Cribriform plate of ethmoid, olfactory bulb and olfactory acuity in forty species of bats

Relationships between the cribriform plate of the ethmoid, the olfactory bulb, and olfactory acuity were explored using material from 13 of the 17 bat families. All megachiropteran cribriform plates were entirely perforated. In contrast, microchiropteran plates showed distinct perforated portions dorsally and nonperforated portions ventrally. The plates of frugivorous species had more foramina than those of insectivorous ones. Bats with mixed dietary habits were intermediate. Our data suggest that the Chilonycterinae were originally frugivorous, and have only secondarily reverted to an insectivorous diet. Trend analyses show that wherever dietary preference appears to favor a more acute sense of smell, bulb diameter tends to be larger. In general, frugivorous bats tend to have bulbs exceeding 2 mm in diameter; insectivorous bats tend to have bulb diameters of 2 mm or less. The number of foramina in the plates and total cribriform plate area tends to increase as a function of bulb area, but the plate area the foramina occupied increases as a function of bulb volume. The ratio of the size of the bulb to the size of the cerebral hemisphere does not predict olfactory acuity in bats.     

Olfaction at depth: Cribriform plate size declines with dive depth and duration in aquatic arctoid carnivorans

It is widely accepted that obligate aquatic mammals, specifically toothed whales, rely relatively little on olfaction. There is less agreement about the importance of smell among aquatic mammals with residual ties to land, such as pinnipeds and sea otters. Field observations of marine carnivorans stress their keen use of smell while on land or pack ice. Yet, one dimension of olfactory ecology is often overlooked: while underwater, aquatic carnivorans forage “noseblind,” diving with nares closed, removed from airborne chemical cues. For this reason, we predicted marine carnivorans would have reduced olfactory anatomy relative to closely related terrestrial carnivorans. Moreover, because species that dive deeper and longer forage farther removed from surface scent cues, we predicted further reductions in their olfactory anatomy. To test these hypotheses, we looked to the cribriform plate (CP), a perforated bone in the posterior nasal chamber of mammals that serves as the only passageway for olfactory nerves crossing from the periphery to the olfactory bulb and thus covaries in size with relative olfactory innervation. Using CT scans and digital quantification, we compared CP morphology across Arctoidea, a clade at the interface of terrestrial and aquatic ecologies. We found that aquatic carnivoran species from two lineages that independently reinvaded marine environments (Pinnipedia and Mustelidae), have significantly reduced relative CP than terrestrial species. Furthermore, within these aquatic lineages, diving depth and duration were strongly correlated with CP loss, and the most extreme divers, elephant seals, displayed the greatest reductions. These observations suggest that CP reduction in carnivorans is an adaptive response to shifting selection pressures during secondary invasion of marine environments, particularly to foraging at great depths. Because the CP is fairly well preserved in the fossil record, using methods presented here to quantify CP morphology in extinct species could further clarify evolutionary patterns of olfactory loss across aquatic mammal lineages that have independently committed to life in water.     

Carnivore olfactory bulb size: allometry, phylogeny and ecology

Olfactory bulb size was measured in 146 species of Carnivora in order to examine whether recently observed functional patterns for overall brain size were similar for component parts of the brain. Comparative measures were analysed in relation to various allometric characters (body, brain and skull size), phylogeny, behaviour and ecology. Olfactory bulbs are significantly and positively correlated with all allometric variables, but indices of skull size correlate slightly more closely than other variables. This probably relates to functional aspects of skull size, facial proportions, and anterior elements of the brain. Phylogenetic associations were examined by two comparative methods: the method of independent contrasts and phylogenetic autoregression. Both revealed similar phylogenetic correlation at generic and familial levels. Using calculated values from either method, relative olfactory bulb size only correlates with zonation among seven behavioural and ecological variables; aquatic otters have smaller bulb sizes than carnivores of other zonal types. This agrees with discussion about the diminution of olfactory communication in aquatic environments. Also, olfactory bulb size correlates with home range size, which is consistent with a recent model on the use of olfaction for foraging in designated home ranges. Generally, comparative differences in olfactory bulb size in carnivores do not associate with functional variables found in other comparative studies. Nevertheless, future analyses of specific brain components in mammals may be more useful than overall brain size for testing evolutionary hypotheses of mammalian brain size.     

The ethmoid and presphenoid of cetaceans

A cribriform plate, a perpendicular plate, and two lateral masses are major components of the ethmoid bone of mammals. Notwithstanding the noticeable bone, virtually sitting in the center of the skull, extensive modifications of the skull of modern cetaceans, especially odontocetes (toothed whales), and the lack of clarity as to what characteristics delimit each element of the ethmoid has made the problem of the nature of the cetacean ethmoid more complicated and elusive than in other, less modified mammals. Furthermore, contention as to whether a perpendicular plate of the ethmoid, or the mesethmoid, exists in all mammals including cetaceans has remained unsettled. In odontocetes, the mesethmoid has been variably identified not only as the osseous nasal septum but also as the mediodorsal region of the posterior wall of the nasal passage below the nasals, as a mass of bone encased by the vomer in front of the osseous nasal cavity at the base of the rostrum, and as a combination of some portions mentioned above. The presence or absence of the mesethmoid in various groups of mammals has attracted the attention of some biologists, and here, I demonstrate that cetaceans have no mesethmoid. The close inspection of the ontogenetic changes of the basicranial elements in cetaceans reveals that a mass of bone ensheathed by the vomer in front, or at the level of the osseous nasal cavity is actually the presphenoid. It is highly likely that in odontocetes the posterior wall of the nasal passages below the nasals consists of the combination of the frontal, the imperforated cribriform plate, the paired ectethmoids, and the vomer, the latter three of which partially concealing the presphenoid dorsally and laterally as the ontogeny proceeds. In contrast, mysticetes clearly display ethmoturbinates and a cribriform plate, which are morphologically similar to those in terrestrial mammals. J. Morphol. 277:1661–1674, 2016. © 2016 Wiley Periodicals, Inc.     

Olfactory receptor gene repertoires in mammals

In mammals, olfaction is mediated by two distinct organs that are located in the nasal cavity: the main olfactory epithelium (MOE) that binds volatile odorants is responsible for the conscious perception of odors, and the vomeronasal organ (VNO) that binds pheromones is responsible for various behavioral and neuroendocrine responses between individuals of a same species. Odorants and pheromones bind to seven transmembrane domain G-protein-coupled receptors that permit signal transduction. These receptors are encoded by large multigene families that evolved in mammal species in function of specific olfactory needs.     

From Jumbo to Dumbo: Cranial Shape Changes in Elephants and Hippos During Phyletic Dwarfing

Members of the mammalian families Elephantidae and Hippopotamidae (extant and extinct elephants and hippos) include extinct dwarf species that display up to 98% decrease in body size compared to probable ancestral sources. In addition to differences in body mass, skulls of these species consistently display distinctive morphological changes, including major reduction of pneumatised areas in dwarf elephants and shortened muzzles in dwarf hippos. Here we build on previous studies of island dwarf species by conducting a geometric morphometric analysis of skull morphology and allometry in target taxa, living and extinct, and elaborate on the relation between skull size and body size. Our analysis indicates that skull size and body size within terrestrial placental mammals scale almost isometrically (PGLS major axis slope 0.906). Furthermore, skull shape in dwarf species differed from both their ancestors and the juveniles of extant species. In insular dwarf hippos, the skull was subject to considerable anatomical reorganisation in response to distinct selection pressures affecting early ontogeny (the “island syndrome”). By contrast, skull shape in adult insular dwarf elephants can be explained well by allometric effects; selection on size may thus have been the main driver of skull shape in dwarf elephants. We suggest that a tightly constrained growth trajectory, without major anatomical reorganization of the skull, allowed for flexible adaptations to changing environments and was one of the factors underlying the evolutionary success of insular dwarf elephants.     

Variability,morphometrics, and co-variation of the os lacrimale in Cervidae

In Ruminantia, the lacrimal bone forms a considerable part of the facial skeleton, and the morphology of its facial facet is highly variable when compared to other mammals. In this study, we quantify the species-specific variability in size and shape of the lacrimal facial facet in species of Cervidae (deer) and relate it to systematics and various aspects of their ecology and behavior. We sampled 143 skull specimens from 10 genera; 12 Moschus and 3 Tragulus specimens were used as outgroups. We find that size and shape of the lacrimal facial facet allow differentiating most species analyzed here, except for Mazama gouazoubira and Capreolus capreolus. Size and shape of the lacrimal facial facet vary widely across Cervidae regardless of their systematic relationships, ecology or behavior. Thus, we could not detect a unique signature of adaptational criteria in lacrimal morphology. Our data indicate that the lacrimal facial facet scales allometrically with skull size, in particular, the lacrimojugal length scales positively and the lacrimomaxillar length scales negatively. However, correlation analyses did not reveal any differences in the integration of the lacrimal bone with any specific skull module in any of the species compared. Lastly, we could not ascertain any correlation between the size and position of the preorbital depression with the size and shape of the lacrimal facial facet. We conclude that the lacrimal facial facet is highly flexible and may rapidly adjust to its surrounding bones. Its allometric growth appears to be an example of exaptation: changes in size and shape in the context of the increase of the skull length provide lacrimal contacts, in particular, a lacrimojugal one, which may serve to reduce mechanical loads resulting from increasingly larger antlers in large cervids.     

Allometry in the distribution of material properties and geometry of the felid skull: why larger species may need to change and how they may achieve it

Extant members of the cat family (Felidae) have been considered behaviourally and morphologically conservative, i.e., despite great differences in size, there is relatively little variation in either the shape of the felid skull and dentition across species, or in the way in which these structures are used to kill and dismember prey. Consequently felids have been considered an appropriate focus for a number of investigations into the influence of allometry on craniomandibular mechanics and morphology. However, although previous treatments have considered the role of shape, they have not investigated the influence of differences in the distribution of relatively stiff cortical and more compliant cancellous bone on performance. Here, using models that incorporate material properties for both cortical and cancellous bone, we apply three-dimensional (3D) finite element analysis (FEA) to models representing the skulls of seven extant felid species. Our objectives being to determine allometric trends regarding both overall geometry and the relative distributions of cortical and cancellous bone tissue. We also more comprehensively assess variation in the efficiency with which muscular force is converted to bite force and the capacity to resist associated stresses. Our results show that the cheetah (Acinonyx jubatus) may be exceptional regarding both the efficiency with which muscular force is converted to bite force and the distribution of stress. We found a negative allometric trend between cortical bone volume and total skull bone volume, and positive allometry between the total skull bone volume and skull surface area. Results gained from mathematical modelling of beam analogies suggest that these trends reflect a need for larger species to respond to physical challenges associated with increased size, and, that changes in skull shape, bone composition, or a combination of both may be required to accommodate these challenges. With geometrical scaling stress increases by the same factor, and displacement by the same factor squared, but the ultimate failure stress of the material is invariant. We find that as species become larger, overall skull bone volume relative to surface area increases by adding a higher proportion of less dense and more compliant cancellous bone. This results in an increased cross-sectional area and second moment of inertia, which acts to reduce the overall stresses. An overall saving in mass is a likely additional consequence. Although we do find evidence that skull stiffness does diminish with size, we also argue that this is at least in part mitigated through the influence of these allometric trends. We further suggest that these trends and the explanations for them may be universal for vertebrates.     

Molecular biology of pheromone perception in mammals

In mammals, olfactory sensory perception is mediated by two anatomically and functionally distinct sensory organs: the main olfactory epithelium (MOE) and the vomeronasal organ (VNO). Pheromones activate the VNO and elicit a characteristic array of innate reproductive and social behaviors, along with dramatic neuroendocrine responses. Recent approaches have provided new insights into the molecular biology of sensory transduction in the vomeronasal organ. Differential screening of cDNA libraries constructed from single sensory neurons from the rat VNO has led to the isolation of a family of genes which are likely to encode mammalian pheromone receptors. The isolation of these receptors from the vomeronasal organ might permit the analysis of the molecular events which translate the bindings of pheromones into innate stereotypic behaviors and help to elucidate the logic of pheromone perception in mammals.     

Blood flow to long bones indicates activity metabolism in mammals, reptiles and dinosaurs

The cross-sectional area of a nutrient foramen of a long bone is related to blood flow requirements of the internal bone cells that are essential for dynamic bone remodelling. Foramen area increases with body size in parallel among living mammals and non-varanid reptiles, but is significantly larger in mammals. An index of blood flow rate through the foramina is about 10 times higher in mammals than in reptiles, and even higher if differences in blood pressure are considered. The scaling of foramen size correlates well with maximum whole-body metabolic rate during exercise in mammals and reptiles, but less well with resting metabolic rate. This relates to the role of blood flow associated with bone remodelling during and following activity. Mammals and varanid lizards have much higher aerobic metabolic rates and exercise-induced bone remodelling than non-varanid reptiles. Foramen areas of 10 species of dinosaur from five taxonomic groups are generally larger than from mammals, indicating a routinely highly active and aerobic lifestyle. The simple measurement holds possibilities offers the possibility of assessing other groups of extinct and living vertebrates in relation to body size, behaviour and habitat.     

Vergleichend quantitative Untersuchungen an Organen von Caviomorpha

Comparative quantitative investigations of organs of Caviomorpha . The weights of the organs: heart, liver, lung, spleen, kidneys and stomach-intestines were measured in nine species of caviomorph rodents. The sizes of these organs were compared using allometry formulae calculated for each organ from the Caviomorpha to the allometric lines of mammals (mouse-to-elephant line). The slopes of the two sets were nearly equal, except for the lung/body slope. In organ sizes, independently from body size, most of the caviomorph species were found to be outlying with respect to the reference Mammalia. Thus, compared to the general mammalian-line the organ sizes of the caviomorph vary between an increase of 47% in the kidneys of Microcavia australis and a decrease of 68% in the spleen weight of Lagostomus maximus . It is argued that these differences in size suggest a strong correlation to the functional importance of the organs, and that the large range of variation in organ size is regarded as a process of functional adaptation to different life styles within caviomorph rodents.     

The structure of the nasal chemosensory system in squamate reptiles. 2. Lubricatory capacity of the vomeronasal organ

The vomeronasal organ is a poorly understood accessory olfactory organ, present in many tetrapods. In mammals, amphibians and lepidosaurian reptiles, it is an encapsulated structure with a central, fluid-filled lumen. The morphology of the lubricatory system of the vomeronasal organ (the source of this fluid) varies among classes, being either intrinsic (mammalian and caecilian amphibian vomeronasal glands) or extrinsic (anuran and urodele nasal glands). In the few squamate reptiles thus far examined, there are no submucosal vomeronasal glands. In this study, we examined the vomeronasal organs of several species of Australian squamates using histological, histochemical and ultrastructural techniques, with the goal of determining the morphology of the lubricatory system in the vomeronasal organ. Histochemically, the fluid within the vomeronasal organ of all squamates is mucoserous, though it is uncertain whether mucous and serous constituents constitute separate components. The vomeronasal organ produces few secretory granules intrinsically, implying an extrinsic source for the luminal fluid. Of three possible candidates, the Harderian gland is the most likely extrinsic source of this secretion.     

Localization of choline acetyltransferase and vasoactive intestinal polypeptide-like immunoreactivity in the nervus terminalis of the fetal and neonatal rat

Ganglia of the nervus terminalis have been shown to contain luteinizing hormone-releasing hormone (LHRH) immunoreactive cells in several mammalian species. These cells are always accompanied by clusters of cells non-immunoreactive to antiserum to LHRH. Using immunocytochemical procedures, we found choline acetyltransferase (ChAT) and vasoactive intestinal polypeptide (VIP) present in cell bodies and in nerve processes throughout the peripheral, intracranial and central projections of the nervus terminalis. In addition, a dense plexus of substance P (SP) immunoreactive fibers was seen in the nasal mucosa surrounding the nasal glandular acini and blood vessels. A number of SP reactive fibers were traced with the olfactory nerves through the cribriform plate of the ethmoid bone and appeared to enter the brain in the area of the central roots of the nervus terminalis.     

Functional Morphology of the Olfactory Mucosa and Olfactory Bulb in Fossorial Rodents: The East African Root Rat (Tachyoryctes splendens) and the Naked Mole Rat (Heterocephalus glaber)

Optimal functioning of the olfactory system is critical for survival of fossorial rodents in their subterranean lifestyle. This study examines the structure of the olfactory mucosa and olfactory bulb of two fossorial rodents exhibiting distinct social behaviors, the East African root rat and the naked mole rat. The social naked mole rat displayed simpler ethmoturbinates consisting of dorsomedial and broad discoid/flaplike parts that projected rostrally from the ethmoid bone. In the solitary root rat however, the ethmoturbinates were highly complex and exhibited elaborate branching which greatly increased the olfactory surface area. In addition, when correlated with the whole brain, the volume of the olfactory bulbs was greater in the root rat (4.24 × 10^?2) than in the naked mole rat (3.92 × 10^?2). Results of this study suggest that the olfactory system of the root rat is better specialized than that of the naked mole rat indicating a higher level of dependence on this system since it leads a solitary life. The naked mole rat to the contrary may have compensated for its relatively inferior olfactory system by living in groups in a social system. These findings demonstrate that structure of the olfactory system of fossorial mammals is dictated by both behavior and habitat.     

The mechanism of olfactory organ ventilation in Periophthalmus barbarus (Gobiidae, Oxudercinae)

Periophthalmus barbarus Linnaeus, 1766 has many adaptations for amphibious life as a consequence of tidal zone occupation. One of them is the ability to keep a little amount of water and air in mouth while on land or in hypoxic water, correlated with closing a gill lid for gas exchange improvement. It causes that mechanisms of olfactory organ ventilation described in other species of actinopterygians (compression of accessory nasal sac(s) by the skull and jaw elements while mouth and gill lid moving) are not in operation. There is a specific mechanism of olfactory organ ventilation independent on jaw and skull elements movements. Compression of accessory nasal sacs is possible by a0 contraction and it is a movement effect on bones combined by ligaments. This process can be observed on P. barbarus as lifting the rostral part of the head.     

Of mice and wrens: the relation between abundance and geographic range size in British mammals and birds

We examine the relation between population size and geographic range size for British breeding birds and mammals. As for most other assemblages studied, a strong positive interspecific correlation is found in both taxa. The relation is also recovered once the phylogenetic relatedness of species has been controlled for using an evolutionary comparative method. The slope of the relation is steeper for birds than for mammals, but this is due in large part to two species of mammals that have much higher population sizes than expected from their small geographic ranges. These outlying mammal species are the only ones in Britain to be found only on small offshore islands, and so may be exhibiting density compensation effects. With them excluded, the slope of the abundance–range size relation for mammals is not significantly different to that for birds. However, the elevation of the relation is higher for mammals than for birds, indicating that mammals are approximately 30 times more abundant than birds of equivalent geographic range size. An earlier study of these assemblages showed that, for a given body mass, bats had abundances more similar to birds than to non-volant mammals, suggesting that the difference in abundance between mammals and birds might be due to constraints of flight. Our analyses show that the abundance–range size relation for bats is not different for that from other mammals, and that the anomalously low abundance of bats for their body mass may result because they have smaller than expected geographic extents for their size. Other reasons why birds and mammals might have different elevations for the relation between population size and geographic range size are discussed, together with possible reasons for why the slopes of these relations might be similar.     

Expression and function of the empty spiracles gene in olfactory sense organ development of Drosophila melanogaster

In Drosophila, the cephalic gap gene empty spiracles plays key roles in embryonic patterning of the peripheral and central nervous system. During postembryonic development, it is involved in the development of central olfactory circuitry in the antennal lobe of the adult. However, its possible role in the postembryonic development of peripheral olfactory sense organs has not been investigated. Here, we show that empty spiracles acts in a subset of precursors that generate the olfactory sense organs of the adult antenna. All empty spiracles-expressing precursor cells co-express the proneural gene amos and the early patterning gene lozenge. Moreover, the expression of empty spiracles in these precursor cells is dependent on both amos and lozenge. Functional analysis reveals two distinct roles of empty spiracles in the development of olfactory sense organs. Genetic interaction studies in a lozenge-sensitized background uncover a requirement of empty spiracles in the formation of trichoid and basiconic olfactory sensilla. MARCM-based clonal mutant analysis reveals an additional role during axonal targeting of olfactory sensory neurons to glomeruli within the antennal lobe. Our findings on empty spiracles action in olfactory sense organ development complement previous studies that demonstrate its requirement in olfactory interneurons and, taken together with studies on the murine homologs of empty spiracles, suggest that conserved molecular genetic programs might be responsible for the formation of both peripheral and central olfactory circuitry in insects and mammals.     

脊椎动物中的一种解剖学新模型——黄鳝外周嗅觉系统

除单鼻型的圆口类外, 脊椎动物的左、右两侧嗅觉器官和嗅神经皆互为独立地分布于头前端, 而且它们的前鼻孔(外鼻孔)、嗅腔、嗅觉副囊腔(部分鱼具嗅觉副囊)与后鼻孔(或内鼻孔)也都互为相通, 且多呈开放状态。它们还通常具有一个体积相对较大且较稳定的嗅腔, 而嗅上皮则多位于嗅腔的一侧。此外, 鱼类的嗅囊与鼻窝之间通常也无明显间隙。然而, 运用常规的解剖学方法发现, 黄鳝(Monopterus albus)外周嗅觉系统(嗅觉器官和嗅神经)在解剖结构上已发生如下重大变化: (1)虽然具有前、后鼻孔, 但两者互不相通, 而嗅腔仅靠前鼻孔通至外界; (2)两侧嗅囊的末端及两侧嗅神经的前段均分别发生了合并。此外, 在该鱼上还发现:(1)嗅囊为一柔软而扁塌的长管囊结构, 其唯一的开口(即位于前鼻孔球上的前鼻孔)却常呈关闭状, 故此时该嗅腔实际上是一个体积被压扁到最小且暂时被封闭的空间; (2)嗅囊纵向地贴附于长鼻窝的内侧壁上, 它仅占鼻窝的一小部分空间, 故鼻窝显得相对很宽敞; (3)嗅觉副囊不与嗅腔相通, 而与鼻窝共同经后鼻孔通至外界; (4)两侧嗅囊的末端相向地穿越鼻窝内侧壁, 进入筛骨与额骨之间的“筛-额横管”, 在那里发生嗅囊合并;(5)嗅囊壁周缘几乎都内衬着嗅上皮, 且具数个褶窝(说明该嗅囊有扩张的可能)。因此, 黄鳝的这套解剖学特征不同于包括鱼类在内的所有脊椎动物的外周嗅觉系统。研究所发现的黄鳝这套形态学特征不仅为脊椎动物外周嗅觉系统的研究提供了一个独特的解剖学新模型, 同时也为动物进化研究提供了一个有关前、后鼻孔互不相通的进化特例。此外, 研究还依据上述发现提出嗅囊扩张-压缩假说以解释气味媒质进出于黄鳝这种特殊嗅腔的动力学机制。       

The role of olfaction throughout juvenile development: Functional adaptations in elasmobranchs

Seven elasmobranch species, a group known for their highly‐developed sense of smell, were examined for developmental changes in the number of olfactory lamellae, the size of the surface area of the sensory olfactory epithelium and the mass of both the olfactory rosettes (primary input to the CNS), and the olfactory bulbs. Within each species, juveniles possessed miniature versions of the adult olfactory organs, visually not distinguishable from these and without any obvious structural differences (e.g., with respect to the number of lamellae and the extent of secondary folding) between differently sized individuals. The size of the olfactory organs was positively correlated with body length and body mass, although few species showed proportional size scaling. In Aetobatus narinari and Aptychotrema rostrata, olfactory structures increased in proportion to body size. With respect to the growth of the olfactory bulb, all species showed allometric but not proportional growth. Olfaction may be of particular importance to juveniles in general, which are often subjected to heavy predation rates and fierce inter/intraspecific competition. Accordingly, it would be advantageous to possess a fully functional olfactory system early on in development. Slow growth rates of olfactory structures could then be attributed to a greater reliance on other sensory systems with increasing age or simply be regarded as maintaining an already optimized olfactory system. J. Morphol., 2010. © 2009 Wiley‐Liss, Inc.     

Habitat loss and mammalian extinction patterns: are the reserves in the Quadrilátero Ferrífero, southeastern Brazil, effective in conserving mammals?

Habitat loss is considered to be the principal cause of the local extinction of mammals worldwide. We assessed the extinction pattern of medium- and large-sized mammals caused by the effects of habitat loss in reserves in the Quadrilátero Ferrífero, southeastern Brazil, and discussed the effectiveness of these natural remnants for conserving mammals. A literature review and field collections were conducted from 2006 to 2011 to estimate the composition and richness of mammals in nine remnants of different sizes, including reserves and non-protected areas. A species–area relation and a nested subset analysis were performed, and a degree of sensitivity to habitat loss was obtained for each species according to its frequency of occurrence. Forty-five species of mammals were recorded. There was a strong species–area relation involving the legal size of reserves. High species richness was associated with large reserves, and the z value was within the range of very isolated continental remnants. The mammalian community exhibited a nested occurrence pattern, suggesting that most species were part of a more continuous ecosystem and that non-random extinction caused by habitat loss occurred in southeastern Brazil. The negative relation found between species frequencies and body weights suggested that selective species loss is associated with decreases in the size of the reserves. The estimated viable size required to conserve all of the sensitive species is greater than the size of the largest reserve inventoried. We recommend the aggregation of neighboring natural remnants and the creation of new reserves to reduce extinction risks.