Enhanced regional connectivity between western North American national parks will increase persistence of mammal species diversity.

Many protected areas worldwide increasingly resemble habitat isolates embedded in human-modified landscapes. However, establishing linkages among protected areas could significantly reduce species-loss rates. Here we present a novel method having broad applicability for assessing enhanced regional connectivity on persistence of mammal diversity. We combine theoretically-derived species relaxation rates for mammal communities with empirically-derived pathways. We assess the value of enhanced regional connectivity for two hypothetical networks of national parks in western North America: the Yellowstone-Glacier network and the Mount Rainier-North Cascades network. Linking the Yellowstone and Glacier park assemblages by eliminating barriers to movement in identified mammal dispersal pathways and by incorporating adjacent wilderness areas and known ungulate migratory routes into a protected area network would greatly enlarge available habitat. This would enhance medium to large mammal species persistence time by factor of 4.3, on average, or ~ 682 generations relative to individual parks. Similarly, linking Mount Rainier and North Cascades park assemblages would enhance mammal species persistence time by a factor of 4.3, on average, or ~305 generations relative to individual parks. Enhancing regional connectivity among western North America parks could serve as an important template for landscape-scale conservation in the 21st century.

Elevational distribution of montane Afrotropical butterflies is influenced by seasonality and habitat structure.

Determinants of elevational distribution of butterfly species richness and abundance in the tropics are poorly understood. Here we assess the combined effects, both additive and interactive, of seasonality and habitat structure on the elevational distribution of butterflies in the Uluguru Mountains, Tanzania. We sampled butterflies along a 1100 m elevational gradient that extended from 1540 to 2639 m using a time-constrained fixed-area method during the short to long rains and long to short rains transitions, and in habitat structure classified as closed or open. We used semi-parametric generalized linear mixed models to assess the relation between butterfly species richness or abundance, and seasonality, habitat structure, family and elevation. For all species combined, species richness declined with elevation in both open and closed habitats during the long to short rains transition. During the short to long rains transition, species richness displayed a mid-elevation peak across habitats. Among the three focal families (Nymphalidae, Papilionidae and Pieridae) similar patterns in the elevational distribution of species richness were observed. Species abundance declined or remained stable with elevation across seasons and habitat structure; the exception being species abundance in open habitat during the short to long rain transition and increased slightly with elevation. Abundance by family did not vary significantly by habitat structure or season. Our results indicate that seasonality and habitat structure shape species richness and abundance of butterflies along an elevational gradient in the Uluguru Mountains. These patterns are important for informing conservation actions because temperature as well as annual and seasonal variation in precipitation are predicted to increase in East Africa as a result of climate change, important determinants of seasonality, while habitat disturbance may increase due to a projected doubling in Tanzania's population over the next 27 years.

Afrotropical montane birds experience upslope shifts and range contractions along a fragmented elevational gradient in response to global warming.

Global warming is predicted to result in upslope shifts in the elevational ranges of bird species in montane habitats. Yet few studies have examined changes over time in the elevational distribution of species along fragmented gradients in response to global warming. Here, we report on a resurvey of an understory bird community in the Usambara Mountains in Tanzania, along a forested elevational gradient that has been fragmented over the last 200 years. In 2019, we resurveyed seven sites, ranging in elevation from 360 m to 2110 m, that were originally surveyed between 1979 and 1981. We calculated differences in mean elevation and lower and upper range limits for 29 species between the two time periods and corrected for possible differences in elevation due to chance. Over four decades, we documented a significant mean upslope shift across species of 93 m. This shift was smaller than the 125 m expected shift due to local climate warming. Of the 29 focal species, 19 shifted upslope, eight downslope, and two remained unchanged. Mean upslope shifts in species were driven largely by contracting lower range limits which moved significantly upslope on average across species by 183 m, while upper range limits shifted non-significantly upslope by 72 m, leading to a mean range contraction of 114 m across species. Community composition of understory bird species also shifted over time, with current communities resembling communities found historically at lower elevations. Past forest fragmentation in combination with the limited gap-crossing ability of many tropical understory bird species are very likely important contributory factors to the observed asymmetrical shifts in lower and upper elevational range limits. Re-establishing forested linkages among the largest and closest forest fragments in the Eastern Arc Mountains are critical to permitting species to shift upslope and to reduce further elevational range contractions over time.

Temperature-associated decreases in demographic rates of Afrotropical bird species over 30 years.

Tropical mountains harbor globally significant levels of biodiversity and endemism. Climate change threatens many tropical montane species, yet little research has assessed the effects of climate change on the demographic rates of tropical species, particularly in the Afrotropics. Here, we report on the demographic rates of 21 Afrotropical bird species over 30 years in montane forests in Tanzania. We used mark-recapture analyses to model rates of population growth, recruitment, and apparent survival as functions of annual mean temperature and annual precipitation. For over one-half of focal species, decreasing population growth rates were associated with increasing temperature. Due to the trend in temperature over time, we substituted a time covariate for the temperature covariate in top-ranked population growth rate models. Temperature was a better explanatory covariate than time for 6 of the 12 species, or 29% of all focal species. Population growth rates were also lower for species found further below their elevational midpoint and for smaller-bodied species. Changes in population growth rates were more closely tied to changes in recruitment than to changes in apparent survival. There were no consistent associations between demographic rates and precipitation. This study demonstrates temperature-associated demographic impacts for 6 (29%) of 21 focal species in an Afrotropical understory bird community and highlights the need to incorporate the impacts of climate change on demographic rates into conservation planning across the tropics.

Heterogeneous Matrix Habitat Drives Species Occurrences in Complex, Fragmented Landscapes.

A fundamental tenet of modern ecology and conservation science is the fact that species occurrence in habitat patches can be determined by patch area and isolation. But such island biogeographic models often poorly predict actual species occurrences in structurally complex landscapes that typify most ecosystems. Recent advances in circuit theory have enhanced estimates of species dispersal and can provide powerful ways to predict landscape-scale distribution of species assemblages through integration with island biogeography. Applying such an integrative analytical framework to 43 bird species in Tanzania improved model fit by an average of 2.2-fold over models where patch isolation was estimated without accounting for landscape matrix heterogeneity. This approach also allowed us to assess species-specific dispersal rates and quantify differences among land cover types in their permeability to animal movement. These results reaffirm the utility of foundational island biogeographic principles, yet with an important caveat. Two-thirds of the variance in species occurrence in habitat fragments can be explained simply by patch area and isolation, conditional on isolation explicitly accounting for the spatial configuration of different land cover types in the landscape matrix.

Targeted habitat restoration can reduce extinction rates in fragmented forests.

The Eastern Arc Mountains of Tanzania and the Atlantic Forest of Brazil are two of the most fragmented biodiversity hotspots. Species-area relationships predict that their habitat fragments will experience a substantial loss of species. Most of these extinctions will occur over an extended time, and therefore, reconnecting fragments could prevent species losses and allow locally extinct species to recolonize former habitats. An empirical relaxation half-life vs. area relationship for tropical bird communities estimates the time that it takes to lose one-half of all species that will be eventually lost. We use it to estimate the increase in species persistence by regenerating a forest connection 1 km in width among the largest and closest fragments at 11 locations. In the Eastern Arc Mountains, regenerating 8,134 ha of forest would create >316,000 ha in total of restored contiguous forest. More importantly, it would increase the persistence time for species by a factor of 6.8 per location or ∼2,272 years, on average, relative to individual fragments. In the Atlantic Forest, regenerating 6,452 ha of forest would create >251,000 ha in total of restored contiguous forest and enhance species persistence by a factor of 13.0 per location or ∼5,102 years, on average, relative to individual fragments. Rapidly regenerating forest among fragments is important, because mean time to the first determined extinction across all fragments is 7 years. We estimate the cost of forest regeneration at $21-$49 million dollars. It could provide one of the highest returns on investment for biodiversity conservation worldwide.

Dynamics of extinction debt across five taxonomic groups.

Species extinction following habitat loss is well documented. However, these extinctions do not happen immediately. The biodiversity surplus (extinction debt) declines with some delay through the process of relaxation. Estimating the time constants of relaxation, mainly the expected time to first extinction and the commonly used time for half the extinction debt to be paid off (half-life), is crucial for conservation purposes. Currently, there is no agreement on the rate of relaxation and the factors that it depends on. Here we find that half-life increases with area for all groups examined in a large meta-analysis of extinction data. A common pattern emerges if we use average number of individuals per species before habitat loss as an area index: for mammals, birds, reptiles and plants, the relationship has an exponent close to a half. We also find that the time to first determined extinction is short and increases slowly with area.

Evolution along the Great Rift Valley: phenotypic and genetic differentiation of East African white-eyes (Aves, Zosteropidae).

The moist and cool cloud forests of East Africa represent a network of isolated habitats that are separated by dry and warm lowland savannah, offering an opportunity to investigate how strikingly different selective regimes affect species diversification. Here, we used the passerine genus Zosterops (white-eyes) from this region as our model system. Species of the genus occur in contrasting distribution settings, with geographical mountain isolation driving diversification, and savannah interconnectivity preventing differentiation. We analyze (1) patterns of phenotypic and genetic differentiation in high- and lowland species (different distribution settings), (2) investigate the potential effects of natural selection and temporal and spatial isolation (evolutionary drivers), and (3) critically review the taxonomy of this species complex. We found strong phenotypic and genetic differentiation among and within the three focal species, both in the highland species complex and in the lowland taxa. Altitude was a stronger predictor of phenotypic patterns than the current taxonomic classification. We found longitudinal and latitudinal phenotypic gradients for all three species. Furthermore, wing length and body weight were significantly correlated with altitude and habitat type in the highland species Z. poliogaster. Genetic and phenotypic divergence showed contrasting inter- and intraspecific structures. We suggest that the evolution of phenotypic characters is mainly driven by natural selection due to differences in the two macro-habitats, cloud forest and savannah. In contrast, patterns of neutral genetic variation appear to be rather driven by geographical isolation of the respective mountain massifs. Populations of the Z. poliogaster complex, as well as Z. senegalensis and Z. abyssinicus, are not monophyletic based on microsatellite data and have higher levels of intraspecific differentiation compared to the currently accepted species.

Long-term demographic consequences of habitat fragmentation to a tropical understory bird community.

Tropical deforestation continues to cause population declines and local extinctions in centers of avian diversity and endemism. Although local species extinctions stem from reductions in demographic rates, little is known about how habitat fragmentation influences survival of tropical bird populations or the relative importance of survival and fecundity in ultimately shaping communities. We analyzed 22 years of mark-recapture data to assess how fragmentation influenced apparent survival, recruitment, and realized population growth rate within 22 forest understory bird species in the Usambara Mountains, Tanzania. This represents the first such effort, in either tropical or temperate systems, to characterize the effect of deforestation on avian survival across such a broad suite of species. Long-term demographic analysis of this suite of species experiencing the same fragmented environment revealed considerable variability in species' responses to fragmentation, in addition to general patterns that emerged from comparison among species. Across the understory bird community as a whole, we found significantly lower apparent survival and realized population growth rate in small fragments relative to large, demonstrating fragmentation effects to demographic rates long after habitat loss, Demographic rates were depressed across five feeding guilds, suggesting that fragmentation sensitivity was not limited to insectivores. Seniority analyses, together with a positive effect of fragmentation on recruitment, indicated that depressed apparent survival was the primary driver of population declines and observed extinctions. We also found a landscape effect, with lower vital rates in one mountain range relative to another, suggesting that fragmentation effects may add to other large-scale drivers of population decline. Overall, realized population growth rate (lambda) estimates were < 1 for most species, suggesting that future population persistence, even within large forest fragments, is uncertain in this biodiversity hotspot.

Habitat fragmentation reduces nest survival in an Afrotropical bird community in a biodiversity hotspot.

Ecologists have long hypothesized that fragmentation of tropical landscapes reduces avian nest success. However, this hypothesis has not been rigorously assessed because of the difficulty of finding large numbers of well-hidden nests in tropical forests. Here we report that in the East Usambara Mountains in Tanzania, which are part of the Eastern Arc Mountains, a global biodiversity hotspot, that daily nest survival rate and nest success for seven of eight common understory bird species that we examined over a single breeding season were significantly lower in fragmented than in continuous forest, with the odds of nest failure for these seven species ranging from 1.9 to 196.8 times higher in fragmented than continuous forest. Cup-shaped nests were particularly vulnerable in fragments. We then examined over six breeding seasons and 14 study sites in a multivariable survival analysis the influence of landscape structure and nest location on daily nest survival for 13 common species representing 1,272 nests and four nest types (plate, cup, dome, and pouch). Across species and nest types, area, distance of nest to edge, and nest height had a dominant influence on daily nest survival, with area being positively related to nest survival and distance of nest to edge and nest height being both positively and negatively associated with daily nest survival. Our results indicate that multiple environmental factors contribute to reduce nest survival within a tropical understory bird community in a fragmented landscape and that maintaining large continuous forest is important for enhancing nest survival for Afrotropical understory birds.

The need for integrative approaches to understand and conserve migratory ungulates.

Over the last two centuries overhunting, anthropogenic barriers and habitat loss have disrupted many ungulate migrations. We review the literature on ungulate migration disruptions and find that for many species the disruption of migratory routes causes a rapid population collapse. Previous research has focused on the proximal ecological factors that might favour migration, particularly spatiotemporal variation in resources and predation. However, this does not provide an adequate basis for understanding and mitigating anthropogenic effects on migratory populations. Migration is a complex behaviour and we advocate an integrative approach that incorporates population dynamics, evolution, genetics, behaviour and physiology, and that borrows insights and approaches from research on other taxa. We draw upon research on avian migration to illustrate research approaches that might also be fruitful in ungulates. In particular, we suggest that the migratory cycle should be evaluated in the context of seasonal population limitation, an approach we highlight with a preliminary demographic perturbation analysis of the Serengeti wildebeest (Connochaetes taurinus) population. We provide suggestions for avenues of future research and highlight areas where we believe rapid progress can be made by applying recent advances in theory, technology and analytical approaches.

A 16-year study of forest disturbance and understory bird community structure and composition in Tanzania.

I compared understory bird community structure and composition among primary, slightly disturbed, and moderately disturbed forest in the East Usambara Mountains in Tanzania. Comparisons were conducted at two spatiotemporal scales: short term, conducted in 1999 and 2000, in which treatments (disturbance levels) were replicated, and long term, conducted from 1989-2004 along the same disturbance gradient of a subset of the short-term sites. I used capture-recapture models to assess the probability of detection and estimate species richness and population growth rates. The probability of detection of species did not vary significantly among disturbance levels but did vary significantly among species. Over the short- and long-term surveys, estimated species richness did not vary significantly among disturbance levels. Temporal variability in estimated species richness and the relative abundance of guilds did vary significantly among disturbance levels yet was contingent on survey length. The coefficient of variation in species richness over the short-term survey was <5% across all disturbance levels, whereas over the long-term survey it was 35% in slightly disturbed forest, 11% in moderately disturbed forest, and 0% in primary forest. In the short-term survey, zero of seven feeding guilds varied significantly in relative abundance among disturbance levels, whereas over the long-term survey four of seven (57%) feeding guilds did so. Terrestrial insectivores were most adversely affected by forest disturbance. Population growth rates (lamda) between 1989 and 2004 for the more common species of terrestrial insectivores did not vary significantly among disturbance levels and for these species were significantly < or = 1.00, indicating the recovery time for terrestrial insectivores in slightly and moderately disturbed forest is very long. These results illustrate the importance of long-term studies in assessing ecological impacts of forest disturbance and the importance of protecting primary forest in the Eastern Arc Mountains.