Aim: The Sunda pangolin (Manis javanica) is the most widely distributed Asian pangolin species. It is one of the most trafficked mammals in the world, which not only negatively impacts wild Sunda pangolin populations, but also poses a potential disease risk to other species, including humans and livestock. Despite the imminent threat to the species’ survival and its prevalence in the wildlife trade, the phylogeography and evolution of the Sunda pangolin is not well understood. We aimed to investigate the species’ phylogeography across its distribution to improve our understanding of the species’ evolutionary history, elucidate any taxonomic uncertainties and enhance the species’ conservation genetic management and wildlife forensics applications. Location: Southeast Asia and southern China. Methods: We sequenced mtDNA genomes from 23 wild Sunda pangolins from Borneo and Peninsular Malaysia. We used these data in conjunction with previous generated mtDNA and nuclear datasets from across the species’ range to perform various phylogenetic and population genetic analyses. Results: We identified an evolutionarily distinct mtDNA lineage in north Borneo, which was estimated to be ~1.6 million years divergent from lineages in west/south Borneo and the mainland, comparable to the divergence time of the Palawan pangolin. There appeared to be mitonuclear discordance, with no apparent genetic structure across Borneo based on analysis of nuclear SNPs. Main conclusions: These findings are consistent with the ‘out of Borneo hypothesis’, whereby Sunda pangolins diversified in Borneo before subsequently migrating throughout Sundaland, and/or a secondary contact scenario between mainland and Borneo. We have elucidated possible taxonomic issues in the Sunda/Palawan pangolin complex, and highlight the critical need for additional georeferenced samples to accurately apportion its range-wide genetic variation into appropriate taxonomic and conservation units. Additionally, these data have improved forensic species identification testing involving these species and permit the implementation of geographic provenance testing in some scenarios.

Scale and tempo of brain expansion in the course of human evolution implies that this process was driven by a positive feedback. The ‘cultural drive’ hypothesis suggests a possible mechanism for the runaway brain-culture coevolution wherein high-fidelity social learning results in accumulation of cultural traditions which, in turn, promote selection for still more efficient social learning. Here we explore this evolutionary mechanism by means of computer modeling. Simulations confirm its plausibility in a social species in a socio-ecological situation that makes the sporadic invention of new beneficial and cognitively demanding behaviours possible. The chances for the runaway brain-culture coevolution increase when some of the culturally transmitted behaviours are individually beneficial while the others are group-beneficial. In this case, ‘cultural drive’ is possible under varying levels of between-group competition and migration. Modeling implies that brain expansion can receive additional boost if the evolving mechanisms of social learning are costly in terms of brain expansion (e.g., rely on complex neuronal curcuits) and tolerant to the complexity of information transferred, that is, make it possible to transfer complex skills and concepts easily. Human language presumably fits this description. Modeling also confirms that the runaway brain-culture coevolution can be accelerated by additional positive feedback loops via population growth and lifespan extension, and that between-group competition and cultural group selection can facilitate the propagation of group-beneficial behaviours and remove maladaptive cultural traditions from the population’s culture, which individual selection is unable to do.

Competition between the sympatric harbour (Phoca vitulina) and grey seals (Halichoerus grypus) is thought to underlie some recent local declines of the former while the population of the latter remains stable or increases. A better understanding of the interactions between these two species is critical to elucidate current changes. This study aims at identifying and quantifying the niche overlap between harbour and grey seals at their Southern European limit range, in the baie de Somme (Eastern English Channel, France), in a context of exponential increase in the number of resident harbour seals and visiting grey seals. Isotopic niche overlap was quantified between both species using whisker δ13C and δ15N isotopic values, taking intra- and interindividual variability into account. Dietary overlap was quantified from scat contents using hierarchical clustering. A high degree of trophic niche overlap was identified between both species. The narrower isotopic niche of harbour seals was nested within that of grey seals (58.2% [CI95%: 22.7-100%] overlap). Six diet clusters were identified from scat content analysis. Two of them gathered most of harbour seals’ scats (85.5 % [80.3-90.2%]) and around half of grey seals’ ones (46.8% [35.1-58.4%]) that almost exclusively contained benthic flatfish. Consumption of this type of prey was identified here to be the root cause of trophic overlap. This highlighted the potential for competition between the two species at their Southern European limit range, linked to foraging on benthic flatfish, in coastal waters close to their haulout sites, especially during spring/summer. We suggest that (1) interspecific competition for prey could occur/increase in the future if the number of grey and harbour seals still increase and/or if flatfish supply decrease in this area, and (2) harbour seals would be disadvantaged in such a case if they do not adapt, as being specialised on flatfish at the colony scale.

Parasites form an integral part of food webs, however, mechanistic insights into the role of parasites for energy flow and community dynamics is currently limited by a lack of conceptual studies investigating host-parasite interactions in a community context. In aquatic systems, chytrids constitute a major group of fungal parasites and their free-living infective stage (zoospores) forms a highly nutritional food source to zooplankton. Consumption of zoospores can create an energy pathway from otherwise inedible phytoplankton to zooplankton (“mycoloop”). The impact of such parasite-mediated energy pathways on community dynamics and energy transfer to higher trophic levels is of high importance considering eutrophication and global warming induced shifts to dominance of unfavourable prey such as cyanobacteria. We theoretically investigated community dynamics and energy transfer in a food web consisting of an edible-nonhost and an inedible-host phytoplankton species, a fungal parasite, and a zooplankton species grazing on edible phytoplankton and fungi. Food web dynamics were investigated along a nutrient gradient for two cases: (1) non-adaptive zooplankton species representative for filter feeders like cladocerans and (2) zooplankton with the ability to actively adapt their feeding preferences like many copepod species. For both feeding strategies, the importance of the mycoloop for zooplankton increases with nutrient availability. This increase is smooth for non-adaptive consumers. For a consumer with an adaptive feeding preference, we observe an abrupt shift from almost exclusive preference for edible phytoplankton (dominant prey) at low nutrient levels to a strong preference for parasitic fungi at high nutrient levels. The model predicts that parasitic fungi can contribute up to 50% of the zooplankton diet in nutrient rich environments, agreeing with empirical observations on zooplankton gut content from eutrophic systems during cyanobacterial blooms. Our findings highlight the role of parasite-mediated energy pathway for predictions on energy flow and community composition under environmental change.

Many populations of long-distance migrant shorebirds are declining rapidly. Since the 1970s, the Lesser Yellowlegs (Tringa flavipes) has experienced a pronounced reduction in abundance of ~63%. The potential causes of the species’ decline are complex and interrelated yet understanding the timing of migration and seasonal routes used by this species will aid in directing conservation planning to address potential threats. During 2018–2021, we tracked 118 adult Lesser Yellowlegs using GPS satellite tags deployed on birds from five breeding and two migratory stopover locations spanning the boreal forest of North America from Alaska to eastern Canada. Our objectives were to quantify migratory connectivity and identify key stopover and non-breeding locations. Individuals tagged in Alaska and central Canada followed similar southbound migratory routes through the Prairie Pothole Region of North America, whereas birds tagged in eastern Canada completed multi-day transoceanic flights covering distances of >4,000 km across the Atlantic between North and South America. Upon reaching their non-breeding locations, Lesser Yellowlegs populations overlapped, resulting in weak migratory connectivity. Lastly, freshwater and agricultural habitats of the Prairie Pothole region supported the highest proportion of Lesser Yellowlegs during southbound migration. Our findings suggest that while Lesser Yellowlegs travel long distances and traverse numerous political boundaries each year, the breeding population from which an individual originates likely has the greatest influence on which threats birds experience during migration. Further, the species’ dependence on wetlands in agricultural landscapes during migration may make them vulnerable to threats related to agricultural practices, such as pesticide exposure.

Insect decline is a major threat for ecosystems around the world as they provide many important functions, such as pollination or pest control. Pollution is one of the main reasons for the decline, besides changes in land use, global warming, and invasive species. While negative impacts of pesticides are well studied, there is still a lack of knowledge about the effects of other anthropogenic pollutants, such as airborne particulate matter, on insects. To address this, we exposed workers of the bumblebee Bombus terrestris to sublethal doses of diesel exhaust particles (DEPs) and brake dust, orally or via air. After seven days, we looked at the composition of the gut microbiome and tracked changes in gene expression. While there were no changes in the other treatments, oral DEP exposure significantly altered the structure of the gut microbiome. In particular, the core bacterium Snodgrassella had a decreased abundance in the DEP treatment. Similarly, transcriptome analysis revealed changes in gene expression after oral DEP exposure, but not in the other treatments. The changes are related to metabolism and signal transduction which indicates a general stress response. Taken together, our results suggest potential health effects of DEP exposure on insects, here shown in bumblebees, as gut dysbiosis may increase the susceptibility of bumblebees to pathogens, while a general stress response may lower available energy resources. However, experiments with multiple stressors and on colony level are needed to provide a more comprehensive understanding of the impact of DEPs on insects.

Resistance (host capacity to reduce parasite burden) and tolerance (host capacity to reduce impact on its health for a given parasite burden) manifest two different lines of defence. Tolerance can be independent from resistance, traded-off against it, or the two can be positively correlated because of redundancy in underlying (immune) processes. We here tested whether this coupling between tolerance and resistance could differ upon infection with closely related parasite species. We tested this in experimental infections with two parasite species of genus Eimeria. We measured proxies for resistance (the (inverse of) number of parasite transmission stages (oocysts) per gram of feces at the day of maximal shedding) and tolerance (the slope of maximum relative weight loss compared to day of infection on number of oocysts per gram of feces at the day of maximal shedding for each host strain) in four inbred mouse strains and four groups of F1 hybrids belonging to two mouse subspecies, Mus musculus domesticus and M. m. musculus. We found a negative correlation between resistance and tolerance against E. falciformis, while the two are uncoupled against E. ferrisi. We conclude that resistance and tolerance against the first parasite species might be traded off, but evolve more independently in different mouse genotypes against the latter. We argue that evolution of the host immune defences can be studied largely irrespective of parasite isolates if resistance-tolerance coupling is absent or weak (E. ferrisi) but host-parasite coevolution is more likely observable and best studied in a system with negatively correlated tolerance and resistance (E. falciformis).

Understanding the factors that regulate the functioning of our ecosystems in response to environmental changes can help to maintain the stable provisioning of ecosystem services to mankind. This is especially relevant given the increased variability of environmental conditions due to human activities. In particular, maintaining a stable production and plant biomass during the growing season (intra-annual stability) despite pervasive and directional changes in temperature and precipitation through time can help to secure food supply to wild animals, livestock, and humans. Here, we conducted a 29-year field observational study in a temperate grassland to explore how the intra-annual stability of primary productivity is influenced by biotic and abiotic variables through time. In particular, we analyzed the relationship of community biomass intra-annual stability with plant diversity and seasonal distribution patterns of temperature and precipitation. We found that lower accumulated precipitation between June and September during the 29-year investigated contributed to lower intra-annual community stability because of a decrease in compensatory mechanisms among species (species asynchrony). Additionally, higher precipitation in July contributed to higher intra-annual stability because higher species richness with higher precipitation led to higher average intra-annual stability of all species in the community (species stability). In contrast, we found no evidence that temperature influenced community intra-annual stability. Our results indicates that ongoing reduced seasonal precipitation leading to reduced intra-annual stability in the temperate grassland, which has important theoretical significance for us to take active measures to deal with climate change.

The sharp rise in anthropogenic activities and climate change have caused the extensive degradation of grasslands worldwide, jeopardising ecosystem function and threatening human well-being. Toxic weeds have been constantly spreading in recent decades; indeed, their occurrence is considered to provide an early sign of land degeneration. Policy makers and scientific researchers often focus on the negative effects of toxic weeds, such as how they inhibit forage growth, kill livestock and cause economic losses. However, toxic weeds can have several potentially positive ecological impacts on grasslands, such as promoting soil and water conservation, improving nutrient cycling and biodiversity conservation, and protecting pastures from excessive damage by livestock. We reviewed the literature to detail the adaptive mechanisms underlying toxic weeds and to provide new insight into their roles in degraded grassland ecosystems. The findings highlight that the establishment of toxic weeds may provide a self-protective strategy of degenerated pastures that does not require special interventions. Consequently, policy makers, managers and other personnel responsible for managing grasslands need to take appropriate actions to assess the long-term trade-offs between the development of animal husbandry and the maintenance of ecological services provided by grasslands.

1.Large carnivore conservation is complex and remains a massive challenge across the world. Owing to their wide-ranging habits, large carnivores encounter various anthropogenic pressures which may potentially lead to conflict. Animal movement is linked with individual fitness as it is important for various biological processes. Therefore, studying how large carnivores adapt their movement to dynamic landscape conditions is vital for management and conservation policy. 2.We first quantified the movement parameters of four large carnivores in and outside protected-areas in India (tiger, leopard, dhole and wolf). We then tested the effects of human pressures like human density, road density and land use types on the movement of the species. Finally, we examined the configuration of core areas as a strategy to exploit human-dominated landscape. 3.Our findings suggest that the mean hourly displacement of 4 large carnivores differed across habitats. Mean displacement of large carnivores varied from 77.58m/h for leopards to 665.3m/h for wolves. Tigers outside PAs exhibited higher displacement as compared to tigers inside PAs. Displacement during day and night were significantly different for tigers inside and outside PAs (P=0.03), and wolf whereas no difference was found for leopard and dholes. The movement and ranging patterns of species outside PAs were influenced by anthropogenic factors such as human population, road network density, and landuse. All carnivores showed multiple areas of intensive use or cores in their home ranges. The range of the core area sizes was greater for species outside PAs (tiger and wolf) in human-altered landscapes. 4.Movement ecology of large carnivores has not been explored using such an exhaustive dataset in India. Our study attempts to extend theoretical concepts to applied management problems. This study can be a starting point for rigorous studies on interlinking animal movement and landscape management for large carnivore conservation and policy-making in the Anthropocene.

Changing climatic conditions and unsustainable land use are perceived as major threats to savannas worldwide. In the past, land use in African savannas was dominated by livestock-farming as one of the major economic products, which led to degraded, shrub encroached pastures in many regions. One response to this widespread degradation is a shift from land use dominated by cattle to strategies characterized by animal compositions with more mixed feeding regimes and higher browser proportions. However, the consequences for ecosystem properties and processes remain so far largely unclear. We used the ecohydrological, spatially explicit savanna model EcoHyD to assess the impacts of two contrasting, herbivore-related land use strategies on a Namibian savannah: grazing versus browsing herbivores. We varied the densities of grazers and browsers and determined the resulting composition and diversity of the plant community, total vegetation cover, soil moisture and water use by plants. Our results show that properties making plants less attractive to herbivores were best adapted to different densities of grazing (cattle) or browsing (pure browsing wildlife) animals. Also, properties leading to a competitive advantage under limited water availability were among the dominant ones. Overall, the results are in line with our expectations: we found heavy shrub encroachment with a loss of the perennial grass matrix under high stocking rates of cattle. A novel and unexpected result was that regardless of the density of browsers, grass cover and plant functional diversity were significantly higher in wildlife scenarios. This increased grass cover, but also the higher total cover improved water uptake by plants. We conclude that in contrast to grazers, browsers even in high densities do not lead to ecosystem degradation, but rather sustain a diverse vegetation with high cover of perennial grasses over a long time, implying also a lower erosion risk and higher provision of ecosystem services.

1. Fruit bats (Family: Pteropodidae) are animals of great ecological and economic importance, yet their populations are threatened by ongoing habitat loss and human persecution. A lack of ecological knowledge for the vast majority of Pteropodid bat species presents additional challenges for their conservation and management. 2. In Australia, populations of flying-fox species (Genus: Pteropus) are declining and management approaches are highly contentious. Australian flying-fox roosts are exposed to management regimes involving habitat modification, either through human-wildlife conflict management policies, or vegetation restoration programs. Details on the fine-scale roosting ecology of flying-foxes are not sufficiently known to provide evidence-based guidance for these regimes and the impact on flying-foxes of these habitat modifications is poorly understood. 3. We seek to identify and test commonly held understandings about the roosting ecology of Australian flying-foxes to inform practical recommendations and guide and refine management practices at flying-fox roosts. 4. We identify 31 statements relevant to understanding of flying-fox roosting structure, and synthesise these in the context of existing literature. We then contribute contemporary data on the fine-scale roosting structure of flying-fox species in south-eastern Queensland and north-eastern New South Wales, presenting a 13-month dataset from 2,522 spatially referenced roost trees across eight sites. 5. We show evidence of sympatry and indirect competition between species, including spatial segregation of black and grey-headed flying-foxes within roosts and seasonal displacement of both species by little red flying-foxes. We demonstrate roost-specific annual trends in occupancy and abundance and provide updated demographic information including the spatial and temporal distributions of males and females within roosts. 6. Insights from our systematic and quantitative study will be important to guide evidence-based recommendations on restoration and management and will be crucial for the implementation of priority recovery actions for the preservation of these species into the future.

In the Anopheles genus, various mosquito species are able to transmit Plasmodium parasites responsible for malaria, while others are non-vectors. In an effort to better understand the biology of Anopheles species and to quantify transmission risk in an area, the identification of mosquito species collected on the field is an essential but problematic task. Morphological identification requires expertise, well-preserved specimens and high-quality equipment, and it does not allow any subsequent verification when samples are later used in a destructive treatment. Moreover, it involves physical manipulations that are not compatible with experiments requiring fast sampling and processing of specimens, hence species identification is often based on DNA sequencing of reference genes or region such as the Internal Transcribed Spacer 2 (ITS2) region of nuclear ribosomal DNA. Sequencing ITS2 for numerous samples is costly, but the design of species-specific PCR primers is not always possible when local species diversity is high. Here, we introduce a molecular technique of species identification based on precise determination of ITS2 length combined with a simple visual observation, the color of mosquito hindleg tip. DNA extracted from field-collected Anopheles mosquitoes was amplified with universal Anopheles ITS2 primers and analyzed with a capillary electrophoresis device, which precisely determines the size of the fragments. We defined windows of amplicon sizes combined with fifth hind tarsus color, which allow to discriminate the major Anopheles species found in our collections. We validated our parameters via Sanger sequencing of the ITS2 amplicons. This method can be particularly useful in situations with a moderate species diversity, i.e. when the number of local species is too high to define species-specific primers but low enough to avoid individual ITS2 sequencing. This tool will be of interest to evaluate local malaria transmission risk and this approach may further be implemented for other mosquito genera.

Understanding the drivers of morphological convergence requires investigation into its relationship with behavior and niche-space, and such investigations in turn provide insights into evolutionary dynamics, functional morphology, and life history. Mygalomorph spiders (trapdoor spiders and their kin) have long been associated with high levels of homoplasy, and many convergent features can be intuitively associated with different behavioral niches. Using genus-level phylogenies based on recent genomic studies and a newly assembled matrix of discrete behavioral and somatic morphological characters, we reconstruct the evolution of burrowing behavior in the Mygalomorphae, compare the influence of behavior and evolutionary history on somatic morphology, and test hypotheses of correlated evolution between specific morphological features and behavior. Our results reveal the simplicity of the mygalomorph adaptive landscape, with opportunistic, web-building taxa at one end, and burrowing/nesting taxa with structurally-modified burrow entrances (e.g., a trapdoor) at the other. Shifts in behavioral niche, in both directions, are common across the evolutionary history of the Mygalomorphae, and several major clades include taxa inhabiting both behavioral extremes. Somatic morphology is heavily influenced by behavior, with taxa inhabiting the same behavioral niche often more similar morphologically than more closely-related but behaviorally-divergent taxa, and we were able to identify a suite of 11 somatic features that show significant correlation with particular behaviors. We discuss these findings in light of the function of particular morphological features, niche dynamics within the Mygalomorphae, and constraints on the mygalomorph adaptive landscape relative to other spiders.

The current COVID-19 pandemic has forced the global higher education community to rapidly adapt to partially- or fully-online course offerings. For field- or lab-based courses in ecological curricula, this presents unique challenges. Fortunately, a diverse set of active learning techniques exist, and these techniques translate well to online settings. However, limited guidance and resources exist for developing, implementing, and evaluating active learning assignments that fulfil specific objectives of ecology-focused courses. To address these informational gaps, we (1) identify broad learning objectives across a variety of ecology-focused courses, (2) provide examples, based on our collective online teaching experience, of active learning activities that are relevant to the identified ecological learning goals, and (3) provide guidelines for successful implementation of active learning assignments in online courses. Using The Wildlife Society’s list of online higher education ecology-focused courses as a guide, we obtained syllabi from 45 ecology-focused courses, comprising a total of 321 course-specific learning objectives. We classified all course-specific learning objectives into at least one of five categories: (1) Identification, (2) Application of Concepts/Hypotheses/Theories, (3) Management of Natural Resources, (4) Development of Professional Skills, or (5) Evaluation of Concepts/Practices. We then provided two examples of active learning activities for each of the five categories, along with guidance on their implementation in online settings. We suggest that, when based on sound pedagogy, active learning techniques can enhance the online student’s experience by activating ecological knowledge; moreover, active learning techniques should also be incorporated into in-person offerings once the current COVID-19 crisis has abated.

Scientists are increasingly using volunteer efforts of citizen scientists to classify images captured by motion-activated trail-cameras. The rising popularity of citizen science reflects its potential to engage the public in conservation science and accelerate processing of the large volume of images generated by trail-cameras. While image classification accuracy by citizen scientists can vary across species, the influence of other factors on accuracy are poorly understood. Inaccuracy diminishes the value of citizen science derived data and prompts the need for specific best practice protocols to decrease error. We compare the accuracy between three programs that use crowdsourced citizen scientists to process images online: Snapshot Serengeti, Wildwatch Kenya, and AmazonCam Tambopata. We hypothesized that habitat type and camera settings would influence accuracy. To evaluate these factors, each photo was circulated to multiple volunteers. All volunteer classifications were aggregated to a single best answer for each photo using a plurality algorithm. Subsequently, a subset of these images underwent expert review and were compared to the citizen scientist results. Classification errors were categorized by the nature of the error (e.g. false species or false empty), and reason for the false classification (e.g. misidentification). Our results show that Snapshot Serengeti had the highest accuracy (97.9%), followed by AmazonCam Tambopata (93.5%), then Wildwatch Kenya (83.4%). Error type was influenced by habitat, with false empty images more prevalent in open-grassy habitat (27%) compared to woodlands (10%). For medium to large animal surveys across all habitat types, our results suggest that to significantly improve accuracy in crowdsourced projects, researchers should use a trail-camera set up protocol with a burst of three consecutive photos, a short field of view, and consider appropriate camera sensitivity. Accuracy level comparisons such as this study can improve reliability of future citizen science projects, and subsequently encourage the increased use of such data.

Sex is one of the important factors affecting gut microbiota. As key predators in agro-forestry ecosystem, many spider species show dramatically different activity habits and nutritional requirements between female and male. However, how sex affects gut microbiota of spiders is still unclear. Therefore, in this study, the compositions and diversities of gut bacteria, based on bacterial 16S rRNA gene sequencing, were compared between female and male Pardosa astrigera. We found that bacterial richness indices (P < 0.05) in female were significantly lower than male, meanwhile, β-diversity showed significantly different between female and male (P < 0.05). The relative abundance of Actinobacteriota and Rhodococcus (belongs to Actinobacteria) were significantly higher in female than male (P < 0.05). Whereas, the relative abundance of Firmicutes and Acinetobacter (belongs to Proteobacteria), Ruminococcus and Fusicatenibacter (all belong to Firmicutes), were significantly higher in male than female (P < 0.05). The results of PICRUSt2 showed that amino acid and lipid metabolisms were significantly higher in female than male (P < 0.05), whereas glycan biosynthesis and metabolism was significantly higher in male than female (P < 0.05). Our results imply that sexual variation is a crucial factor in shaping gut bacterial community in P. astrigera. Male P. astrigera dispersed more widely than the female hence the male had a higher bacterial diversity. While the distinct differences of bacterial composition mainly due to their different nutritional and energy requirements.

The structure of Apteryx’s eggshell has generated much debate over the decades because it does not fit well with most allometric predictions. Apteryx eggshells are unusually thin and have been reported to be 60% less porous than expected. It has been suggested that these adaptations are compensations for a very long incubation period. Most studies so far have been carried out in what has been reported as Apteryx australis, and using infertile eggs or eggs laid in captivity. However, A. australis once comprised all kiwi with brown plumage, now separated into three distinct species: Brown Kiwi (A.mantelli), Rowi (A.rowi), and Tokoeka (A.australis). These three species use different habitats and live at different latitudes and altitudes. In addition, captive eggs are much smaller than wild laid eggs. These confounding factors make necessary to revise the assumptions made for Apteryx in the past. In this study, we analysed the physical characteristics of the Apteryx eggshells making a comparison between the three species of brown coloured kiwi and for some of the analysis we included some specimens of Roroa (A. haastii, Great Spotted Kiwi). We found that shell characteristics are different between the different species studied. The pore area of Apteryx eggshells was higher than previously suggested, and the water vapour conductance was much closer to what is expected for an egg that size. We found several new features such as triangular mineral particles composing the cuticle, only reported for a cretaceous Theropod, and the presence of plugs and caps on the eggshell pores. We suggest that the characteristics of the eggshells of the different species relate to the mating system of each species in addition to environmental variables, particularly pluviosity. We also suggest that the erosion of the cuticle during incubation is an adaptation to a long incubation period in a burrow.