetic or recreational use, despite many projects encouraging community participation. Globally, restoration goals differed among regions. For example, in North America, projects were primarily conservation oriented, whereas in Asia and Africa projects frequently focused on restoring cultural harvesting. From 1995 to 2014, the restoration ecology literature contained few references to cultural values or use. We argue that restoration practitioners are missing a vital component for reassembling functional ecosystems. Inclusion of sustainably harvestable areas within restored landscapes may allow for the continuation of traditional practices that shaped ecosystems for millennia, and also aid project success by ensuring community support.

Cooperation-mediated plasticity in dispersal and colonization.

Kin selection theory predicts that costly cooperative behaviors evolve most readily when directed toward kin. Dispersal plays a controversial role in the evolution of cooperation: dispersal decreases local population relatedness and thus opposes the evolution of cooperation, but limited dispersal increases kin competition and can negate the benefits of cooperation. Theoretical work has suggested that plasticity of dispersal, where individuals can adjust their dispersal decisions according to the social context, might help resolve this paradox and promote the evolution of cooperation. Here, we experimentally tested the hypothesis that conditional dispersal decisions are mediated by a cooperative strategy: we quantified the density-dependent dispersal decisions and subsequent colonization efficiency from single cells or groups of cells among six genetic strains of the unicellular Tetrahymena thermophila that differ in their aggregation level (high, medium, and low), a behavior associated with cooperation strategy. We found that the plastic reaction norms of dispersal rate relative to density differed according to aggregation level: highly aggregative genotypes showed negative density-dependent dispersal, whereas low-aggregation genotypes showed maximum dispersal rates at intermediate density, and medium-aggregation genotypes showed density-independent dispersal with intermediate dispersal rate. Dispersers from highly aggregative genotypes had specialized long-distance dispersal phenotypes, contrary to low-aggregation genotypes; medium-aggregation genotypes showing intermediate dispersal phenotype. Moreover, highly aggregation genotypes showed evidence for beneficial kin-cooperation during dispersal. Our experimental results should help to resolve the evolutionary conflict between cooperation and dispersal: cooperative individuals are expected to avoid kin-competition by dispersing long distances, but maintain the benefits of cooperation by dispersing in small groups.

Tolerance for nutrient imbalance in an intermittently feeding herbivorous cricket, the Wellington tree weta.

Organisms that regulate nutrient intake have an advantage over those that do not, given that the nutrient composition of any one resource rarely matches optimal nutrient requirements. We used nutritional geometry to model protein and carbohydrate intake and identify an intake target for a sexually dimorphic species, the Wellington tree weta (Hemideina crassidens). Despite pronounced sexual dimorphism in this large generalist herbivorous insect, intake targets did not differ by sex. In a series of laboratory experiments, we then investigated whether tree weta demonstrate compensatory responses for enforced periods of imbalanced nutrient intake. Weta pre-fed high or low carbohydrate: protein diets showed large variation in compensatory nutrient intake over short (<48 h) time periods when provided with a choice. Individuals did not strongly defend nutrient targets, although there was some evidence for weak regulation. Many weta tended to select high and low protein foods in a ratio similar to their previously identified nutrient optimum. These results suggest that weta have a wide tolerance to nutritional imbalance, and that the time scale of weta nutrient balancing could lie outside of the short time span tested here. A wide tolerance to imbalance is consistent with the intermittent feeding displayed in the wild by weta and may be important in understanding weta foraging patterns in New Zealand forests.

Artefacts, biology and bias in museum collection research.

Museum collections are increasingly subjected to scientific scrutiny, including molecular, isotopic and trace-element analyses. Recent advances have extended analyses from natural history specimens to historical artefacts. We highlight three areas of concern that can influence interpretation of data derived from museum collections: sampling issues associated with museum collection use, methods of analysis, and the value of cross-referencing data with historical documents and data sets. We use a case study that focuses on kiwi (Apteryx spp.) feather samples from valuable 19th century Maori cloaks in New Zealand to show how sampling and analysis challenges need to be minimized by careful design. We argue that aligning historical records with scientific data generated from museum collections significantly improves data interpretation.

Isotopic fractionation in a large herbivorous insect, the Auckland tree weta.

Determining diet and trophic position of species with stable isotopes requires appropriate trophic enrichment estimates between an animal and its potential foods. These estimates are particularly important for cryptic foragers where there is little comparative dietary information. Nonetheless, many trophic enrichment estimates are based on related taxa, without confirmation of accuracy using laboratory trials. We used stable isotope analysis to investigate diet and to resolve trophic relationships in a large endemic insect, the Auckland tree weta (Hemideina thoracica White). Comparisons of isotopes in plant foods fed to captive wetas with isotope ratios in their frass provided variable results, so frass isotope values had limited usefulness as a proxy indicator of trophic level. Isotopic values varied between different tissues, with trophic depletion of (15)N highest in body fat and testes. Tissue fractionation was consistent in captive and wild caught wetas, and isotopic values were not significantly different between the two groups, suggesting that this weta species is primarily herbivorous. Whole-body values in captive wetas demonstrated trophic depletion (Δδ) for δ(15)N of about -0.77 ‰ and trophic enrichment of 4.28 ‰ for δ(13)C. These values differ from commonly estimated trophic enrichments for both insects and herbivores and indicate the importance of laboratory trials to determine trophic enrichment. Isotopic values for femur muscles from a number of local wild weta populations did not vary consistently with body weight or size, suggesting that juveniles eat the same foods as adults. Considerable variation among individuals within and between populations suggests that isotopic values are strongly influenced by food availability and individual foraging traits.

Traditional plant harvesting in contemporary fragmented and urban landscapes.

Ecosystem fragmentation and destruction can lead to restrictive administration policies on traditional harvesting by indigenous peoples from remaining ecosystem tracts. In New Zealand, concerns about endangered species and governmental policies that focus on species and ecosystem preservation have resulted in severely curtailed traditional harvesting rights. Although provision has been made for limited gathering of traditional plants from government-administered conservation lands, it is unclear how much harvesting is undertaken on these lands and elsewhere and what this harvest might consist of. We interviewed seven expert Maori elders from the Waikato, New Zealand, to identify plant species they currently harvested and from where. We compared these data with the data we collected on permits issued for plant collecting on conservation lands in the same region. We sought to gain information on indigenous plant harvesting to determine the extent of permitted harvesting from conservation lands in the Waikato and to identify issues that might affect plant harvesting and management. Elders identified 58 species they harvest regularly or consider culturally important; over 50% of these species are harvested for medicinal use. Permit data from 1996 to 2006 indicated no apparent relationship between species of reported cultural significance and the number of permits issued for each of these species. Currently, few plant species are harvested from conservation lands, although some unofficial harvesting occurs. Elders instead reported that medicinal plants are frequently collected from urban and other public areas. They reported that plant species used for dyeing, carving, and weaving are difficult to access. Elders also discussed concerns such as spraying of roadsides, which resulted in the death of medicinal species, and use of commercial hybrids in urban planning. Local government may have an increasingly important role in supporting native traditions through urban planning, which takes account of cultural harvesting needs while potentially reducing future harvesting pressure on conservation lands. We suggest that active participation by the Maori community in the development and management of urban harvesting resources will result in positive outcomes.

Indigenous ancestral sayings contribute to modern conservation partnerships: examples using Phormium tenax.

Traditional ecological knowledge (TEK) is central to indigenous worldviews and practices and is one of the most important contributions that indigenous people can bring to conservation management partnerships. However, researchers and managers may have difficulty accessing such knowledge, particularly where knowledge transmission has been damaged. A new methodological approach analyzes ancestral sayings from Maori oral traditions for ecological information about Phormium tenax, a plant with high cultural value that is a dominant component in many threatened wetland systems, and frequently used in restoration plantings in New Zealand. Maori ancestral sayings record an association with nectar-feeding native parrots that has only rarely been reported, as well as indications of important environmental parameters (rainfall and drought) for this species. These sayings provide evidence of indigenous management that has not been reported from interviews with elders, including evidence of fire use to create Phormium cultivations. TEK in Maori ancestral sayings imply landscape-scale processes in comparison to intensive, small-scale management methods often reported in interviews. TEK in ancestral sayings can be used to generate new scientific hypotheses, negotiate collaborative pathways, and identify ecological management strategies that support biodiversity retention. TEK can inform restoration ecology, historical ecology, and conservation management of species and ecosystems, especially where data from pollen records and archaeological artifacts are incomplete.