Both termites and large mammalian herbivores (LMH) are savanna ecosystem engineers that have profound impacts on ecosystem structure and function. Both of these savanna engineers modulate many common and shared dietary resources such as woody and herbaceous plant biomass, yet few studies have addressed how they impact one another. In particular, it is unclear how herbivores may influence the abundance of long-lived termite mounds via changes in termite dietary resources such as woody and herbaceous biomass. While it has long been assumed that abundance and areal cover of termite mounds in the landscape remains relatively stable, most data are observational, and few experiments have tested how termite mound patterns may respond to biotic factors such as changes in large herbivore communities. Here, we use a broad tree density gradient and two landscape-scale experimental manipulations—the first a multi-guild large herbivore exclosure experiment and the second a tree removal experiment– to demonstrate that patterns in termite mound abundance and cover are unexpectedly dynamic. Termite mound abundance, but not areal cover not significantly, is positively associated with experimentally controlled presence of cattle, but not wild mesoherbivores (15-1000 kg) or megaherbivores (elephants and giraffes). Herbaceous productivity and tree density, termite dietary resources that significantly affected by different LMH treatments, are both positive predictors of termite mound abundance. Experimental reductions of tree densities are associated with lower abundances of termite mounds. These results reveal a richly interacting web of relationships among multiple savanna ecosystem engineers and suggest that termite mound abundance and areal cover is intimately tied to herbivore-driven resource availability.
To study the effect of the invasion of Bursaphelenchus xylophilus on the functional relationship between woody plants and insect communities, the populations of tree species and insect communities were investigative in the Masson pine forests with different infestation durations of B. xylophilus. In this study, the number of Pinus massoniana began to decrease sharply, whereas the total number of other tree species in the arboreal layer increased gradually with the infestation duration of B. xylophilus. The principal component analysis ordination biplot shows that there was a significant change in the spatial distribution of woody plant species in different Masson pine forest stands. Additionally, a total of 7,188 insect specimens was obtained. The insect population showed an upward trend in stand types with the increase of pine wilt disease infection periods, which demonstrated that the insect community had been significantly affected by the invasion of B. xylophilus. The structure of insect functional groups changed from herbivorous (He) > omnivorous (Om) > predatory (Pr) > parasitic (Pa) > detritivorous (De) in the control stand to He > Pa > Om, De > Pr after B. xylophilus infestation in the forests. The results showed that the populations of He, Pa, and De increased after the invasion of B. xylophilus, but the populations of Pr decreased. Moreover, the redundancy analysis ordination bi-plots reflected the complicated functional relationship between woody plant communities and insects after the invasion of B. xylophilus. The present study provides insights into the changes in the community structure of woody plants and insects, as well as the functional relationship between woody plant communities and insect communities after invasion of B. xylophilus.
Dens are places for cavernicolous animals to hibernate, reproduce, and avoid predators and harsh weather conditions, and thus they have a vital impact on their survival. M. himalayana is the main large cavernicolous rodent on the Qinghai-Tibet Plateau. The analysis of den traits and their ecological functions can reveal mechanisms by which marmots have adapted to their environment, which is important for further understanding the ecological significance of this species. From July to August 2019 (warm season), we used unmanned aerial vehicles to fly at low altitudes and slow speeds to locate 131 marmot burrows (45 on shaded slopes, 51 on sunny slopes, and 35 on flat areas) in the northeastern Qinghai-Tibet Plateau. We then measured the physical characteristics (den density, entrance size, first tunnel length, volume, orientation and plant characteristics near the den entrance) of these dens on site. The physical parameters of the M. himalayana dens showed that they function to protect the marmots from natural enemies and bad weather, provide good drainage, and maintain a stable microclimate around the entrance. This is a result of the marmot’s adaptation to the harsh environment (cold and humidity) of the Qinghai-Tibet Plateau.
1. The description and analysis of animal behaviour over long periods of time is one of the most important challenges in ecology. However, most of these studies are limited due to the time and cost required by human observers. The collection of data via video recordings allows observation periods to be extended. However, their evaluation by human observers is very time-consuming. Progress in automated evaluation, using suitable deep learning methods, seems to be a forwardlooking approach to analyse even large amounts of video data in an adequate time frame. 2. In this study we present amulti-step convolutional neural network system for detecting animal behaviour states, which works with high accuracy. An important aspect of our approach is the introduction of model averaging and post-processing rules to make the system robust to outliers. 3. Our trained system achieves an in-domain classification accuracy of >0.92, which is improved to >0.96 by a postprocessing step. In addition, the whole system performs even well in an out-of-domain classification task with two unknown types, achieving an average accuracy of 0.93. We provide our system at https://github.com/Klimroth/Video-Action-Classifier-for-African-Ungulates-in-Zoos/tree/main/mrcnn_based so that interested users can train their own models to classify images and conduct behavioural studies of wildlife. 4. The use of a multi-step convolutional neural network for fast and accurate classification of wildlife behaviour facilitates the evaluation of large amounts of image data in ecological studies and reduces the effort of manual analysis of images to a high degree. Our system also shows that post-processing rules are a suitable way to make species-specific adjustments and substantially increase the accuracy of the description of single behavioural phases (number, duration). The results in the out-of-domain classification strongly suggest that our system is robust and achieves a high degree of accuracy even for new species, so that other settings (e.g. field studies) can be considered.
The mass die-off of Caribbean corals has transformed many of this region’s reefs to macroalgal-dominated habitats since systematic monitoring began in the 1970s. Although attributed to a combination of local and global human stressors, the lack of long-term data on Caribbean reef coral communities has prevented a clear understanding of the causes and consequences of coral declines. We integrated paleoecological, historical, and modern survey data to track the prevalence of major coral species and life history groups throughout the Caribbean from the pre-human period to present. The regional loss of Acropora corals beginning by the 1960s from local human disturbances resulted in increases in the prevalence of formerly subdominant stress-tolerant and weedy scleractinian corals and the competitive hydrozoan Millepora beginning in the 1970s and 1980s. These transformations have resulted in the homogenization of coral communities within individual countries. However, increases in stress-tolerant and weedy corals have slowed or reversed since the 1980s and 1990s in tandem with intensified coral bleaching. These patterns reveal the long history of increasingly stressful environmental conditions on Caribbean reefs that began with widespread local human disturbances and have recently culminated in the combined effects of local and global change.
Males have the ability to compete for fertilizations through both pre-copulatory and post-copulatory intrasexual competition. Pre-copulatory competition has selected for large weapons and other adaptations to maximize access to females and mating opportunities while post-copulatory competition has resulted in ejaculate adaptations to maximize fertilization success. Negative associations between these strategies support the hypothesis that there is a trade-off between success at pre- and post-copulatory mating success. Recently, this trade-off has been demonstrated with experimental manipulation. Male leaf-footed cactus bugs, Narnia femorata, that lose a weapon by autotomy during development invest instead in large testes. While evolutionary outcomes of the trade-offs between pre- and post-copulatory strategies have been identified, less work has been done to identify proximate mechanisms by which the trade-off might occur, perhaps because the systems in which the trade-offs have been investigated are not ones that have the molecular tools required for exploring mechanism. Here we applied knowledge from a related model species for which we have developmental knowledge and molecular tools, the milkweed bug Oncopeltus fasciatus, to investigate the proximate mechanism by which autotomized N. femorata males developed larger testes. Autotomized males had evidence of a higher rate of transit amplification divisions in the spermatogonia, which would result in greater sperm numbers. Identification of mechanisms underlying a trade-off can help our understanding of the direction and constraints on evolutionary trajectories and thus the evolutionary potential under multiple forms of selection.
Artiodactyl prey species of Chile, especially guanacos (Lama guanicoe) are reported to be very susceptible to predation by pack hunting feral dogs. It has been previously suggested that guanacos and endemic South American deer may have evolved in the absence of pack-hunting cursorial predators. However, the paleoecology of canid presence in southern South America and Chile is unclear. Here, we review the literature on South American and Chilean canids, their distributions, ecologies and hunting behaviour. We consider both wild and domestic canids, including Canis familiaris breeds. We establish two known antipredator defense behaviours of guanacos: predator inspection of ambush predators, e.g. Puma concolor, and rushing at and kicking smaller cursorial predators, e.g. Lycalopex culpaeus. We propose that since the late Pleistocene extinction of hypercarnivorous group-hunting canids east of the Andes, there were no native species creating group-hunting predation pressures on guanacos. Endemic deer of Chile may have never experienced group hunting selection pressure from native predators. Even hunting dogs (or other canids) used by indigenous groups in the far north and extreme south of Chile (and presumably the center as well) appear to have been used primarily within ambush hunting strategies. This may account for the susceptibility of guanacos and other prey species to feral dog attacks. We detail seven separate hypotheses that require further investigation in order to assess how best to respond to the threat posed by feral dogs to the conservation of native deer and camelids in Chile and other parts of South America.
Diffusible iodine-based contrast-enhanced Computed-Tomography (diceCT) visualizes soft-tissue from microCT (µCT) scans of specimens to uncover internal features and natural history information without incurring physical damage via dissection. Unlike hard-tissue imaging, diceCT datasets are currently limited to a few individual specimens and taxonomically underrepresented. To initiate best practices for diceCT in a non-model group, we outline a guide for staining and high-throughput µCT scanning in snakes. We scanned the entire body and one region of interest (i.e., head) for 23 specimens representing 23 species from the clades Aniliidae, Dipsadinae, Colubrinae, Elapidae, Lamprophiidae and Viperidae. We generated 82 scans that include 1.25% Lugols iodine stained (soft tissue) and unstained (skeletal) data for each specimen. We found that duration of optimal staining time increased linearly with body size; head radius was the best indicator. Post-reconstruction of scans, optimal staining was evident by evenly distributed grayscale values and clear differentiation among soft-tissue anatomy. Under and over stained specimens produced poor contrast among soft-tissues, which was often exacerbated by user bias during “digital dissections” (i.e., segmentation). Regardless, all scans produced usable data from which we assessed a range of downstream analytical applications within ecology and evolution (e.g., predator-prey interactions, life history, and morphological evolution). Ethanol de-staining reversed the known effects of iodine on the exterior appearance of physical specimens, but required substantially more time than reported for other de-staining methods. We discuss the feasibility of implementing diceCT techniques for a new user, including approximate financial and temporal commitments, required facilities, and potential effects of staining on specimens. We present the first high-throughput workflow for full-body skeletal and diceCT scanning in snakes, which can be generalized to any elongate vertebrates, and increases publicly available diceCT scans for reptiles by an order of magnitude.
Aim The aim of this study is to model the past, current and future distribution of J. phoenicea s.s., J. turbinata and J. canariensis, based on bioclimatic variables using a maximum entropy model (MaxEnt) in the Mediterranean and Macaronesian regions. Location Mediterranean and Macaronesian Taxon Cupressaceae, Juniperus Methods Data on the occurrence of the J. phoenicea complex was obtained from the GBIF, the literature, herbaria, and the authors’ field notes. The bioclimatic variables were obtained from the WorldClim database (http://worldclim.org/) and Paleoclim (http://www.paleoclim.org/). The climate data related to species localities were used for predictions of niches by implementation of MaxEnt and we evaluated the model with ENMeval. Results The potential niches of Juniperus phoenicea during the LIG, LGM and MH covered 30%, 10% and almost 100%, respectively, of the current potential niche. Climate warming could reduce potential niches by 30% and 90% in scenarios RCP2.6 and RCP8.5, respectively. The potential niches of Juniperus turbinata had a broad circum-Mediterranean and Canarian distribution during the LIG and the MH, extending its distribution during the LGM when it was found in more areas than at present; the predicted warming in scenario RCP2.6 and RCP8.5 could reduce the current potential niche by 30% and 50%, respectively. The model did not find suitable niches for J. canariensis during the LIG and the LGM, but during the MH its potential niche was 30% larger than at present. The climate warming scenario RCP2.6 indicates a reduction of the potential niche by 30%, while RCP8.5 does so by almost 60%. Main conclusions This research can provide information to increase the protection of the juniper forest and to try to counteract the phenomenon of local extinctions caused by anthropic pressure and climate changes.
Seed dispersal by ants is an important means of migration for plants. Although many 34 myrmecochorous plants have seeds containing elaiosome, a nutritional reward for ants, some 35 non-myrmecochorous seeds without elaiosomes are also dispersed by ant species. However, the 36 mechanism by which seeds without elaiosomes enable efficient dispersal by ants is scarcely 37 investigated. The seeds of the achlorophyllous and myco-heterotrophic herbaceous plant 38 Monotropastrum humile are very small without elaiosomes and require a fungal host for 39 germination and survival. We performed a bioassay using seeds of M. humile and the ant 40 Nylanderia flavipes to demonstrate ant-mediated seed dispersal. We also analyzed the volatile 41 odors emitted from M. humile seeds and conducted bioassays using dummy seeds coated with 42 seed volatiles. Although elaiosomes were absent from the M. humile seeds, the ants carried the 43 seeds to their nests. They also carried the dummy seeds coated with the seed volatile mixture to 44 the nest, and left some dummy seeds inside the nest and discarded the rest of the dummy seeds 45 outside the nest with a bias toward locations with moisture conditions, which might be 46 conducive to germination. We concluded that seeds of M. humile were dispersed by the ants, 47 and that seed odors were sufficient to induce directed dispersal even without elaiosomes. It is 48 probable that the fleshy fruit producing genus Monotropastrum evolved from the related 49 anemochorous genus Monotropa, which produces capsule fruit. This transformation from 50 anemochory to myrmecochory presents a novel evolutionary pathway toward ant-mediated seed 51 dispersal in an achlorophyllous plant.
The muskrat (Ondatra zibethicus) is an iconic species in Canada, valued for both its fur and its integral role in wetland ecosystems, and widely regarded for its perseverance. However, the resilience of this semi-aquatic mammal seems to be in question now as increasing evidence points to widespread population declines. Recent analyses of harvest data across North America suggest a reduction in their numbers, but this has not been widely corroborated by population surveys. In this study we replicated historic muskrat house count surveys at two large Great Lakes coastal wetlands and present confirmation that declines in muskrat harvest correspond to actual declines in muskrat abundance. At the Point Pelee National Park marsh and the Matchedash Bay-Gray Marsh wetland we found that mean muskrat house counts declined by 93% and 91% respectively between historic surveys 40-50 years ago and contemporary surveys over the past five years. The factors responsible for these dramatic declines remain unclear but there may be a relationship with changes in the habitat quality of these wetlands that have occurred over the same time frame. Not only is the loss of muskrats an issue for the resulting loss of the wetland ecosystem services they provide, but it may be an indication of broader marsh ecosystem degradation. As such, a scarcity of muskrats should be considered a red flag for the state of biodiversity in our wetlands. Continued surveys and ongoing research are needed to shed more light on the current status of muskrat populations and their marsh habitats across their native range. Keywords: Fur harvest; Muskrat; Ondatra; Population decline; Typha; Wetlands
Populations adapt to novel environmental conditions by genetic changes or phenotypic plasticity. Plastic responses are generally faster and can buffer fitness losses under variable conditions. Plasticity is typically modelled as random noise and linear reaction norms that assume simple one-to-one genotype-phenotype maps and no limits to the phenotypic response. Most studies on plasticity have focused on its effect on population viability. However, it is not clear, whether the advantage of plasticity depends solely on environmental fluctuations or also on the genetic and demographic properties (life histories) of populations. Here we present an individual-based model and study the relative importance of adaptive and non-adaptive plasticity for populations of sexual species with different life histories experiencing directional stochastic climate change. Environmental fluctuations were simulated using differentially autocorrelated climatic stochasticity or noise color, and scenarios of directional climate change. Non-adaptive plasticity was simulated as a random environmental effect on trait development, while adaptive plasticity as a linear, logistic, or sinusoidal reaction norm. The last two imposed limits to the plastic response and emphasized flexible interactions of the genotype with the environment. Interestingly, this assumption led to (i) smaller phenotypic than genotypic variance in the population and the coexistence of polymorphisms, (ii) many-to-one genotype-phenotype map, and (iii) the maintenance of higher genetic variation – compared to linear reaction norms and genetic determinism – even when the population was exposed to a constant environment for several generations. Limits to plasticity led to genetic accommodation, when costs were negligible, and to the appearance of cryptic variation when limits were exceeded. We found that adaptive plasticity promoted population persistence under red noise stochasticity and was particularly important for life histories with low fecundity. Populations producing more offspring could cope with environmental fluctuations solely by genetic changes or random plasticity, unless environmental change was too fast.
Alternative reproductive tactics (ARTs) have provided valuable insights into how sexual selection and life history tradeoffs can lead to variation within a sex. However, the possibility that tactics may constrain evolution through intralocus tactical conflict (IATC) is rarely considered. In addition, when IATC has been considered, the focus has often been on the genetic correlations between the ARTs, while evidence that the ARTs have different optima for associated traits and that at least one of the tactics is not at its optima is often missing. Here we investigate selection on three traits associated with the ARTs in the swordtail fish Xiphophorus multilineatus; body size, body shape and the sexually selected trait for which these fishes were named, sword length (elongation of the caudal fin). All three traits are tactically dimorphic, with courter males being larger, deeper bodied and having longer swords, and the sneaker males being smaller, more fusiform and having shorter swords. Using measures of reproductive success in a wild population we calculated selection differentials, linear and quadratic gradients, demonstrate that the tactics have different optima and at least one of the tactics is not at its optima for body size and sword length. Our results provide the first evidence of selection in the wild on the sword, an iconic trait for sexual selection. In addition, given the high probability that these traits are genetically correlated to some extent between the two tactics, our study suggests that IATC is constraining both body size and the sword from reaching their phenotypic optima. We discuss the importance of considering the role of IATC in the evolution of tactical dimorphism, how this conflict can be present despite tactical dimorphism, and how it is important to consider this conflict when explaining not only variation within a species but differences across species as well.
Many insects possess the plastic ability to either develop directly to adulthood, or enter diapause and postpone reproduction until the next year, depending on environmental cues (primarily photoperiod) that signal the amount of time remaining until the end of the growth season. These two developmental pathways often differ in co-adapted life history traits, e.g. with slower development and larger size in individuals headed for diapause. The developmental timing of these differences may be of adaptive importance: if pathways diverge late, the scope for phenotypic differences is smaller, whereas if pathways diverge early, the risk is higher of expressing a maladaptive phenotype if the selective environment changes. Here we explore the effects of changes in photoperiodic information during life on pupal diapause and associated life history traits in the butterfly Pararge aegeria. We find that both pupal diapause and larval development rate are asymmetrically regulated: while exposure to long days late in life (regardless of earlier experiences) was sufficient to produce nondiapause development and accelerate larval development accordingly, more prolonged exposure to short days was required to induce diapause and slow down pre-diapause larval development. While the two developmental pathways diverged early in development, development rates could be partially reversed by altered environmental cues. Meanwhile, pathway differences in body size were more inflexible, despite emerging late in development. Hence, in P. aegeria several traits are regulated by photoperiod, along subtly different ontogenies, into an integrated phenotype that strikes a balance between flexibility and phenotype-environment matching.
Hybridization is a common and important stage in species formation in plants and animals. The evolutionary consequences of hybridization depend not only on reproductive compatibility between sympatric species, but also on factors like vulnerability to each other’s predators and parasites. We examine infection patterns of the blood parasite Haemoproteus lophortyx, a causative agent of avian malaria, at a site in the contact zone between California quail (Callipepla californica) and Gambel’s quail (C. gambelii). We tested whether species identity, sex, and year predicted infection status and intensity. While we found no effect of sex on the status or intensity of infection, we found differences in infection status and intensity across species and between years. The prevalence of infection in California and hybrid quail was lower than in Gambel’s quail. Once infected, however, California and hybrid quail had higher infection intensities than Gambel’s quail. California and hybrid quail exhibited no significant differences in prevalence or intensity of infection. These findings suggest that infection by H. lophortyx has the potential to influence species barrier dynamics in this system, however, more work is necessary to determine the exact evolutionary consequences of this blood parasite.
Kauri dieback, caused by Phytophthora agathidicida, is an ecosystem disturbance that poses a recent threat to the survival of kauri (Agathis australis) forests in New Zealand. Throughfall and stemflow play an important role in meeting the nutrient requirements of kauri forests. However, the effects of kauri dieback on canopy nutrient deposition remain unknown. Here we measured throughfall, stemflow and forest floor water yield and nutrient concentrations and fluxes (potassium, calcium, magnesium, manganese, silicon, sulphur, sodium, iron) of ten kauri trees differing in soil P. agathidicida DNA concentration and health status. We did not observe an effect of soil P. agathidicida DNA concentration on throughfall and stemflow water yield. Throughfall and forest floor nutrient concentrations and fluxes tended to decrease (up to 50%) with increasing soil P. agathidicida DNA concentration. Significant effects were found for potassium and manganese fluxes in throughfall, and calcium and silicon fluxes in forest floor leachate. The decline in nutrient input will have implications on plant nutrition, tree health and susceptibility to future pathogen infection in these ecologically unique kauri forests. Given our findings and the increasing spread of Phytophthora species worldwide, research on the underlying physiological mechanisms linking dieback and plant-soil nutrient fluxes is critical.
Understanding what variables affect ungulate neonate survival is imperative to successful conservation and management of the species. Predation is commonly cited as a cause-specific source of mortality and ecological covariates often influence neonate survival. However, variation in survival estimates related to capture methodology has been documented with opportunistically captured neonates generally displaying greater survival than those captured via aid of vaginal implant transmitters (VITs), likely because of increased left truncation observed in the opportunistically captured datasets. Our goal was to assess if 3- and 6-month survival estimates varied by capture method while simultaneously assessing if capture method affected model selection and interpretation of ecological covariates for white-tailed deer neonates captured from three study sites in North Dakota and South Dakota, USA. We found survival varied by capture method for 3-month neonate survival with opportunistically captured neonates displaying up to 26% greater survival than their counterparts captured via VITs; however, this relationship was not present for 6-month survival. We also found model selection and subsequent interpretation of ecological covariates varied when analyzing datasets comprised of neonates captured via VITs, neonates captured opportunistically, and all neonates combined regardless of capture method. When interpreting results from our VIT only analysis for 3-month survival, we found survival varied by three time intervals and was lowest in the first two weeks of life. Capture method did not affect 6-month survival which was most influenced by total precipitation occurring during 3 – 8 weeks of a neonate’s life and percent canopy cover found at a neonate’s capture site. Our results support previous research that capture method must be accounted for when deriving survival estimates for ungulate neonates as it can impact derived estimates and subsequent interpretation of results.
Seed recruitment is a major driver of mangrove restoration globally. It is hypothesized that soil condition and channel hydrology can accelerate seedling recruitment and regeneration after a major disturbance. Species abundance, diversity indices, microbial and chemical concentrations in sand-filled mangrove forest was studied. Eight plots (area = 3902.16 m2) were established with ten transects in each plot in a random block design to investigate the effect of soil conditions on seedling growth. A total of 1, 886 seedlings were physically counted. Seedling abundance was significantly different between red (Rizophora racemosa), white (Laguncularia racemosa) and black (Avicennia germinans) mangroves and nypa palm (nypa fruticans). The most dominant species was black mangroves and the least dominant species was nypa palm. Muddy soils had the most abundant species while sandy soils had the least abundant species. Furthermore, semi-muddy soils had the highest species diversity (H = 0.948) whereas muddy soils had the least species diversity (H = 0.022). The soil metal concentration has no correlation with seed abundance and occur in the order Iron>Nitrate>Copper>Cadmium. Soil with high species diversity had high soil microbial population; however, seedling abundance was correlated with soil nutrients and not heavy metals. Small seeds are easily recruited while good soil condition plus existing hydrological connection facilitated natural seedling regeneration in the disturbed mangrove forest.