Marcus Vinícius Vieira

Does the type of matrix matter? A quantitative review of the evidence

It has been increasingly recognized that the type of matrix surrounding habitat patches can affect biodiversity in landscapes, but there were only qualitative reviews of the subject focused on particular taxonomic groups. We present a quantitative review of studies from 1985 to 2008 that compared effects of different matrix types on individuals, populations and communities. We compiled 104 studies, most on animals, covering a broad range of landscape types and spatial scales. Most studies were empirical, focused on individuals and communities, and evaluated abundance/richness in the patch as the dependent variable. The type of matrix surrounding habitat patches influenced the studied parameters in 95% of the studies, but such effects were overall smaller compared to patch size or isolation effects. Matrix type effects were strongly species-specific, with different species responding differently to matrix type in 96% of studies comparing species or group of species. In 88% of studies, matrix types more similar in structure to the patch had higher quality for the studied organisms from the point of view of functional connectivity. Overall, the type of matrix is important, but patch size and isolation are the main determinants of ecological parameters in landscapes. Matrix quality generally increases with increasing structural similarity with habitat patches, a pattern that could be used as a general guideline for management of the matrix in fragmented landscapes.

Support diameter, incline, and vertical movements of four didelphid marsupials in the Atlantic forest of Brazil

Didelphids use vertical strata in different ways, suggesting the existence of a wider variety of niches than simply arboreal or terrestrial. This variety could be related to the differential ability to deal with support diameters and inclines, and might be important for the coexistence of species in local assemblages. From data obtained with the spool-and-line technique, the use of the vertical space and above-ground supports is described in four species of an assemblage of didelphid marsupials in south-east Brazil. The stratification in this and other assemblages in the Atlantic forest and in the Amazon is also compared. Animals were trapped in Serra dos Orgaos, state of Rio de Janeiro, and equipped with a spool-and-line device before release. The paths were tracked by measuring variables related to support diameter, incline, distance and height moved above ground. The diameter of supports used was positively related to body size, and the incline to the dominant direction of movement (horizontal vs vertical). The more cursorial species, Metachirus nudicaudatus, only once moved above ground in > 3200 m of paths followed. Didelphis aurita moved mostly on the ground, but was the only species that occasionally reached the canopy. Philander frenata also moved mostly on the ground, occasionally used the understorey, but never reached the canopy. The more arboreal species, Marmosops incanus, moved mostly in the understorey, but never in the canopy. This pattern of stratification is similar to that observed in other sites in the Atlantic forest and in the Amazon.

THE EFFECTS OF REPRODUCTIVE AND CLIMATIC SEASONS ON MOVEMENTS IN THE BLACK-EARED OPOSSUM (DIDELPHIS AURITA WIED-NEUWIED, 1826)

Abstract Resource availability and reproductive activity can exert contrasting demands on movements of promiscuous or polygynous mammals. Movements of the black-eared opossum (Didelphis aurita) vary seasonally in the Atlantic Rainforest of Brazil, so our objectives were to test the relative importance of reproductive activity and resource abundance on movement areas and intensity of use by opossums. We used a spool-and-line device to map the path of individuals, to estimate daily movement areas, and to measure intensity of habitat use. Two models were compared by generalized linear models, by grouping movements by reproductive or climatic season. Males used larger areas less intensively in the breeding season, whereas the movement of females did not differ between breeding and nonbreeding season. Differences between movements of females were significant only when grouped by climatic seasons, with females using larger areas less intensively in the dry season. Movements of females were determined by resource availability, whereas movements of males were determined by reproductive season.

Population responses of small mammals to food supply and predators: a global meta-analysis.

1. The relative importance of food supply and predation as determinants of animal population density is a topic of enduring debate among ecologists. To address it, many studies have tested the potential effects of food on population density by experimentally supplementing natural populations, with much focus on terrestrial vertebrates, especially small mammals. 2. Here we perform a meta-analysis of such experiments, testing two complementary hypotheses: (i) small mammal populations are bottom-up limited and (ii) population increases in response to food supplementation are constrained by predation, a top-down limitation. 3. In the 148 experiments recorded, food supplementation had an overall positive and significant effect, increasing population densities by 1.5-fold. Larger population increases occurred when predation was reduced and populations were open to immigration. Predation appeared to be unimportant when populations were closed to immigration. Immigration was the major mechanism underlying increases in abundance by increasing local population density and crowding. Contributions of increased reproductive rate could be detected, but were minor compared to immigration, and no effects were detected from survival. 4. Our analyses support the view that animal population density is determined by both bottom-up and top-down forces. They also suggest the possibility that food supplementation experiments might unintentionally create ecological traps by aggregating both prey and predators in small areas of the landscape. We suggest an alternative experimental design to increase the contribution that food supplementation experiments can make in future.

Movement behaviour within and beyond perceptual ranges in three small mammals: effects of matrix type and body mass.

1. For animal species inhabiting heterogeneous landscapes, the tortuosity of the dispersal path is a key determinant of the success in locating habitat patches. Path tortuosity within and beyond perceptual range must differ, and may be differently affected by intrinsic attributes of individuals and extrinsic environmental factors. Understanding how these factors interact to determine path tortuosity allows more accurate inference of successful movements between habitat patches. 2. We experimentally determined the effects of intrinsic (body mass and species identity) and extrinsic factors (distance to nearest forest fragment and matrix type) on the tortuosity of movements of three forest-dwelling didelphid marsupials, in a fragmented landscape of the Atlantic Forest, Brazil. 3. A total of 202 individuals were captured in forest fragments and released in three unsuitable matrix types (mowed pasture, abandoned pasture and manioc plantation), carrying spool-and-line devices. 4. Twenty-four models were formulated representing a priori hypotheses of major determinants of path tortuosity, grouped in three scenarios (only intrinsic factors, only extrinsic factors and models with combinations of both), and compared using a model selection approach. Models were tested separately for individuals released within the perceptual range of the species, and for individuals released beyond the perceptual range. 5. Matrix type strongly affected path tortuosity, with more obstructed matrix types hampering displacement of animals. Body mass was more important than species identity to determine path tortuosity, with larger animals moving more linearly. Increased distance to the fragment resulted in more tortuous paths, but actually reflects a threshold in perceptual range: linear paths within perceptual range, tortuous paths beyond. 6. The variables tested explained successfully path tortuosity, but only for animals released within the perceptual range. Other factors, such as wind intensity and direction of plantation rows, may be more important for individuals beyond their perceptual range. 7. Simplistic scenarios considering only intrinsic or extrinsic factors are inadequate to predict path tortuosity, and to infer dispersal success in heterogeneous landscapes. Perceptual range represents a fundamental threshold where the effects of matrix type, body mass and individual behaviour change drastically.

A device and standard variables to describe microhabitat structure of small mammals based on plant cover

Studies quantifying habitat structure generally use several instruments. This paper aims to propose a new and efficient device to characterize microhabitat structure of small mammals. Seven measurements were taken: plant cover, litter cover, rock cover, canopy cover, and vegetative obstruction at three heights. The device is a 0.25 m2 square wooden frame (0.50 m x 0.50 m) divided in to 100 open squares by wire mesh. Average time spent to measure each trapping station was six minutes. This new device is efficient, i.e., quick, practical, simple, and reliable. It can be used in any kind of forest. We propose this method as a standard method to describe habitat structure.

Nests and nesting sites of Brazilian forest rodents (Nectomys squamipes andOryzomys intermedius) as revealed by a spool-and-line device

Patterns of nest construction and nest site selection of two murid rodents of the Brazilian Atlantic rainforest were described. The animals were captured in an area of about 1.5 ha of primary forest, marked, equipped with a spool-and-line device and released at the capture point. The next day we followed the thread to find their nests. In total 11 nests ofNectomys squamipes (Brants, 1827) and 13 ofOryzomys intermedius (Leche, 1886) were found. They were all egg-shaped, about 15 cm long and 10 cm wide, without any apparent entrance and consisted of dry leaves and grass. The two species differed in the selection of nest sites. The nests ofN. squamipes were close to streams and built inside decomposed fallen logs or using natural entrances between the soil and tangled roots or rocky crevices. Nests ofO. intermedius were built under or inside fallen logs and inside aerial root systems of palms. The spool-and-line technique used here is useful to provide detailed information on several aspects of the ecology and natural history of small mammals.

Scale-dependent habitat selection in three didelphid marsupials using the spool-and-line technique in the Atlantic forest of Brazil

Habitat selection can be detected at different scales. For small mammals, habitat studies normally evaluate micro- or meso-habitat selection, but rarely are multiple scales analysed simultaneously. These studies use quantitative data of structural characteristics around the trap stations to evaluate selection. We test an alternative method to evaluate habitat selection in three didelphid marsupials (Didelphis aurita, Philander frenatus and Metachirus nudicaudatus), using a spool-and-line device, and measure habitat selection at two spatial scales. Habitat was characterized by seven habitat variables measured at ten points of direction change along the path traversed by each individual, and at 2 5 trap stations distributed in a 1 -ha area. Micro-habitat selection was estimated at each point by comparing habitat measures between the direction selected against other directions. Meso-habitat selection was estimated by comparing habitat measurements on the whole path of an individual against habitat measures on the 25 trap stations. Patterns of selection for each species could only be detected at the meso-habitat scale, although a few individuals in each species were selective at the micro-habitat scale. Studies of habitat selection need to address the scale of study quantitatively, if possible comparing two or more scales to determine the scale of selection.

Perception of a fragmented landscape by neotropical marsupials effects of body mass and environmental variables

Perceptual range is used as a measure of the ability of animals to perceive the landscape, and can be used to inter functional connectivity between habitat patches such as forest remnants. A relationship of perceptual range with body mass was proposed by Mech and Zollner, but effects of life-history traits and environmental conditions were also acknowledged. We evaluated perceptual ranges and the influence of body mass, wind speed and direction, and grass height in four marsupial species of the Atlantic Forest of south-eastern Brazil. Release experiments were conducted and orientation towards the forest was determined using the spool-and-line technique. In 111 animals released, Didelphis aurita had the highest perceptual range (200 m), followed by Philander frenatus and Micoureus paraguayanus (100 m), and by Marmosops incanus (< 100 m). None of the species presented homing behaviour, and there was no difference in orientation abilities between sexes. Perceptual range was overestimated based on the equations of Mech and Zollner, but species were ordered in perceptual range according to their body size. Distance to the forest was a common determinant of the orientation in P. frenatus and M. paraguayanus, but grass height and wind direction were also important. Orientation of Didelphis aurita was determined by wind direction, showing anemotactic behaviour. Body mass is likely to affect perceptual ranges in didelphid marsupials, but only in interspecific comparisons. Within a species perception depends on the environmental context. Variables other than distance must be considered to estimate functional connectivity of the landscape based on perceptual range.

Stride lengths and frequencies of arboreal walking in seven species of didelphid marsupials

Didelphid marsupials differ in their use of the forest strata, with corresponding differences in morphology and arboreal walking performances. Similar performances may be reached by different combinations of stride length and frequency, but it has been suggested that arboreal walkers increase velocity by longer strides. Our objective was to determine how stride length and frequency contribute to the velocity in the arboreal walking of seven species of didelphid marsupials of the Atlantic Forest of Brazil. Animals were stimulated to cross five 3-m long horizontal supports of different diameters. The cycle of maximum velocity was chosen to measure relative stride length, frequency, and relative velocity. Except forCaluromys philander, the more arboreal species were faster than the terrestrial species, but maximum velocity of arboreal species was reached by two strategies, increasing stride frequency (Gracilinanus microtarsus, Micoureus demerarae, andDidelphis aurita), or reducing frequency and increasing stride length (Marmosops incanus andC. philander). Increasing velocity in arboreal walking by more frequent strides may reduce oscillations of the body, whereas longer strides may reduce branch swaying. Among the terrestrial species,Philander frenatus performed similarly to more arboreal species, suggesting a potential ability to use the canopy, undetected in field observations.