Long-term, climate-driven phenological shift in a tropical large carnivore

Compared to their main competitors, African wild dogs (Lycaon pictus) have inferior competitive abilities and interspecific competition is a serious fitness-limiting factor. Lions (Panthera leo) are the dominant large carnivore in African savannah ecosystems and wild dogs avoid them both spatially and temporally. Wild dog young are particularly vulnerable and suffer high rates of mortality from lions. Since lions do not utilize all parts of the landscape with an equal intensity, spatial variation in lion densities can be exploited by wild dogs both during their general ranging behaviour, but more specifically when they are confined to a den with vulnerable young. Since patches of rugged terrain are associated with lower lion densities, we hypothesized that these comparatively safe habitats should be selected by wild dogs for denning. We investigated the relationship between the distribution of 100 wild dog den sites and the occurrence of rugged terrain in four wild dog populations located in Tanzania, Zimbabwe and South Africa. A terrain ruggedness index was derived from a 90 m digital elevation model and used to map terrain ruggedness at each site. We compared characteristics of actual and potential (random) den sites to determine how wild dogs select den sites. The distributions of wild dog dens were strongly associated with rugged terrain and wild dogs actively selected terrain that was more rugged than that available on average. The likelihood of encountering lions is reduced in these habitats, minimizing the risk to both adults and pups. Our findings have important implications for the conservation management of the species, especially when assessing habitat suitability for potential reintroductions. The simple technique used to assess terrain ruggedness may be useful to investigate habitat suitability, and even predict highly suitable denning areas, across large landscapes.

African wild dogs ( Lycaon pictus ) are endangered largely because their population‐density is low under all conditions. Interspecific competition with larger carnivores may be a factor limiting wild dog density. The density of wild dogs on a 2600‐km 2 area of the Selous Game Reserve (Tanzania) was 0.04 adults/km 2 . Spotted hyaena ( Crocuta crocuta ) density for the same area was estimated by audio playbacks as 0.32 hyaenas/km 2 . Lion ( Panthera leo ) density, determined from the ratio of hyaenas to lions, was 0.11 lions/km 2 . Across six ecosystems including Selous, there were strong negative correlations between wild dog and hyaena densities (r = −0.92; p = 0.01) and between wild dog and lion densities (r = −0.91; p = 0.03). Hyaenas out‐numbered wild dogs by ratios ranging from 8:1 to 122:1. Ratios of lions to wild dogs ranged from 3:1 to 21:1. The diets of hyaenas and wild dogs overlap extensively; those of wild dogs and lions show less overlap. Where hyaenas are common and visibility is good, interference competition from hyaenas at wild dog kills is common and reduces wild dogs’ feeding time. Where hyaena density is lower and visibility is poor, interference competition at wild dog kills is rare. Wild dogs are commonly killed by lions and occasionally by hyaenas. These data suggest that competition with spotted hyaenas may limit or exclude wild dogs when hyaena density is high. Competition with lions appears less intense, but direct predation by lions on wild dogs is important. Competition and predation by larger carnivores may be of broad importance to the conservation of wild dogs and other medium‐sized carnivores.

Interaction among species through competition is a principle process structuring ecological communities, affecting behavior, distribution, and ultimately the population dynamics of species. High competition among large African carnivores, associated with extensive diet overlap, manifests in interactions between subordinate African wild dogs (Lycaon pictus) and dominant lions (Panthera leo) and spotted hyenas (Crocuta crocuta). Using locations of large carnivores in Hluhluwe-iMfolozi Park, South Africa, we found different responses from wild dogs to their two main competitors. Wild dogs avoided lions, particularly during denning, through a combination of spatial and temporal avoidance. However, wild dogs did not exhibit spatial or temporal avoidance of spotted hyenas, likely because wild dog pack sizes were large enough to adequately defend their kills. Understanding that larger carnivores affect the movements and space use of other carnivores is important for managing current small and fragmented carnivore populations, especially as reintroductions and translocations are essential tools used for the survival of endangered species, as with African wild dogs.

Hot or hungry? A tipping point in the effect of prey depletion on African wild dogs

Most African wild dog (Lycaon pictus) populations are in decline and, due to habitat fragmentation and conflict rates in areas of higher anthropogenic land-use, are primarily restricted to protected areas. As a species that occurs at low densities, with a strict reproductive social structure, wild dogs rely on long-range dispersal to facilitate colonization, reproduction, and pack formation. In Malawi, large carnivores have been subject to widespread population decline and several protected areas have a reduced large carnivore guild, including the loss of resident wild dog populations. Here, during a biodiversity monitoring camera trap survey, we captured a novel record of wild dogs in Vwaza Marsh Wildlife Reserve (Vwaza), Malawi. The 11 photographic captures of potentially three individual wild dogs represent the first documented evidence of the species in Vwaza and the first record since an unconfirmed report in 2011. We hypothesize that this group of wild dogs moved into Vwaza through the Malawi-Zambia Transfrontier Conservation Area (MZTFCA), with the MZTFCA linking protected areas in Malawi with the Luangwa Valley, Zambia. The evidence provided here, and similar documentation in Kasungu National Park, Malawi, show that large carnivores can potentially move through the MZTFCA into protected areas in Malawi. We argue that the MZTFCA provides an important dispersal corridor that could help facilitate the recolonization of wild dogs, and other large carnivores, in Malawian protected areas. However, further research is needed to assess the permeability and status of the MZTFCA corridor into Malawi. We show that camera traps can be useful to document novel records of rare species and can be used to inform conservation management planning.

Predators of similar size often compete over prey. In semi-arid ecosystems where water is a limiting resource, prey availability can be affected by water distribution, which further increases resource competition and exacerbate conflict among predators. This can have implications for carnivore dietary competition. Hence, we evaluated the dynamics of food resource competition between African wild dogs and four competing predators (cheetahs, leopards, lions and spotted hyaenas) in different seasons and across areas with different waterhole densities in Hwange National Park, Zimbabwe. We used the frequency of occurrence of prey items found in predators' scats to analyse diet composition, overlap and prey preference. For most predators, kudu was most frequently consumed and preferred. Low and medium water-dependent prey (medium and small-sized) were mostly consumed by wild dogs, leopards and cheetahs. Wild dog diet overlap was high with all predators, particularly with hyaenas and lions. There were no seasonal differences in the predators diet. The diet overlap of wild dogs with lions was highest in the low waterhole density area, and wild dog diet composition did not differ significantly from the diet of lions and hyaenas. In the low waterhole density area, wild dogs and hyaenas broadened their niche breadth, and predators diet had a higher proportion of low water-dependent prey. A low density of waterholes increased food resource competition. However, high density of waterholes, where there is more prey availability, can increase the aggregation and density of predators, and hence, increase the risks involved in interspecific competition on wild dogs. To reduce food resource competition on wild dogs, we propose to conserve larger-bodied prey that are less dependent on water (e.g. kudu, reedbuck, eland and gemsbok). As the use of water pumping is common practice, we propose maintaining water management heterogeneity where prey which is less dependent on water can also thrive.

<h3>Abstract</h3> Climate change has induced pronounced shifts in the reproductive phenology of plants, with the timing of first flowering advancing in most species. Indeed, population persistence may be threatened by the inability to track climate change phenologically. Nevertheless, substantial variation exists in biological responses to climate change across taxa. Here, we explore the consequences of climate change for flowering phenology by integrating data from a long-term observational study and a manipulative experiment under contemporary conditions. Dissecting the environmental factors that influence phenological change will illuminate why interspecific variation exists in responses to climate change. We examine a 43-year record of first flowering for six species in subalpine meadows of Colorado in conjunction with a 3-year snow manipulation experiment on the perennial mustard <i>Boechera stricta</i> from the same site. We analyze shifts in the onset of flowering in relation to environmental drivers known to influence phenology: the timing of snowmelt, the accumulation of growing degree days, and photoperiod. At our study site, climate change is reducing snowpack and advancing the timing of spring snowmelt. We found that variation in phenological responses to climate change depended on the sequence in which species flowered, with early-flowering species flowering faster, at a lower heat sum, and under increasingly disparate photoperiods in comparison to species that flower later in the season. Furthermore, climate change is outpacing phenological change for all species. Early snow removal treatments confirm that the timing of snowmelt governs observed trends in flowering phenology of <i>B. stricta</i> and that climate change can reduce the probability of flowering, thereby depressing fitness. Shorter-term studies would not have captured the trends that we document in our observational and experimental datasets. Accurate predictions of the biological responses to climate change require a thorough understanding of the specific environmental factors driving shifts in phenology.

Climate change imposes an urgent need to recognise and conserve the species likely to be worst affected. However, while ecologists have mostly explored indirect effects of rising ambient temperatures on temperate and polar species, physiologists have predicted direct impacts on tropical species. The African wild dog (Lycaon pictus), a tropical species, exhibits few of the traits typically used to predict climate change vulnerability. Nevertheless, we predicted that wild dog populations might be sensitive to weather conditions, because the species shows strongly seasonal reproduction across most of its geographical range. We explored associations between weather conditions, reproductive costs, and reproductive success, drawing on long-term wild dog monitoring data from sites in Botswana (20°S, 24years), Kenya (0°N, 12years), and Zimbabwe (20°S, 6years). High ambient temperatures were associated with reduced foraging time, especially during the energetically costly pup-rearing period. Across all three sites, packs which reared pups at high ambient temperatures produced fewer recruits than did those rearing pups in cooler weather; at the non-seasonal Kenya site such packs also had longer inter-birth intervals. Over time, rising ambient temperatures at the (longest-monitored) Botswana site coincided with falling wild dog recruitment. Our findings suggest a direct impact of high ambient temperatures on African wild dog demography, indicating that this species, which is already globally endangered, may be highly vulnerable to climate change. This vulnerability would have been missed by simplistic trait-based assessments, highlighting the limitations of such assessments. Seasonal reproduction, which is less common at low latitudes than at higher latitudes, might be a useful indicator of climate change vulnerability among tropical species.

Large carnivores have experienced widespread extirpation and species are now threatened globally. The ecological impact of the loss of large carnivores has been prominent in Gorongosa National Park, Mozambique, after most were extirpated during the 1977–92 civil war. To remedy this, reintroductions are now being implemented in Gorongosa, initiating with endangered African wild dogs (Lycaon pictus), hereafter ‘wild dogs’. We describe the first transboundary translocation and reintroduction of founding packs of wild dogs to Gorongosa over a 28-month study period and evaluate the success of the reintroduction based on five key indicator categories. We also assess how wild dog space use and diet influenced their success. We found that pre-release, artificial pack formation in holding enclosures aided group cohesion and alpha pair establishment. Post-release, we also observed natural pack formations as a result of multiple dispersal events. Founder and naturally formed packs produced pups in two of the three breeding seasons and packs successfully recruited pups. Survival rate for all wild dogs was 73% and all mortality events were from natural causes. Consequently, the population grew significantly over the study period. All indicators of success were fully achieved and this study documents the first successful reintroduction of wild dogs into a large, unfenced landscape in Mozambique and only the second on the continent. Potential mechanisms underlying these early successes were the avoidance of habitats intensively used by lions, dietary partitioning with lion, avoidance of human settlements, and Gorongosa’s management strategy. We predict further population expansion in Gorongosa given that 68% of the park is still unused by wild dogs. This expansion could be stimulated by continued reintroductions over the short- to medium-term. Recovery of wild dogs in Gorongosa could aid in the re-establishment of a larger, connected population across the greater Gorongosa-Marromeu landscape.

Large carnivores have experienced widespread extirpation and species are now threatened globally. The ecological impact of the loss of large carnivores has been prominent in Gorongosa National Park, Mozambique, after most were extirpated during the 1977-92 civil war. To remedy this, reintroductions are now being implemented in Gorongosa, initiating with endangered African wild dogs (Lycaon pictus), hereafter 'wild dogs'. We describe the first transboundary translocation and reintroduction of founding packs of wild dogs to Gorongosa over a 28-month study period and evaluate the success of the reintroduction based on five key indicator categories. We also assess how wild dog space use and diet influenced their success. We found that pre-release, artificial pack formation in holding enclosures aided group cohesion and alpha pair establishment. Post-release, we also observed natural pack formations as a result of multiple dispersal events. Founder and naturally formed packs produced pups in two of the three breeding seasons and packs successfully recruited pups. Survival rate for all wild dogs was 73% and all mortality events were from natural causes. Consequently, the population grew significantly over the study period. All indicators of success were fully achieved and this study documents the first successful reintroduction of wild dogs into a large, unfenced landscape in Mozambique and only the second on the continent. Potential mechanisms underlying these early successes were the avoidance of habitats intensively used by lions, dietary partitioning with lion, avoidance of human settlements, and Gorongosa's management strategy. We predict further population expansion in Gorongosa given that 68% of the park is still unused by wild dogs. This expansion could be stimulated by continued reintroductions over the short- to medium-term. Recovery of wild dogs in Gorongosa could aid in the re-establishment of a larger, connected population across the greater Gorongosa-Marromeu landscape.

African wild dogs (Lycaon pictus) are the most endangered large carnivores in southern Africa. Direct and indirect persecution by farmers causes significant conservation challenges. Farmer– wild dog conflict in Botswana commonly occurs as a result of cattle and stocked game depredation by wild dogs, affecting farmer livelihood and causing economic and emotional distress. Although wild dogs predate livestock at lower levels than other carnivores, they continue to be killed both indiscriminately and in retaliation for incidents of depredation. Investigating farmer–wild dog conflict is a necessary step towards establishing appropriate conflict mitigation strategies. Eighty livestock and game farmers were interviewed in order to examine farmers’ value of, perceptions of and experiences with wild dogs as well as their insights on wild dog impacts and conservation in the eastern Kalahari region of Botswana. Interviews were semi-structured and used open-ended questions to capture complexities surrounding farmer–wild dog relations. This research contributes baseline data on wild dogs in understudied tribal land and commercial livestock and game farms in eastern Kalahari. It confirms the presence of wild dogs, livestock and stocked game depredation by wild dogs and negative perspectives amongst farmers towards wild dogs and their conservation. Mean losses were 0.85 livestock per subsistence farmer, 1.25 livestock per commercial livestock farmer, while game farmers lost 95.88 game animals per farmer during January 2012 through June 2013. Proportionally, more subsistence farmers than commercial livestock farmers and game farmers held negative perspectives of wild dogs (χ ² = 9.63, df = 2, p &lt; 0.05). Farmer type, education level, socioeconomic status and land tenure, as well as positive wild dog characteristics should be considered when planning and operationalising conflict mitigation strategies. As such, conservation approaches should focus on conservation education schemes, improved wild prey base for wild dogs, poverty alleviation, and community engagement in order to offer long-term opportunities for addressing farmer–wild dog conflict in Botswana.Conservation implications: Our research contributes to wild dog conservation in Botswana by confirming the presence of wild dogs and the occurrence of livestock and stocked game depredation in previously understudied tribal land and commercial livestock and game farms in eastern Kalahari. To improve predominately negative perceptions of wild dogs and reduce conflict, practitioners should focus their efforts on conservation education schemes, improved wild prey base for wild dogs, poverty alleviation, and community engagement.

African wild dogs (Lycaon pictus) always live at low population densities relative to sympatric large carnivores. This suggests that there are basic ecological reasons for the wild dog's endangered status. We examined the effects of intraspecific and interspecific competition, infectious diseases, foraging success, genetics and human activities on wild dogs. We present data from wild dogs in the Selous Game Reserve, and make comparisons with other populations to identify the limiting factors that are broadly important. The high density of wild dogs in Selous (40 adults/1000 km2) is associated with weak competition from lions and spotted hyenas. Across ecosystems, population density is negatively related to the intensity of interference competition with larger carnivores. Predation by lions and hyenas accounted for 13% of known-cause deaths in Selous and 33–50% in other populations. Intraspecific competition caused 69% of known-cause deaths in Selous, through infanticide and fights between packs, although most of the victims were juveniles with low reproductive value. Infectious diseases had little apparent impact in Selous (4% of deaths) or Kruger National Park (5%), but did play a role in the extinction of a small population in Serengeti. Infectious diseases and competition will generally interact because competitors harbor and transmit the diseases that affect wild dogs. Human activities caused 12% of deaths in Selous, even though it is large (43 000 km2) and does not border large human or livestock populations. Humans were the major agent of mortality in some populations. Foraging success varied little across ecosystems and was not apparently limiting. Mitochondrial DNA (mtDNA) genotypes revealed clinal variation between Selous and distant populations (rather than geographically isolated subspecies, as previously suggested). All wild dog populations have a genetically effective size (Ne) less than 500, so gene flow is necessary to maintain genetic diversity within populations.

African wild dogs (Lycaon pictus) always live at low population densities relative to sympatric large carnivores. This suggests that there are basic ecological reasons for the wild dog's endangered status. We examined the effects of intraspecific and interspecific competition, infectious diseases, foraging success, genetics and human activities on wild dogs. We present data from wild dogs in the Selous Game Reserve, and make comparisons with other populations to identify the limiting factors that are broadly important. The high density of wild dogs in Selous (40 adults/1000 km2) is associated with weak competition from lions and spotted hyenas. Across ecosystems, population density is negatively related to the intensity of interference competition with larger carnivores. Predation by lions and hyenas accounted for 13% of known‐cause deaths in Selous and 33–50% in other populations. Intraspecific competition caused 69% of known‐cause deaths in Selous, through infanticide and fights between packs, although most of the victims were juveniles with low reproductive value. Infectious diseases had little apparent impact in Selous (4% of deaths) or Kruger National Park (5%), but did play a role in the extinction of a small population in Serengeti. Infectious diseases and competition will generally interact because competitors harbor and transmit the diseases that affect wild dogs. Human activities caused 12% of deaths in Selous, even though it is large (43 000 km2) and does not border large human or livestock populations. Humans were the major agent of mortality in some populations. Foraging success varied little across ecosystems and was not apparently limiting. Mitochondrial DNA (mtDNA) genotypes revealed clinal variation between Selous and distant populations (rather than geographically isolated subspecies, as previously suggested). All wild dog populations have a genetically effective size (Ne) less than 500, so gene flow is necessary to maintain genetic diversity within populations.

Climate change has induced pronounced shifts in the reproductive phenology of plants, yet we know little about which environmental factors contribute to interspecific variation in responses and their effects on fitness. We integrate data from a 43yr record of first flowering for six species in subalpine Colorado meadows with a 3yr snow manipulation experiment on the perennial forb Boechera stricta (Brassicaceae) from the same site. We analyze shifts in the onset of flowering in relation to environmental drivers known to influence phenology: the timing of snowmelt, the accumulation of growing degree days, and photoperiod. Variation in responses to climate change depended on the sequence in which species flowered, with early-flowering species reproducing faster, at a lower heat sum, and under increasingly disparate photoperiods relative to later-flowering species. Early snow-removal treatments confirm that the timing of snowmelt governs observed trends in flowering phenology of B.stricta and that climate change can reduce the probability of flowering, thereby depressing fitness. Our findings suggest that climate change is decoupling historical combinations of photoperiod and temperature and outpacing phenological changes for our focal species. Accurate predictions of biological responses to climate change require a thorough understanding of the factors driving shifts in phenology.

Plants respond to climate change through shifts in traits such as height, leaf width, and flowering time. However, little is known about how grass species in semiarid ecosystems are responding. In this study, we tested three hypotheses: (1) grass species are experiencing shifts in their vegetative and reproductive organs through time, (2) precipitation is the primary driver of these morphological shifts, and (3) the reproductive period of annual grasses changes through years in the Brazilian semiarid region. We analyzed morphological and phenological data from 590 herbarium specimens of four annual grass species collected between 1859 and 2022, along with climate data from 1960 onwards. Using simple and multiple linear regressions, we assessed relationships between morphological, phenological, climatic, and temporal variables. We tested changes in phenological synchronicity related to two periods of land use alterations. Our results revealed morphological changes throughout 1859-2022: three species showed reductions in plant height, two species exhibited shorter leaves and inflorescences, and one species presented smaller spikelets. Phenological times were delayed with increasing temperatures, although no consistent directional change in reproductive phenology was observed over the last 163 years. We also found a reduction in phenological synchronicity correlated with increasing land use shift. These findings contribute to understanding morphological and phenological shifts of grasses from semiarid ecosystems in parallel to climate and land use changes.