skip to main content


Title: Vertical stratification patterns of tropical forest vertebrates: a meta‐analysis
ABSTRACT

Tropical forests harbour the highest levels of terrestrial biodiversity and represent some of the most complex ecosystems on Earth, with a significant portion of this diversity above ground. Although the vertical dimension is a central aspect of the ecology of forest communities, there is little consensus as to prominence, evenness, and consistency of community‐level stratification from ground to canopy. Here, we gather the results of 62 studies across the tropics to synthesise and assess broad patterns of vertical stratification of abundance and richness in vertebrates, the best studied taxonomic group for which results have not been collated previously. Our review of the literature yielded sufficient data for bats, small mammals, birds and amphibians. We show that variation in the stratification of abundance and richness exists within and among all taxa considered. Bat richness stratification was variable among studies, although bat abundance was weighted towards the canopy. Both bird richness and abundance stratification were variable, with no overriding pattern. On the contrary, both amphibians and small mammals showed consistent patterns of decline in abundance and richness towards the canopy. We descriptively characterise research trends in drivers of stratification cited or investigated within studies, finding local habitat structure and food distribution/foraging to be the most commonly attributed drivers. Further, we analyse the influence of macroecological variables on stratification patterns, finding latitude and elevation to be key predictors of bird stratification in particular. Prominent differences among taxa are likely due to taxon‐specific interactions with local drivers such as vertical habitat structure, food distribution, and vertical climate gradients, which may vary considerably across macroecological gradients such as elevation and biogeographic realm. Our study showcases the complexity with which animal communities organise within tropical forest ecosystems, while demonstrating the canopy as a critical niche space for tropical vertebrates, thereby highlighting the inherent vulnerability of tropical vertebrate communities to forest loss and canopy disturbance. We recognise that analyses were constrained due to variation in study designs and methods which produced a variety of abundance and richness metrics recorded across different arrangements of vertical strata. We therefore suggest the application of best practices for data reporting and highlight the significant effort required to fill research gaps in terms of under‐sampled regions, taxa, and environments.

 
more » « less
NSF-PAR ID:
10390546
Author(s) / Creator(s):
 ;  ;  ;  
Publisher / Repository:
Wiley-Blackwell
Date Published:
Journal Name:
Biological Reviews
Volume:
98
Issue:
1
ISSN:
1464-7931
Page Range / eLocation ID:
p. 99-114
Format(s):
Medium: X
Sponsoring Org:
National Science Foundation
More Like this
  1. BACKGROUND The Republic of Madagascar is home to a unique assemblage of taxa and a diverse set of ecosystems. These high levels of diversity have arisen over millions of years through complex processes of speciation and extinction. Understanding this extraordinary diversity is crucial for highlighting its global importance and guiding urgent conservation efforts. However, despite the detailed knowledge that exists on some taxonomic groups, there are large knowledge gaps that remain to be filled. ADVANCES Our comprehensive analysis of major taxonomic groups in Madagascar summarizes information on the origin and evolution of terrestrial and freshwater biota, current species richness and endemism, and the utilization of this biodiversity by humans. The depth and breadth of Madagascar’s biodiversity—the product of millions of years of evolution in relative isolation —is still being uncovered. We report a recent acceleration in the scientific description of species but many remain relatively unknown, particularly fungi and most invertebrates. DIGITIZATION Digitization efforts are already increasing the resolution of species richness patterns and we highlight the crucial role of field- and collections-based research for advancing biodiversity knowledge in Madagascar. Phylogenetic diversity patterns mirror that of species richness and endemism in most of the analyzed groups. Among the new data presented, our update on plant numbers estimates 11,516 described vascular plant species native to Madagascar, of which 82% are endemic, in addition to 1215 bryophyte species, of which 28% are endemic. Humid forests are highlighted as centers of diversity because of their role as refugia and centers of recent and rapid radiations, but the distinct endemism of other areas such as the grassland-woodland mosaic of the Central Highlands and the spiny forest of the southwest is also important despite lower species richness. Endemism in Malagasy fungi remains poorly known given the lack of data on the total diversity and global distribution of species. However, our analysis has shown that ~75% of the fungal species detected by environmental sequencing have not been reported as occurring outside of Madagascar. Among the 1314 species of native terrestrial and freshwater vertebrates, levels of endemism are extremely high (90% overall)—all native nonflying terrestrial mammals and native amphibians are found nowhere else on Earth; further, 56% of the island’s birds, 81% of freshwater fishes, 95% of mammals, and 98% of reptile species are endemic. Little is known about endemism in insects, but data from the few well-studied groups on the island suggest that it is similarly high. The uses of Malagasy species are many, with much potential for the uncovering of useful traits for food, medicine, and climate mitigation. OUTLOOK Considerable work remains to be done to fully characterize Madagascar’s biodiversity and evolutionary history. The multitudes of known and potential uses of Malagasy species reported here, in conjunction with the inherent value of this unique and biodiverse region, reinforce the importance of conserving this unique biota in the face of major threats such as habitat loss and overexploitation. The gathering and analysis of data on Madagascar’s remarkable biota must continue and accelerate if we are to safeguard this unique and highly threatened subset of Earth’s biodiversity. Emergence and composition of Madagascar’s extraordinary biodiversity. Madagascar’s biota is the result of over 160 million years of evolution, mostly in geographic isolation, combined with sporadic long distance immigration events and local extinctions. (Left) We show the age of the oldest endemic Malagasy clade for major groups (from bottom to top): arthropods, bony fishes, reptiles, flatworms, birds, amphibians, flowering plants, mammals, non-flowering vascular plants, and mollusks). Humans arrived recently, some 10,000 to 2000 years (top right) and have directly or indirectly caused multiple extinctions (including hippopotamus, elephant birds, giant tortoises, and giant lemurs) and introduced many new species (such as dogs, zebu, rats, African bushpigs, goats, sheep, rice). Endemism is extremely high and unevenly distributed across the island (the heat map depicts Malagasy palm diversity, a group characteristic of the diverse humid forest). Human use of biodiversity is widespread, including 1916 plant species with reported uses. The scientific description of Malagasy biodiversity has accelerated greatly in recent years (bottom right), yet the diversity and evolution of many groups remain practically unknown, and many discoveries await. 
    more » « less
  2. Abstract

    Neotropical countries receive financing and effort from temperate nations to aid the conservation of migratory species that move between temperate and tropical regions. If allocated strategically, these resources could simultaneously contribute to other conservation initiatives. In this study, we use novel distribution maps to show how those resources could aid planning for the recovery of threatened resident vertebrates.

    Using eBird‐based relative abundance estimates, we first identified areas with high richness of Neotropical migrant landbirds of conservation concern (23 species) during the stationary non‐breeding period. Within these areas, we then identified threatened species richness, projected forest loss and conducted a prioritization for 1,261 red‐listed vertebrates using Terrestrial Area‐of‐Habitat maps.

    Richness for migrants was greatest along a corridor from the Yucatan peninsula south to the northern Andes but also included south‐west Mexico and Hispaniola. Protected areas account for 22% of this region while 21% is at risk of forest loss. Within this focal region for migrants, all four vertebrate groups showed hotspots of threatened species richness along the west and east Andean slopes. Taxa‐specific hotspots included montane areas of southern Mexico and central Guatemala (amphibians/reptiles) and the entire east slope of the Colombian East Andes (mammals).

    Our prioritization highlighted several areas of importance for conservation due to high threatened species richness and projected forest loss including (a) the Pacific dry forests of south‐west Mexico, (b) montane regions of northern Central America and (c) the west Andean slope of Colombia and Ecuador. At a landscape scale in southern Colombia, we show how conservation efforts for six Neotropical migrants could benefit 56 threatened residents that share a similar elevational range.

    Synthesis and applications. Funding and effort for migratory bird conservation also has potential to benefit threatened resident vertebrates in the Neotropics. Our study highlights how novel, high‐resolution information on species distributions and risk of forest loss can be integrated to identify priority areas for the two groups at regional and landscape scales. The approach and data can be further modified for more specific goals, such as within‐country initiatives.

     
    more » « less
  3. Abstract Aim

    Physiological tolerances and biotic interactions along habitat gradients are thought to influence species occurrence. Distributional differences caused by such forces are particularly noticeable on tropical mountains, where high species turnover along elevational gradients occurs over relatively short distances and elevational distributions of particular species can shift among mountains. Such shifts are interpreted as evidence of the importance of spatial variation in interspecific competition and habitat or climatic gradients. To assess the relative importance of competition and compression of habitat and climatic zones in setting range limits, we examined differences in elevational ranges of forest bird species among four Bornean mountains with distinct features.

    Location

    Bornean mountains Kinabalu, Mulu, Pueh and Topap Oso.

    Taxon

    Rain forest bird communities along elevational gradients.

    Methods

    We surveyed the elevational ranges of rain forest birds on four mountains in Borneo to test which environmental variables—habitat zone compression or presence of likely competitors—best predicted differences in elevational ranges of species among mountains. For this purpose, we used two complementary tests: a comparison of elevational range limits between pairs of mountains, and linear mixed models with naïve occupancy as the response variable.

    Results

    We found that lowland species occur higher in elevation on two small mountains compared to Mt. Mulu. This result is inconsistent with the expectation that distributions of habitats are elevationally compressed on small mountains, but is consistent with the hypothesis that a reduction in competition (likely diffuse) on short mountains, which largely lack montane specialist species, allows lowland species to occur higher in elevation. The relative influence of competition changes with elevation, and the correlation between lower range limits of montane species and the distribution of their competitors was weaker than in lowland species.

    Main conclusions

    These findings provide support for the importance of biotic interactions in setting elevational range limits of tropical bird species, although abiotic gradients explain the majority of distribution patterns. Thus, models predicting range shifts under climate change scenarios must include not only climatic variables, as is currently most common, but also information on potentially resulting changes in species interactions, especially for lowland species.

     
    more » « less
  4. Small-mammal faunas of tropical savannas consist of endemic assemblages of murid rodents, small marsupials, and insectivores on four continents. Small mammals in tropical savannas are understudied compared to other tropical habitats and other taxonomic groups (e.g., Afrotropical megafauna or Neotropical rainforest mammals). Their importance as prey, ecosystem engineers, disease reservoirs, and declining members of endemic biodiversity in tropical savannas compels us to understand the factors that regulate their abundance and diversity. We reviewed field studies published in the last 35 years that examined, mostly experimentally, the effects of varying three primary endogenous disturbances in tropical savanna ecosystems—fire, large mammalian herbivory (LMH), and drought—on abundance and diversity of non-volant small mammals. These disturbances are most likely to affect habitat structure (cover or concealment), food availability, or both, for ground-dwelling small mammalian herbivores, omnivores, and insectivores. Of 63 studies (included in 55 published papers) meeting these criteria from the Afrotropics, Neotropics, and northern Australia (none was found from southern Asia), 29 studies concluded that small mammals responded (mostly negatively) to a loss of cover (mostly from LMH and fire); four found evidence of increased predation on small mammals in lower-cover treatments (e.g., grazed or burned). Eighteen studies concluded a combination of food- and cover-limitation explained small-mammal responses to endogenous disturbances. Only two studies concluded small-mammal declines in response to habitat-altering disturbance were caused by food limitation and not related to cover reduction. Evidence to date indicates that abundance and richness of small savanna mammals, in general (with important exceptions), is enhanced by vegetative cover (especially tall grass, but sometimes shrub cover) as refugia for these prey species amid a “landscape of fear,” particularly for diurnal, non-cursorial, and non-fossorial species. These species have been called “decreasers” in response to cover reduction, whereas a minority of small-mammal species have been shown to be “increasers” or disturbance-tolerant. Complex relationships between endogenous disturbances and small-mammal food resources are important secondary factors, but only six studies manipulated or measured food resources simultaneous to habitat manipulations. While more such studies are needed, designing effective ones for cryptic consumer communities of omnivorous dietary opportunists is a significant challenge. 
    more » « less
  5. null (Ed.)
    Background Previous research has shown diverse vertical space use by various taxa, highlighting the importance of forest vertical structure. Yet, we know little about vertical space use of tropical forests, and we often fail to explore how this three-dimensional space use changes over time. Methods Here we use canopy tower systems in French Guiana and passive acoustic monitoring to measure Neotropical bat activity above and below the forest canopy throughout nine nights. We use a Bayesian generalized linear mixed effect model and kernel density estimates to demonstrate patterns in space-use over time. Results We found that different bats use both canopy and understory space differently and that these patterns change throughout the night. Overall, bats were more active above the canopy (including Cormura brevirostris, Molossus molossus, Peropteryx kappleri and Peropteryx macrotis ), but multiple species or acoustic complexes (when species identification was impossible) were more active in the understory (such as Centronycteris maximiliani, Myotis riparius, Pteronotus alitonus and Pteronotus rubiginosus ). We also found that most bats showed temporally-changing preferences in hourly activity. Some species were less active (e.g., P. kappleri and P. macrotis ), whereas others were more active ( Pteronotus gymnonotus, C. brevirostris , and M. molossus ) on nights with higher moon illuminance. Discussion Here we show that Neotropical bats use habitat above the forest canopy and within the forest understory differently throughout the night. While bats generally were more active above the forest canopy, we show that individual groups of bats use space differently over the course of a night, and some prefer the understory. This work highlights the need to consider diel cycles in studies of space use, as animals use different habitats during different periods of the day. 
    more » « less