SMALL MAMMAL INVENTORIES IN AN EASTERN BRAZILIAN PARK Jody R. Stallings* ABSTRACT Small mammal inventories were conducted between September 1985 and February 1987 in native forest, exotic forest, and open field habitats in a state forestry park in southeastern Brazil. In 40,490 trap nights, 17 species of non-volant small mammals were captured. This small mammal fauna was composed of 6 species of didelphid marsupials and 11 species of r6dents. Open field habitats were dominated, in terms of species richness and relative abundance, by rodents. One species, Akodon cursor, represented 85% of the captures in this habitat type. Forested habitats, both native and exotic, were composed of more species of rodents, but higher relative densities of didelphid marsupials. Didelphid marsupials Metachirus nudicaudams, Marmosa incana, and M. cinerea were the three most common and ubiquitous forest-dwelling species captured during the inventory. Exotic Euca6pms forests with native species subcanopy help to maintain the species diversity of small mammals in a landscape greatly altered by human activities. RESUMO Inventarios de pequenos mamiferos foram realizados entre Setembro de 1985 e Fevereiro de 1987 em habitats de florestas nativa, exotica e campo situados no Parque Estadual da Floresta do Rio Doce no Estado de Minas Gerais, Brasil. Em 40.490 armadilhas-noite, foram capturadas 17 esp6cies de pequenos mamiferos terrestres e arb6reos. Esta fauna de pequenos mamfferos compreendeu 6 esp6cies de marsupiais didelfideos e 11 esp6cies de roedores. Os habitats de campo foram dominados em termos de esp6cies e relativa abundancia, pelos roedores. Uma especie, Akodon cursor, representou 85% das capturas neste tipo de habitat. Os habitats de floresta, tanto nativa como exotica, apresentou um maior numero de esp6cies de roedores, porem com uma maior diversidade relativa de mai'supiais didelfideos. Os marsupiais didelfideos Metachirus nudicaudatus, Marmosa incana, e M. cinerea foram as tres espEcies mais comuns capturadas durante o inventario, no habitat florestal. As florestas exoticas de Euca6ptus, com um sub-bosque de esp6cies nativas, auxiliam na manutengao da diversidade de esp6cies de pequenos mamfferos num ambiente grandemente alterado pela aqao antropica. * The author is in the Departamento de Zoologia, instituto de Ciencias Biologicas, Universidade. Federal de Minas Gerais, Belo Horizonte, M.G. 30000, Brazil. He was formerg a Post Doctoral Associate in the Department of Wildlife and Range Sciences, School of Forest Resources and Conservation, University of Florida. Stallings, J. R. 1989. Small Mammal Inventories in an Eastern Brazilian Park. Bull. Florida State Mus., Biol. Sci. 34(4):153-200. 160 BULLEI'IN FLORIDA SI'ATE MUSEUM 34(4) TABLE OF CONTENTS Introdi,rtinn 160 Acknowledgements 161 Study Sitp 162 Methrvis 163 RM„ 1,9 168 Species Accounts 173 Discuscion 180 Literature Cited 184 Tphip. 187 Appendiv 1 194 INTRODUCTION Despite the country's great size, few studies have been conducted on small mammal communities in Brazil. Most published reports on Brazilian mammals are preliminary species lists (e.g. Avila-Pires and Gouvea 19'77) or inventories (e.g. Moojen 1952; Vieira 1955). Some Brazilian studies have focused on densities (Emmons 1984), others on abiotic effects on small mammals (Borchert and Hansen 1983; Peterson in press). Still others have addressed such applied subjects as plantation effects (Dietz et al. 1975) and public health needs or mammals that carry human diseases (e.g. Laemment et al. 1946; Botelho and Linardi 1980; Dias 1982). Recently, work in Brazil using mark-release techniques has addressed the use of space, longevity, diversity, and social habits of small mammals (see Alho 1982). As Alho (1982) indicated, most of these studies were concentrated in the xerophitic Cerrado and Caatinga habitats (e.g. Lacher 1981; Mares et al. 1981; Fonseca and Redford 1985; Redford and Fonseca 1986; Nitikman and Mares 1987; Streilein 1982). Fewer studies have been carried out in humid forest. Carvalho (1965) live-trapped small mammals in a tropical humid forest in Sao Paulo, and J. Malcolm (pers. comm.) conducted similar studies in another such forest near Manaus, Amazonia. Fonseca (1988) worked on small mammals in a range of forested habitats in eastern Minas Gerais. The Brazilian Atlantic forest has a highly diverse flora and fauna, with many endemic species of trees (Mori et al. 1981), reptiles (Muller 1973), and birds (Haffer 1974). In-depth mammal inventories in the region are lacking. The mammalian fauna is poorly known. Mittermeier et al. (1982) and Kinzey (1982) reported on the high level of diversity and endemism found in the primates of the region. Preliminary species lists for non-volant mammals in this region also suggest very high diversity and endemism (Moojen 1952; Vieira 1955; Cabrera 1957, 1961; Honacki et al. 1982). For this paper, a preliminary analysis was conducted of the non-volant mammal species that probably occur in the Atlantic Forest region. These data indicate that for the region there are SrALLINGS: BRAZILIAN SMALL MAMMAL INVENTORIES 161 at least 130 species, 54 of which (42%) are endemic. Didelphid marsupials and rodents account for 78% of the endemic species and 82% of the endemic genera. The purpose of this paper is to report the results of intensive inventories of small mammals in the tropical humid Rio Doce State Forestry Park, Minas Gerais, Brazil. These inventories were part of a larger overall project concentrating on small mammal communities in the Brazilian Atlantic Forest. Gustavo A. B. da Fonseca concentrated his efforts on the effect of habitat size and disturbance on small mammal species diversity in eastern Minas Gerais state. The small mammal biology and natural history observations from his research are presented in Fonseca and Kierulff (This volume). I investigated the effect of forest fire on small mammal communities in the Rio Doce State Forestry Park, eastern Minas Gerais state. In-depth small mammal inventories are lacking from this Park. Gastal (1982) and Avila-Pires (1978) reported preliminary results from intermittent small mammal inventories in this sanctuary. ACKNOWLEDGEMENTS Foremost, I thank John Robinson, my Ph. D. committee chairman, for his continued support throughout the various phases of this project and for his suggestion to work on small mammals in Brazil. Dr. Robinson, along with John Eisenberg, Mel Sunquist, Wayne Marion, S. David Webb, and Oliver Pearson made constructive comments on the manuscript. John Robinson and Kent Redford were instrumental in helping me design and implement the project. L. Aguiar, L. P. de Souza Silva, and E. Sabato helped collect the field data. J. Ladeira, H. Silva-Neto, and A. Lopes solved daily logistical problems in the Park. Housing in the Park was provided by the Instituto Estadual de Florestas. G. Fonseca, C. Valle, and I. Santos helped me obtain my visa to work in Brazil and offered laboratory space in the Federal University of Minas Gerais. This project was funded by an Organization of American States Fellowship and by the Program for Studies in Tropical Conservation. Gustavo and Ana Fonseca opened their doors to me and my family, and we will always be grateful for the assistance that they provided in helping us become acclimated to Brazil. This is contribution No. 35 from the Program for Studies in Tropical Conservation. STUDY SITE Small mammal trapping was carried out in the Rio Doce State Forestry Park (19°48'18" and 19°29'24" south latitude and 42°38'30" and 42°28'18" west longitude). The Park was created in 1944 at the request of Dom Helvecio, the bishop of the region (Gilhaus 1986). The State Forestry Institute of the state of Minas Gerais is the present administrative body. The climate of the Park is classified as tropical humid (Gilhaus 1986) with a seasonal pulse of precipitation from November through February and a pronounced dry season from June through August. Average annual rainfall for 162 BULLETIN FLORIDA STATE MUSEUM 34(4) a 20-year period (CETEC 1981) was 1480 mm, although the rainfall recorded during the study year was considerably less (Fig. 1). Mean annual temperature averaged 22°C (CETEC 1981), and mean minimum monthly temperatures varied greatly throughout the year (Fig. 2). The Park boundaries on the north and the east are two rivers, the Rio Piracicaba and the,Rio Doce, respectively (Fig. 3). The southern and western boundaries abut plantations of Euca6ptus spp. The altitude in the Park varies from 230 m to 515 m. CETEC (1981) reported that 21% of the Park is composed of plains, 40% undulating to strongly undulating hills, and 34% strongly undulating hills to mountainous terrain. The predominant terrain derives from dissection of fluvial plains (Gilhaus 1986). A unique feature of the Park is the system of lakes in the Rio Doce Valley. Approximately 40 lakes and numerous marshes within the Park's boundaries were formed by damming the drainage river of the Rio Doce watershed (Saijo and Tundisi 1985). The marshes are the result of the sedimentation of previous lakes. The vegetation of the park is classified as tropical semi-deciduous (Gilhaus 1986). Most of the emergent trees lose their leaves during the cool dry months. Forest fire has been a major threat to the vegetation and the wildlife in the park because of the litter that accumulates during the dry season. In 1964 and 1967, major fires burned approximately 30% of the park (Lopes 1982; Silva-Neto 1984). METHODS Habitats Studied Small mammals were live-trapped in four habitat types within the Park and in an exotic habitat type near the Park boundary. Five forested and five open/field habitats have been described for the Park by Gilhaus (1986). These are related to the four habitats that were sampled in the park (Table 1). I trapped in five sites within the native forested habitat category. Two of the sites, Rio Doce/Campolina (RD/C) and Rio Doce/Turvo (RD/T), were primary forests and corresponded to the Gilhaus (1986) classification of Tall Primary Forest with Epiphytes. All of the forested and open/field habitats have been altered to some extent by fire, with the exception of the Tall Primary Forest with Epiphytes. With respect to this study, two sites, Rio Doce/Hotel (RD/H) and Rio Doce/Misturado (RD/M) were altered by forest fire in 1967 and burned in an intermediate fashion which produced a forest mosaic of short, secondary, and tall forest. This habitat type corresponded to the SrALLINGS: BRAZILIAN SMALL MAMMAL INVENTORIES 163 MONTHLY RAINFALL (MM] MEAN TEMPERATURE (C) MONTHLY RAINFALL (MM) MEAN TEMPERZURE (C) 400 .0 400 140 ~ SURPLUS ~ SURPLUS120 -120 300 100 , 00300 - (3 DEFICIT 00 ~ DEFICIT eo 200 200 80 60 40 40100 100 20 20 O 0 0 0 J»N FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC JAN FEB MAR *PR MAY JUN JUL AUG SEP OCT NOV DEC MONTHS MONTHS CATI FROM 1964-1974 -TA FAOM .86-1.6 A B Figure 1. Walter and Leith climatic diagrams characterizing precipitation surplus and deficit per month in Rio Doce State Forestry Park, Minas Gerais, Brazil. Y axis is monthly rainfall in mm and Y axis in mean temperature CC). A = data collected for a 20-year period (1954-1974), and B = data collected during present study. TEMPERATURE (C ) 35 30 - 25 - 20 I l 15 - 10 - 5- 0 ' ' ' ' 'lilli NOV DEC JAN FEB MAR APR MAY JUN JUL AUG SEP OCT MONTHS MEAN MAXIMUM TEMP -4- MEAN MINIMUM TEMP Figure 2. Temperature graph demonstrating the pronounced decrease in minimum temperature during June, July, and August in the Rio Doce State Forestry Park, Minas Gerais, Brazil. 164 BULLETIN FLORIDA STATE MUSEUM 34(4) BRAZIL RIO 190 29'PIRACIC BA ATLANTIC FOREST PFERD RIO DOCE PFERD 'C RIO TURVO 0 4 KM - 190 48' Figure 3. Map of Rio Doce State Forestry Park, Minas Gerais, Brazil. Rivers Piracicaba and Doce form the northern and southern boundaries of the Park respectively. Trapping sites are indicated by circled letters: E = eucalypt forest with native species subcanopy (RD/E); B = wet meadow habitat (RD/B); F = homogeneous short secondary forest (RD/F); M = mosaic habitat of secondary and primary forest (RD/M); H = mosaic habitat of sec6ndary and primary forest (RD/H); T = primary forest habitat (RD/I); C = primaiy forest habitat (RD/C). SrALLINGS BRAZILIAN SMALL MAMMAL INVENTORIES 165 Medium to Tail Forest with Bamboos and Graminoids. The remaining native forest site, Rio Doce/Fogo (RD/F), was burned completely to the ground in 1967, and the resulting vegetation type corresponded to the Medium Secondary Forest with Bamboos and Graminoids. The wet meadow habitat, Rio Doce/Brejo (RD/B), corresponded to both the Low Woodland and Low Tree and Scrub Tallgrass Savanna classified by Gilhaus (1986). This habitat type occurs between the edge of permanent marshes and secondary forest. Grasses were the dominant vegetative cover and were introduced in the region as food for cattle. The eucalypt forest with native subcanopy habitat, Rio Doce/Eucalypt (RD/E), was planted with Eucalypms saligna in 1954 after the original vegetative cover was removed. The eucalypt forest was harvested selectively in 1964 and again in 1971. However, the eucalypt forest was never clear-cut and the native species were allowed to regenerate, largely through coppicing, and developed into a complex native subcanopy. The result was a homogeneous eucalypt upper canopy and a native species subcanopy or "mata suja." Tall exotic grasses covered the ground. Emergent eucalypt trees reached 20 m in height. Mammal Trapping Small mammals were snap trapped in order to obtain voucher specimens, as well as dietary and reproductive information. Such trapping was carried out exclusively in wet meadow and secondary successional habitats. All specimens were either preserved in 10% formalin or made into museum study skins with corresponding complete skulls. I also made study skins of individuals of species of uncertain taxonomic status that were live-trapped in habitats other than those where snap trapping was carried out. In the wet meadow habitat, live trapping started in February 1986 and continued at monthly intervals through January 1987. I used two parallel trapping lines in this habitat. Each line was 280 m in length and subdivided into 20 trap stations separated by 15 m. All traps were placed on the ground, with even numbered stations having only one Sherman live trap, while odd numbered stations had one Sherman and one locally made small wire live trap. Bait was identical to that used in the homogeneous eucalypt forest. In the eucalypt forest with native subcanopy and the native forested habitats, the trapping design was identical. In each area, I cut three parallel lines 300 m in length through the forest; 16 trapping stations were placed along the line separated by 20 m. I used Sherman live traps and locally made small (15 X 15 X 30 cm) and large (25 X 30 X 60 cm) wire live traps. All traps were placed within 3.5 m of the trapping post. All trapping posts had a terrestrial small live trap. Odd numbered trapping posts had a small wire live trap placed in a tree 166 BULLErIN FLORIDA SrATE MUSEUM 34(4) or bush. Mean arboreal live trap height was 1.2 m. Odd numbered trapping posts had a Sherman live trap alternating between arboreal and terrestrial positions. The exterior trapping lines were identical with respect to number, kind, and placement of traps. I did not use the large live traps on the interior line. Aside from the large live traps, the exterior and interior lines were equal in trap number. However, the positions of the Sherman and small live wire traps were reversed for the interior line. The Sherman live traps were introduced into each of the forested habitats after the study was well underway in an attempt to sample smaller bodied species. I placed Shermans in two of the native forested sites in January 1986 and introduced Shermans in the remaining forested sites in May of the same year. I also experimented with traps that were placed in the canopy by a pulley and platform device. This method was similar to that developed by Malcolm (pers. comm.) for use in the Brazilian Amazon. Mean trap height was 11.2 m. I used these arboreal traps to sample the canopy dwelling small mammals. Traps were located at trapping posts along the established lines. I selected trees that were to have arboreal platforms in a subjective manner. I placed traps in trees that I thought had a high degree of canopy connectivity and upper stratum vine density. I spread 42 arboreal platforms across four native forested sites. The primary forest sites, RD/C and RD/T, and one of the mosaic sites, RD/H, each had 12 arboreal traps, 4 traps per line. The other mosaic site, RD/M, had only 6 traps, because I did not believe that there was sufficient upper strata development to support canopy dwelling species. Arboreal canopy trapping started in June 1986 and continued through October 1986. I followed the same schedule in canopy trapping as I had used for terrestrial and arboreal trapping. I used dry oatmeal, pineapple chunks, and cotton balls soaked with cod-liver oil for bait. Traps were set during the day and remained open for five consecutive nights each month for one calendar year. Calculations A first capture was defined as the first occasion that an ihdividual was trapped and marked. The first capture plus subsequent captures of each individual were considered total captures. Minimum known alive (MKA) was the number of individuals actually captured during a particular month whether the capture was the first capture or a recapture from an earlier session. Trapping success of small mammals was calculated in the following manner. The number of traps was multiplied by the number of nights the traps were baited and armed per site per month to determine the number of trap nights. Trapping success was the number of first captures, MKA, or total captures of all species divided by the number of trap nights and expressed in percentages. STALLINGS: BRAZILIAN SMALL MAMMAL INVENTORIES 167 For example, if 100 individuals were trapped during 1000 trap nights, the trapping success would be (100/1000) X 100 = 10%. Recapture indices were calculated by dividing total captures by first captures; thus indicating the average number of times an individual of each species was captured. I recorded the following information from each small mammal captured: date, location On trapping line, position of trap, species, sex, whether a juvenile or adult, its reproductive condition, general condition, external parasitic load on a relative scale, and behavior upon release. I also recorded standard body measurements for each individual: body length, tail, ear, hind foot, and mass. Numbered metal eartags were placed in the left pinna af each individual upon the initial trapping of the individual. In general, there was very little evidence of tag loss. Individuals eartagged from open/field habitat tended to have higher occurrence of tag loss compared to individuals in forested habitats. My taxonomic determinations, when in doubt, were checked by taxonomists specializing in various small mammal groups. Voucher specimens were distributed to the following people: cricetid rodents and marsupials (genus Mamlosa) were sent to Dr. Phil Myers, University of Michigan; rodents of the subgenus Oecomys to Dr. Guy Musser, American Museum of Natural History, and to Dr. Mike Carleton, U.S. National Museum of Natural History. I used body measurements, mass, reproductive condition, and pelage characteristics to determine the age class (juvenile or adult) of each captured individual. An individual was considered an adult if it was reproductively active. Female rodents were considered reproductively active if they (1) had a perforated vulva, (2) were pregnant, or (3) were lactating. Marsupial females were considered reproductively active if they (1) were lactating or (2) had young attached to the teat field. Male rodents were considered reproductively active if the testes were descended. I could not determine the reproductive status of male marsupials as the testes are permanently descended. However, the activity state of the sternal gland in marsupials can indicate the reproductive time of year. Initially, I used the overall condition and relative size of each individual of each sex to assign age classes. Later, I compared my initial classification with the body measurements and mass. Body measurements and weights were sorted for each sex of each species and plotted according to size. I then assigned a body measurement value as the threshold for separating juvenile and adult age classes. These age classes were then compared to the initial age classes that were assigned in the field. I used the General Linear Program (PC-SAS) ANOVA to test for the equivalence of adult body measurements and mass means between sexes for each. species. This analysis enabled me to determine the extent of sexual dimorphism for the external characters. Statistical significance was set at -£ 0.05. Feeding categories were determined by stomach content analysis (Charles- Dominique et al. 1981) and from the literature. I relied heavily on information gleaned from the literature on food preferences of small neotropical mammals. 168 BULLETIN FLORIDA SrATE MUSEUM 34(4) The use of vertical space (i.e. terrestrial, scansorial, or arboreal) of each species was determined from three data sets. For each species, I compared the proportion of captures in trees to that of captures on the ground. There were more terrestrial than arboreal traps, and this bias was corrected by adjusting the number of total trapping opportunities. For this adjustment, I divided the number of arboreal total captures by the total arboreal opportunities (or total arboreal trap nights). The same was done for terrestrial total captures. Results of these two divisions were summed, and each respective result (e.g. arboreal) was divided into the sum. The result generated adjusted percentage success of arboreal and terrestrial captures. The sum of total captures was multiplied by the adjusted percentage in order to generate adjusted number of captures per trapping stratum (on the ground or in arborescent vegetation). These adjustments also were made for each species at each site as well as pooled adjustments across all sites. I tested the null hypothesis that there was no difference in the proportion of arboreal and terrestrial captures for each species across all habitats as well as within each trapping site. Prior to using Student's t-test, I adjusted the proportions by an arc-sin transformation. I then compared the proportion of arboreal and terrestrial responses upon release for each species across all habitat types. These adjustments were made for all species in each habitat type. I used Student's t-tests to test the null hypothesis that there was no difference in the locomotory response upon release of each species. These two data sets were then compared to determine if there were any differences between where an individual was captured and its locomotory behavior upon release across all habitat types. These tests also were performed for each habitat sampled. A species was considered to be arboreal if that species was found to have a high proportion of arboreal captures and a high proportion of arboreal behavior upon release.. The opposite would be true for a terrestrial species. A species would be scansorial if there were no significant differences in the proportion of spatial captures and no significant differences in the proportion of behaviors exhibited upon release. I then compared my results obtained from the trapping data to the available literature for each species. RESULTS Trapping Results Tables 2,3, and 4 present the capture results by species for the three habitat types: native forested, eucalypt with native species subcanopy, and wet meadow habitats, respectively. As a group, marsupials represented 79.2% of the first and 83.3% of the total captures in native forested sites and 67.7% and 82.9% in STALLINGS: BRAZILIAN SMALL MAMMAL INVENTORIES 169 eucalypt forest with native species subcanopy. Rodents represented 97.3% and 98.4% of the first and total captures in the wet meadow habitat. In the native forested habitat, Mannosa cinerea represented over 40% of the marsupial captures (Table 2), while in the eucalypt forest this species represented more than 58% of the marsupial captures (Table 3). Akodon cursor was the major contributor to the rodent captures in all three habitats. This species only represented about 7% of the total captures in the native forested habitat, but 40% of the rodent captures. In the eucalypt forest, A. cursor represented about 16% of the total captures and 93% of the rodent captures (Table 3). This rodent was the dominant species captured in the wet meadow habitat, representing about 85% of both the total and of the rodent captures. In both the native and eucalypt forested habitats, marsupials in general were recaptured at a high rate. Didelphis marmpialis and Mannosa microtarsus both showed low recapture rates and reflect the small sample size. Especially noteworthy was the relatively high recapture rate of Mannosa cinerea (Tables 2 and 3). No individuals of other species, neither rodent nor marsupial, were recaptured as frequently as individuals of this species. In the native forested habitat, individuals of M. cinerea were recaptured on the average 3.1 times, while in the eucalypt forest the average recapture rate for individuals of this species was 7.7 times. Akodon cursor was the only rodent that had a relatively high number of captures and recaptures (Tables 3 and 4). This species had a recapture rate of 1.6 and 3.0 in the eucalypt and wet meadow habitats, respectively. Trapping Success Table 5 presents the trapping success by habitat type. Trapping success was calculated for small mammals in thp native forested habitat. The platform trapping data were excluded. It must be kept in mind that the sampling effort in each general habitat category was different; however, comparisons of trapping success are the result of the number of captures relative to the number of trapping opportunities or nights. Overall, the wet meadow habitat yielded the highest trapping success (18.8%). Figure 4 compares the progression of trapping success in the native forest habitat (without the platform data), the wet meadow habitat, and the eucalypt forest with subcanopy habitat. Although these three habitats have unequal sampling effort and trapping design, they were sampled for a one-year period and show important temporal trends. From the overall gross comparison portrayed in Figure 4 and the percent trapping success presented in Table 5, in 170 BULLETIN FLORIDA STATE MUSEUM 34(4) PERCENT SUCCESS 50 40 30 20 10 0*--#.I-f---==;r==sz~Ezzs~===»-=~,-''Il'--1.'* NOV DEC JAN FEB MAR APR MAY JUN JUL AUG SEP OCT MONTHS EUCALYPT -- WET MEADOW -*-NATIVE FOREST Figure 4. Comparison of the trapping success of small mammals in three general habitat types: eucalypt forest with native species subcanopy, wet meadow habitat, and native forest habitat in the Rio Doce State Forestry Park, Minas Gerais, Brazil. NUMBER OF CAPTURES 140 120 100 80 60 40 20 0 ''''ll,1 --1 11 NOV DEC JAN FEB MAR APR MAY·JUN JUL AUG SEP OCT MONTHS TOTAL CAPTURES + MKA CAPTURES +-FFRST CAPTURES Figure 5. Small mammal capture curves for all native forested trapping sites in Rio Doce State Forestry Park, Minas Gerais, Brazil. Capture curves include total captures, minimum known alive (MKA), and first captures. STALLINGS: BRAZILIAN SMALL MAMMAL INVENTORIES 171 NUMBER OF CAPTURES 25 20 15 10 5 0 NOV DEC JAN FEB MAR APR MAY JUN JUL AUG SEP OCT MONTHS TOTAL CAPTURES -1- MKA CAPTURES -FIRST CAPTURES Figure 6. Small mammal capture curves for eucalypt forest with a native species subcanopy near the Rio Doce State Forestry Park, Minas Gerais, Brazil. Capture curves include total captures, minimum known alive (MKA), and first captures. NUMBER OF CAPTURES 70 60 50 40 30 20 10 O 1, NOV DEC JAN FEB MAR APR MAY JUN JUL AUG SEP OCT MONTHS TOTAL CAPTURES -1 MKA CAPTURES --FIRST CAPTURES Figure 7. Small mammal capture curves for wet meadow habitat in Rio Doce State Forestry Park, Minas Gerais, Brazil. Capture curves include total captures, minimum known alive (MICA), and first captures. 172 BULLEnN FLORIDA SrATE MUSEUM 34(4) contrast to the wet meadow habitat, it appears that the forested habitats, both native and exotic, were quite similar in overall percent success and monthly trapping success. The wet meadow trapping success fluctuated greatly throughout time, from a high approaching 45% in March and April, to a crash lower than 10% in May, October, and November. I plotted the total captures, minimum known alive, and first captures through time for each of the three habitats. Figure 5 presents the capture curves for the native forested habitat. Capture rate was relatively Iow and stable from November through April, with a noticeable increase in June, July, and the early part of August. After the first of August, capture rate dropped back to the levels observed prior to the increase. Figures 6 and 7 compare the capture rate of small mammals in the eucal#t forest and the wet meadow habitat, respectively. The highest number of captures at any one time in the wet meadow habitat was double the highest in the eucalypt forest. However, the trends were similar. Both habitats showed two pronounced peaks in their respective capture curves that corresponded to the same months throughout the year. There was a peak in February, March, and April followed by a crash, and another peak in June, July, and August followed by another crash. Trap T*es Overall, trapping success was higher in terrestrial traps than in arboreal ones (Table 6). Caution should be used in these comparisons as the number of trap nights are unequal; almost three times the number of terrestrial trap nights as the number of arboreal ones. However, I feel that some degree of comparison can be drawn from this analysis based upon the number of captures relative to the number of trap nights. Trapping success by trap type varied considerably (Table 6). Small terrestrial Sherman live traps were the moit successful (9.1%), medium terrestrial traps represented the trap with the greatest number of captures and trapping opportunities, and large terrestrial live traps were relatively unproductive. Arboreal trap type success varied. Small arboreal Shermans were the least productive arboreal trap type (1.8% success). The arboreal platform traps had a success rate of 6.3%. Table 7 reports the number of total captures of each species and the number of captures per trap type. Odd numbered traps represent terrestrial traps, and even numbered ones represent arboreal traps. In general, the number of captures per species in terrestrial and arboreal traps represented the general use of space for each species. For marsupials, Didelphis marsupialis and Metachims nudicaudams were captured principally in terrestrial medium live traps (3~(IT), while Mannosa incana was captured in all STALLINGS: BRAZILIAN SMALL MAMMAL INVENTORIES 173 trap types except for the large terrestrial and arboreal platform traps. Mannosa cinerea was captured in all trap types except the large terrestrial traps. Caluromys philander was trapped principally in arboreal medium live traps and arboreal platform traps. The sample size for rodents was too small to allow for clear trapping trends. Akodon cuisor is the clear exception. The small terrestrial Sherman live trap was most effective for this species. This was mostly a consequence of approximately 80% of all Akodon captures being made in the wet meadow habitat. 0/yzomys subj7avus was trapped principally in terrestrial small Shermans and medium live traps. SPECIES ACCOUNTS The small mammal fauna of the park consisted of 6 species of marsupials and 11 species of rodents. The following accounts present the essential observations regarding the autecology of each species. Additional natural history observations of many of these same species are presented by Fonseca and Kierulff (This volume). The diet, use of vertical space, and habitat requirements are presented in Table 8 for each species captured during the study. SUBCLASS MARSUPIALIA Family Didelphidae Didelphis marsupialis Unne (1158) (Plate lA) The black-eared opossum ranges widely in South America from the Isthmus of Panama to southern Brazil. This species occurs sympatrically with D. albiventris throughout much of its range (Streilein 1982). In the Rio Doce Valley, however, D. manupialis inhabits moister habitats, while D. albivent,is occurs in the cerrado vegetation (Valle and Varejao 1981). A. Gardner (pers. comm.) suggested that the form of D. marsupialis found in eastern Brazil is distinct and should be referred to as D. azame. This species inhabits brush and forested habitats (Alho 1982; Nowak and Paradiso 1983). Miles et al. (1981) found this species to be nocturnal, with a preference for nesting in tree cavities. This species was captured in all forested habitats in the park. Adult body measurements did not indicate sexual dimorphism (Appendix 1). Females have a well developed pouch. This species is terrestrial. There was a significant percentage of terrestrial captures (Table 9) and terrestrial behavior 174 BULLEnN FLORIDA STATE MUSEUM 34(4) upon release (Table 10). Several juvenile individuals and one adult were observed to climb readily. Charles-Dominique (1983) reported that this species exploits the lower stratum in forests, but can climb. It is basically an opportunistic feeder and feeds upon fruit and animal matter (Charles- Dominique 1983). Metachims nudicaudams Geomoy (1803) (Plate lA) The brown four-eyed opossum has a geographical distribution similar to that of D. marsupialis except that it is not found over much of Venezuela nor in northeastern Brazil (Streilein 1982). M. nudicaudatus can be confused with Philander opossum as both have pale spots above the eyes. In addition, there is considerable confusion over the taxonomy of the two species. Nowak and Paradiso (1983) classified this species as Philander nudicau(lams and Philander oposmn: as Memchirops oposmm. I agreed with Honacki et al. (1982) and followed their classification. This species was captured in all forested habitats (Table 11). Metachims nudicaudams is sexually dimorphic in its mass and hind foot measurements (Appendix 1). Females do not have a pouch. This species is strongly terrestrial, rarely caught in arboreal traps (Table 9), and rarely climbs upon release (Table 10). Miles et al. (1981) found this species to be nocturnal and construct nests on the forest floor or in ground hollows. There are very little data on the feeding habits of this species due to the small numbers that have been reported to be trapped. Preliminary data indicate that this species is an insectivore-omnivore (Robinson and Redford 1986) or frugivore-omnivore (Hunsaker 1977). Mannosa incana (Lund 1840) This mouse opossum is endemic to the Brazilian Atlantic Rainforest (Streilein 1982). M. incana occurs in both secondary and primary forest habitat. This species is small in size (adults average weight = 62 g) and strongly sexually dimorphic in body size and color. Males tended to have larger ears and hind feet (Appendix 1) while females tend to have a more rose colored venter and less pronounced face mask (P. Myers, pers. litt.). Females do not have a true pouch. This species tended to use both the ground and arborescent vegetation. I classify the spatial adaptation of this species as scansorial. There was no significant difference in the proportion of terrestrial and arboreal captures (Table 9, p > 0.90, df = 169), however; individuals tended to remain on the ground upon release (Table 10,p < 0.001, df= 145). No data STALLINGS: BRAZILIAN SMALL MAMMAL INVENTORIES 175 exist on the feeding category of this species. Other species of Mannosa which have similar body mass are classified as insectivore-omnivores. Stomach content analysis (n = 3) showed 100% insects from two orders, Coleoptera and Orthoptera (Table 12). I classify this species as an insectivore-omnivore based on the relationship found between body mass and dietary classification (Robinson and Redford 1986). Mannosa cinerea Temminck (1824) (Plate lB) M. cinerea has a disjunct geographical distribution in South America; it occurs in northern Venezuela through the Guianas, and it occurs in the Brazilian Atlantic Rainforest extending into Paraguay (Streilien 1982). M. cinerea occurs in brush and forested habitat, ranging from secondary to primary. This species is a large bodied Mannosa (Appendix 1, average weight = 105 g) and is highly sexually dimorphic based on external body measurements. Males tended to have larger body, tail, ear, and foot (Appendix 1). Females do not have a pouch. This species is strongly arboreal and exploits the high forest stratum (Miles et al. 1981; Charles-Dominique 1983). Miles et al. (1981) found this species to be nocturnal and to construct open arboreal nests rather than use cavities. M. cinerea tended to be caught a greater proportion of the time in arboreal traps (Table 9,p < 0.001, df=356) and tended to exhibit arboreal rather than terrestrial behavior upon release (Table 10, p < 0.001, df=303). This species is primarily an insectivore- omnivore (Robinson and Redford 1986). Mannosa micromrsus Wagner (1842) This species is restricted to the Brazilian Atlantic Rainforest (Streilein 1982). M. microtarsus differs from its congener M. Ggilis by possessing a pure colored white patch of hairs on the throat and chin (Tate 1933). There are insufficient data in the literature to determine the spatial adaptation of this species. I only recorded one capture during the study, in an intermediately disturbed habitat (Table 11). However, the species has a long prehensile tail and short wide feet which suggest an arboreal lifestyle. This species is probably an insectivore-omnivore. 176 BULLETIN FLORIDA STATE MUSEUM 34(4) Caluromys philander Linne (1758) (Plate lB) This species has a disjunct geographical distribution with populations in Venezuela, the Guianas and northern Brazil and in southeastern Brazil (Streilein 1982). The woolly opossum is classified as a forest dwelling species (Nowak and Paradiso 1983). This species was present in the eucalypt, mosaic, and primary forested habitats (Table 10). Based upon the external body measurements, there was no sexual dimorphism in adults (Appendix 1). Females lack a true pouch. According to Charles-Dominique (1983) and Miles et al. (1981), this species exploits the high forest stratum and is nocturnal. My data (based on 51 captures) showed that there was a higher proportion of arboreal captures (Table 9, p < 0.001, df = 49) and that this species tended to climb more than remain on the ground upon release (Table 10, p < 0.05, df= 18). Fruit makes up a large portion of this species' diet (Charles- Dominique 1983; Robinson and Redford 1986). Suborder Eutheria Order Rodentia Oecomys (O,yzomys) trinitatus = (0. concolor) Wagner (1845) This genus is in need of revision and the subgenus Oecomys is currently being revised (P. Myers, pers. comm.). This species was previously called Olzon:ys concolor and was known, within Brazil, as an Amazonian species (Alho 1982). However, Nitikman and Mares (1987) reported trapping this species in gallery forest in the Brazilian cerrado. I captured this species in all native forested habitats (Table 11). The species is not sexually dimorphic (Appendix 1). Gyldenstolpe (1932) and Moojen (1952) stated that this species is "more or less adapted for arboreal life." My data Suggested that this rat is scansorial; there were no significant differences in the proportion of terrestrial and arboreal captures (Table 9,p > 0.10, df= 19) and no significant differences in terrestrial and arboreal behavior upon release (Table 10, p > 0.20, df= 9). Most species of the genus Oozomys are frugivore-granivores (Robinson and Redford 1986). SI'ALLINGS: BRAZILIAN SMALL MAMMAL INVENTORIES 177 Olzomys subfavus Wagner (1842) (Plate 2A) This species is distributed throughout the Guianas, southeastern Brazil, and eastern Paraguay (Honacki et al. 1982; Alho 1982). In Brazil, it occurs in the cerrado, caatinga, and Atlantic Rainforest (Alho 1982). This species was captured in wet meadow and heavily disturbed secondary habitat (Table 11). There were no differences in body measurements between sexes in adult individuals (Appendix 1). I classify this species as terrestrial. This species tended to be captured more on the ground than in the trees (Table 9, p < 0.01, df=46) and was never observed to climb upon release (Table 10). Stomach content analysis (n= 1) showed 95% grass and 5% fruit (Table 12). Ogzomys capito Olfers (1818) This species has a wide distribution throughout the neotropics and occurs in a variety of habitats ranging from agricultural fields (Moojen 1952) to humid forests (Alho 1982). Oiyzomys capito was primarily captured in humid forests ranging from intermediate levels of disturbance to primary forests in the park (Table 11). There were no significant differences in body measurements between sexes for adults (Appendix 1). My capture and release data are in accordance with Alho's (1982) terrestrial classification for this species. 0. Capito tended to be caught more on the ground than in trees (Table 9, p < 0.05, df= 19) and tended to remain on the ground upon release (Table 10, p < 0.001, df= 11). 00,zomys (Oligoiyzomys) nignpes (eliums) Wagner (1845) This small-bodied rodent (Appendix 1) occurs in grassland, wet meadow, and secondary forest habitat in northern Argentina, eastern Paraguay, southern Brazil, and the Bolivian Beni (Honacki et al. 1982).In the Park, all captures were made in the wet meadow habitat (Table 11). All captures were made on the ground (n = 4), however; the individuals climbed readily in captivity (pers. obs.). The results from the stomach analysis (n=5) revealed a wide range of foodstuffs (Table 12). 178 BULLETIN FLORIDA STATE MUSEUM 34(4) Abrawayaomys ruschii Cunha and Cruz (1979) (Plate 2A) This species is only known from the type locality in Espirito Santo, eastern Brazil. It is endemic to the Brazilian Atlantic Rainforest. The single capture of this species was *recorded in an intermediately disturbed forest (Table 11). There is very little information available regarding the ecology of this species, and there are only three study skins found in museums (A. Gardner, pers. comm.). Rh*idomys mastacalis Lund (1840) (Plate 2B) Climbing mice range south from Margarita and Tobago islands to Venezuela and Guianas to northeastern and east central Brazil (Honacki et al. 1982). This species was only captured in a relatively undisturbed primary forest (Table 11). Sample size was too small to detect any differences between terrestrial and arboreal captures and behavior upon release. However, as the common name implies, this species climbs readily. I captured two individuals in my house in the park, a commonly cited "exotic" habitat for this species (Nowak and Paradiso 1983). Nectomys squamipes Brants (1827) The neotropical water rat occurs in aquatic habitats either in grasslands and wet meadows or in forests. This species' distribution ranges from the Guianas to Colombia to Peru and in Brazil, Paraguay, and northeastern Argentina (Honacki et al. 1982). It tended to be caught more on the ground (Table 9, p < 0.05, df= 17) and exhibited a significant tendency to remain on the ground upon release (Table 10, p < 0.001, df= 10). Stomach content analysis (n=2) showed 50% grass and stems and 50% fruit (Table 12). Akodon cursor Winge (1887) (Plate 2B) This species occurs in several habitat types from southeastern and central Brazil to Uruguay, Paraguay, and northern Argentina. The wet meadow STALLINGS: BRAZILIAN SMALL MAMMAL INVENTORIES 179 habitat in the park was the primary habitat to capture this species (Table 11). A. cursor was formerly included in A. an'iculoides (Honacki et al. 1982). This species is sexually dimorphic in tail (p < 0.01) and body (p < 0.008) length (Appendix 1). A. cursor is strongly terrestrial (Table 9 and Table 10). Analysis of stomach contents (n = 23) revealed a high proportion of insects, seeds and fruit (Table 12). Calomys laucha Olfers (1818) This species occurs in grassland and wet meadows in southern Bolivia, southeastern Brazil, Paraguay, central Argentina, and Uruguay. This species was only captured in the wet meadow habitat in the Park (Table 11). The results from one stomach sample revealed 100% seeds (Table 12). Oxymycterus roberti Thomas (1901) The burrowing mouse occurs in a variety of habitats but is usually associated with moist substrate in open or brush habitats. This species was captured in both wet meadow and secondary forest habitats in the Park (Table 11). This species is endemic to eastern Brazil. This semifossorial mouse is described as an insectivore (Nowak and Paradisio 1983). Stomach analysis (n = 2) revealed 100% insects (Table 12). Family Caviidae Cavia fulgida Wagler (1831) This species of cavy is endemic to the open grasslands and wet meadows of the Atlantic Rainforest of eastern Brazil (Honacki et al. 1982; Nowak and Paradiso 1983). I captured this species in grassland and wet meadow habitats in the Park, however; it was not trapped in site RD/B. For this reason this species does not appear in Table 11. Cavies are terrestrial and are herbivore- grazers (Nowak and Paradiso 1983). 180 BULLETIN FLORIDA SrATE MUSEUM 34(4) Family Echimyidae Euryzygomatomys spinosus Fischer (1814) The single species of guiara is endemic to southeastern Brazil, Paraguay, and northeastern Argentina (Honacki et al. 1982). I captured this species in the wet meadow habitat (Table 11). This species inhabits open grasslands and wet meadows, is considered terrestrial or semifossorial (Alho 1982), and is most probably a herbivore-grazer. DISCUSSION The trapping success realized in this study for neotropical humid forests falls within the range of observed success rates (Table 13). However, one striking difference between this and other neotropical small mammal studies was the high number of marsupial captures relative to rodent captures. All reported studies show rodent biases and usually high captures of rodents relative to marsupials. Only Emmons (1984) reported a marsupial to rodent capture ratio approaching equality. Trapping results from small mammal studies in southeastern Brazil varied considerably with respect to marsupial and rodent biased captures. Only Fonseca (pers. comm.) has reported marsupial biased trapping results from a variety of native forest sites in eastern Minas Gerais. Avila-Pires (1978) captured 245 rodents and 40 marsupials from the Rio Doce Park. Gastal (1982) reported that five species of marsupials were captured in the Park but gave no comparative data for rodents. Dias (1982) trapped more rodents than marsupials in the Rio Doce Valley in Minas Gerais. Davis (1945) trapped 58 didelphid marsupials and 285 rodents in his study site in Rio de Janeiro. Hunsaker (1977) stated that marsupials require considerable effort to trap. Perhaps one explanation for the observed high marsupial capture rate could be due to the habitat type found in the Park. Charles-Dominique (1983) suggested that didelphid marsupials can reach high local densities in areas of abundant food resources. He postulated that these species are r-strategists and are adapted to the "unstable environment of secondary forests." There is very little primary habitat in the Park relative to secondary habitat. Most of the forest habitat in the Park has been altered by fire in the recent past. The primary forest plots that I sampled yielded the lowest species richness and absolute captures of didelphid marsupials relative to the other secondary forested habitats (Stallings 1988). In Panama, Didelphis marsupialis tended to occur at higher densities in primary forest, while Calummys and Philander were found at higher densities in secondary forest than primary forest (Fleming 1972). STALLINGS: BRAZILIAN SMALL MAMMAL INVENrORIES 181 Marsupials were recaptured with relative high frequency in this study, especially Metachirus nudicaudatus, Marmosa incana, and Marmosa cinerea. My recapture data on marsupials agreed with data reported by Fleming (1972, 1973), August (1984), and to some degree with that found by O'Connell (1979). These data were not in accord with Hunsaker (1977), who stated that didelphid marsupials are difficult to recapture. For example, individuals of Mannosa cinerea were captured on the average 7.7 times in the eucalypt forest with native subcanopy habitat. This high recapture rate could be an artifact of the habitat. This trapping site was surrounded by habitat unsuitable for arboreal species. In essence, this habitat was a forested island. On one side there was a monoculture of Eucalyptus saligna with no subcanopy, on another a marsh conyerted to a rice field, and the other two sides of the forest were bounded by pasture. Also, the subcanopy offered more food and resting opportunities than did the homogeneous eucalypt canopy. Perhaps the inability to move far beyond the limits of the forest and the location of the optimal habitat within the forest help explain the observed recapture rate. Another obvious difference between this study and other inventories conducted in neotropical native forests was the absence of echimyid rodents. Species of the genus Proechimys ate ~e most widespread taxa of the family Echymidae in the neotropics (Hershkovitz 1969). These forest species are terrestrial and usually appear on species lists from forest inventories. In Panama, Proechimys was a common forest capture (Eisenberg and Thorington 1973; Glanz 1982). Handley (1976) reported Proechimys as a common species in Venezuela. Emmons (1984) and Terborgh et al. (1986) reported captures of Proechimys from forested sites in Peru and Ecuador. There are several reports of Proechimys captures in Brazilian tropical moist forests. Laemmert et al. (1946), Emmons (1984), Carvalho (1965), Miles et al. (1981) and Malcolm (pers. comm.) reported Proechimys in their inventories from the Brazilian Amazon. In the Atlantic Forest, Davis (1945) and Fonseca (1988) reported two species of Proechimys from their studies in the states of Rio de Janeiro and Minas Gerais, respectively. Dias (1982) trapped one species of Proechimys in three study areas in the Rio Doce Valley. I did not capture one individual of Proechimys from the Park in approximately 35,000 trap nights from 1985 to 1986, nor from an additional 30,000 trap nights from 1986 to 1987 (unpubl. data). One explanation could be the presence of predators in the Park. Eisenberg (1980) speculated that the abundance of rodents in some neotropical sites and the paucity of rodents in other sites could be the result of the absence or presence of top predators, respectively. Hershkovitz (1969) stated that species of the genus Proechimys "are the basic source of protein for lowland predators in the Brazilian subregion." The felid community in the Park is intact. All the felids have been observed by field workers in the recent past. I saw spoor from jaguar, puma, ocelot, Geoffrey's cat, and jaguarundi. 182 BULLETIN FLORIDA SrATE MUSEUM 34(4) The results from the wet meadow habitat were consistent with the literature (Table 13). In every study, there were more rodent captures than marsupial captures. In this study, Akodon cursor was the dominant species in terms of absolute numbers and captures. O'Connell (1981) reported that Zygodontomys, an ecological equivalent of Akodon, was the dominant rodent in grass habitat in Venezuela and represented 85% of the total rodent captures. However, one difference between this study and others was the observed high trapping success. As can be observed from Table 10, Akodon cursor made up 85% of the captures and was largely responsible for the high trapping success. One explanation for the high capture rate could be due to the shape and amount of the available grass or wet meadow habitat in the Park. This habitat type occurs particularly at the bases of hills dnd on the perimeter of the swamps and lake edges. There are no vast areas of this habitat type in the park, and these areas commonly take on a long and narrow shape following the contour lines. There was a high degree of habitat specificity exhibited by the species captured in this habitat. Grassland rodents can achieve high local densities, and perhaps this fact, coupled with the populations being compressed and compacted in narrow and small habitat, helps to explain the observed high trapping success in the wet meadow habitat. The eucalypt forest with native subcanopy habitat yielded surprising results. I did not expect to find many small mammals in this habitat because of "plantation effects." However, seven species of small mammals were captured, five of which were marsupials. In fact, the marsupial/rodent capture ratio was similar to that observed in the native forested habitat (Table 14). Dietz et al. (1975) captured two species of terrestrial rodents, Ofomys nignpes and Akodon cursor, in homogeneous eucalypt forest with grass/bamboo undergrowth. They reported a total of five species, only one of which was not strictly terrestrial, in their two native forested habitats. The plantation habitats in Dietz et al. (1975) and in this study are similar in that both treated eucalypt plantations of similar age and that in both the terrestrial substrate was covered by grass. The major difference was the native species subcanopy in this study. I captured a relatively high number of terrestrial rodents and marsupials and a high number of arboreal marsupials, but no arboreal rodents. The native species subcanopy could be considered a secondaty forest sere, if the emergent eucalypt stratum was ignored. Charles-Dominique's (1983) hypothesis that didelphid marsupial abundance increases in secondary habitat would explain the high numbers of marsupials captured in this stiidy. The temporal capture results from the native Torested habitat suggest that seasonality is important with respect to the trappability of small mammals. The trapping data show a pronounced peak in the total number of captures, MKA, and first captures for the native forested habitat during the cool, dry winter. Davis (1945) reported a similar trend and suggested that this was the result of more younger individuals present in the trapping pool or because of a paucity of natural food items during this time of year. My data do not support STALLINGS: BRAZILIAN SMALL MAMMAL INVENTORIES 183 the hypothesis that more younger individuals explain the pronounced increase; rather it appears that food resource paucity results in the increase (unpubl. data). The eucalypt and wet meadow habitats both produced peaks in the cool, dry winter and again in the late summer. These two habitats might have yielded similar capture curves because they share a grass substrate. Akodon cursor was an important component of both habitats and is an insectivore/omnivore that is reported to use a high proportion of grass and grass seed in its diet (Nowak and Paradiso 1983). Mannosa incana and Metachims nudicaudatus are insectivore/omnivores and frugivore/omnivores, respectively, and perhaps track insect availability in grass substrate. The grass species did not produce seeds until late May. Thus, perhaps the peak observed in February, March and April can be explained by the lack of food for both rodents and marsupials. The second peak, which occurred in June, July, and August, could also be explained in terms of a decrease in food availability. Insect and fruit availability are usually low duiing the hibernal period in seasonal neotropical forests (e.g. Janzen and Schonener 1968). The marsupial species rely heavily on these food resources. The results of a preliminary stomach content analysis on Akodon, suggest that insects are important items in this species diet (unpubl. data). Graminoids in this habitat wefe dry, and seeds were not as readily available as they were during April and May. Data analysis of trap type revealed that terrestrial small Sherman live traps were very productive in the wet meadow habitat but yielded relatively few captures in the forested habitats. Arboreal small Shermans were relatively unproductive in the forested habitats. Large terrestrial live traps were unproductive in the forested habitats. The most productive trap types in the forested habitats were the medium sized terrestrial and arboreal traps and the arboreal platform traps. Some individuals of species that are considered terrestrial were captured in arboreal traps. Apparently these instances resulted when low arboreal traps were easily accessible from the ground by either a vine or log. No additional species were added to the inventory list by using the arboreal platform traps. However, these traps increased the capture frequency for the highly arboreal marsupial Caluromys philander. In total, I recorded 49 captures for this species in both the eucalypt with native species subcanopy and the native forested habitats. In the latter habitat, however, Calitromys was captured only five times in the terrestrial and low arboreal traps, as compared with 29 times in the arboreal platform traps. Clearly, the abundance of this species would have been underestimated if arboreal traps had not been used. Malcolm (per. comm.) obtained similar results with platform traps in Manaus. Mam,osa cinerea was also trapped with relatively high frequency in this trap type. The use of arboreal platforms for trapping small mammals was first described by Davis (1945) and Laemmert et al. (1946). Unfortunately, the relative success of arboreal traps could not be determined from these studies. 184 BULLETIN FLORIDA STATE MUSEUM 34(4) It was surprising to find such a high frequency of Caluromys in the eucalypt forest. Although this arboreal frugivore is quite common in native forests, it was not expected in an exotic monoculture plantation. Presumably its principal food resources came from the native species subcanopy. This interpretation gains support from the fact that this species also was captured in terrestrial and low arboreal traps, suggesting that this species regularly moved down through the subcanopy. The results of this small mammal inventory give substance to previous impressions of a highly diverse and endemic Atlantic Forest fauna. These results also demonstrate the important role that didelphid marsupials play in the community structure of small mammals in one of the largest remaining tracts of native Atlantic Forest in Brazil. Wet meadow habitat in this region supports many rodents, although Akodon cursor dominates. Eucalypt forests with native species subcanopy can play an important role in conserving small mammal communities in a region greatly altered by monocultural plantations. LITERATURE CITED Alho, C. J. & 1982. Brazilian rodents: Their habitats and habits. Pp. 143-165 in MA. Mares and H.H. Genoways (eds.). Mammalian Biology in South America. Spec. Publ. Ser., Pymatuning Lab. Ecol., Univ. Pittsburgh, Pennsylvania. August, P. V. 1984. Population ecology of small mammals in the 11anos of Venezuela. Pp. 71- 104 in R.E. Martin and B.R. Chapman (eds.). Contributions in Mammalogy in Honor of R6bert L. Packard. Spec. Publ. Mus. Texas Tech Univ., Lubbock, Texas. Avila-Pires, F. D. 1978. Fauna de Mamfferos PFERD. Relat6rio parcial das pesquisas ecoldgicas no Parque Florestal Estadual Rio Doce. CErEC, December. , and E. Gouvea. 1977. Mamiferos do Parque Nacional do Itatiata. Bol. Mus. Nac. Rio de Janeiro, 291:1-29. Borchert, M., and R. L. Hanson. 1983. Effects of flooding and wildfire on valley side wet campo rodents in central Brazil. Rev. Brasil. Biol. 43(3):229-240. Botelho, J. R., and P. M. Linardi. 1980. Alguns ectoparasitos de roedores silvestres do municfpio de Caratinga, Minas Gerais, Brazil. I. Relacoes pulga hospedeiro. Rev. Bras. Ent. 24(2):127-130. Cabrera, A 1957. Cat~logo de los mamfferos de America de Sun I. Rev. Mus. Cien. Nat. "Bernadino Rivadavia." IV:1-309. 1961. Catdlogo de los mamfferos de America de Sun II. Rev. Mus. Cien. Nat. "Bernadine Rivadavia." IV:310-732. Carvalho, C. T. 1965. Bionomia de pequenos mamfferos em Boraceia. Rev. Biol. Trop. 13(2):239-257. CETEC. 1981. Vegetagao do Parque Florestal do Rio Doce. Programa de Pesquisas Ecoldgicas no Parque Estadual do Rio Doce, Relatorio Final, 2nd Vol., 277 pp. Fundaqao Centro Tecnoldgico de Minas Gerais, Belo Horizonte, Brazil. Charles-Dominique, P. 1983. Ecology and social adaptions in didelphid marsupials: Comparison with eutherians of similar ecology. Pp. 395-420 in J.F. Eisenberg and D.G. Kleiman (eds.). Advances in the Study of Mammalian Behavior. Spec. Publ. No. 7, Amen Soc. Mamm., Shippensburg, Pennsylvania. , M. Atramentowicz, M. Charles-Dominique, H. Gerard, A. Hladik, C.M. Hladik, and M.F. Prevost. 1981. Les mammiferes frugivores arboricoles nocturnes d'une foret guyannaise: Inter-relations plantes-animaux. Rev. Ecol. 35:341-435. STALLINGS: BRAZILIAN SMALL MAMMAL INVENTORIES 185 Davis, D. E. 1945. The annual cycle of plants, mosquitos, birds and mammals in two Brazilian forests. Ecol. Monog. 15:244-295. Dias, M. 1982. Leishmaniose tegumentar Americana na zona do Rio Doce, Minas Gerais. Aspectos da doenga no homem e estudo de reservatorios. Ph. D. dissertation, Universidade Federal de Minas Gerais, Brazil. 88 pp. Dietz, J. M., E. A. Couto, A. C. Alfenas, A. Faccini, and G. F. da Silva. 1975. Efeitos de duas planta~oes de florestas homog6neas sobre populagoes de mamfferos pequenos. Brasil Florestal 6(23):54-57. Eisenberg, J. F. 1980. The density and biomass of tropical mammals. Pp. 35-56 in M. Soule and B. Wilcox (eds.). Conservation Biology: An Evolutionary Perspective. Sinauer Press, Sunderland, Massachusetts. , and R. W. Thorington. 1973. A preliminary analysis of a neotropical fauna. Biotropica 5(3):150-161. Emmons, L H. 1984. Geographic variation in densities and diversities of non-flying mammals in Amazonia. Biotropica 16(3):210-222. Fleming, T. H. 1972. Aspects of the population dynamics of three species of opossums in the Panama canal zone. 1 Mamm. 53:619-623. . 1973. The reproductive cycles of three species of opossums and other mammals in the Panama canal zone. J. Mamm. 54:439455. Fonseca, G. A. B. 1988. Patterns of species diversity of small mammals in the Brazilian Atlantic Forest. Ph. D. dissertation, Univ. Florida, Gainesville, , and K H. Redford. 1985. The mammals of IBGE's ecological reserve and an analysis of the role of gallery forests in increasing diversity. Rev. Brasil. Biol. 44:517-523. , and M. A. M. Kierulff. 1988. Biology and natural history of Brazilian Atlantic Forest small mammals. Bull. Florida State Mus., Biol. Sci. 34(3):99-152. Gastal, M. L. A. 1982. Nota previa sobre os marsupiais do Parque Estadual do Rio Doce, Minas Gerais. IX Congreso Brasileiro de Zoologia, pp. 30-31. Gilhaus, J. P. 1986. Vegetation survey of the Parque Florestal Estadual do Rio Doce-MG-Brazil. M. S. thesis, Federal University of Vicosa, Brazil, and Agricultural University of Wageningen, The Netherlands. 86 pp. Glanz, W. E. 1982. The terrestrial mammal fauna of Barro Colorado Island: Censuses and long- term changes. Pp. 455468 in E.G. Leigh, Jr., A.S. Rand, and D.M. Windsor (eds.). The Ecology of a Tropical Forest: Seasonal Rhythms and Long-Term Changes. Smithsonian Inst. Press, Washington, D.C. Gyldenstolpe, N. 1932. A manual of Neotropical sigmodont rodents. Kungl. Sv. Vet. Akademiens Handlingar. Band 11. 3:1-164. Haffer, J. 1974. Avian Speciation in Tropical South America. Nuttall Omithol. Club, Cambridge, Massachusetts. 390 pp. Handley, C. 0. Jr. 1976. Mammals of the Smithsonian Venezuela project. Brigham Young Univ. Sci. Bull. 20:1-91. Hershkovitz, P. 1969. The evolution of mammals on southern continents. VI. The recent mammals of the Neotropical region: A zoogeographic and ecological review. Quar. Rev. Bio. 44(1):1-70. Honacki, J. H., K E. Kinman, and J. W. Koeppl. 1982. Mammal Species of the World: A Taxonomic and Geographic Reference. Allen Press, Inc., and the Assoc. System, Lawrence, Kansas. Hunsaker, D. 1977. Ecology of new world marsupials. Pp. 95-158 in D. Hunsaker (ed.). The Biology of Marsupials. Academic Press, New York. Janzen, D. H., and T. W. Schoener. 1968. Differences in insect abundance and diversity between wetter and drier sites during a tropical dry season. Ecology 49:96-110. Kinzey, W. G. 1982. Distribution of primates and forest refuges. Pp. 455482 in G.T. Prance (ed.). Biological Diversification in the Tropics. Columbia Univ. Press, New York. Lacher, T. E. 1981. The comparative social behavior of Kerodon rupesfris and Galea spirii and the evolution of behavior in the Caviidae. Bull. Carnegie Mus. Nat. Hist. 17:1-71. , and C. J. R. Atho. In press. Small mammal communities in the Brazilian pantanal: Population densities, microhabitat affinities, and species interactions. J. Mamm. Laemment, H. W., L. de Castro Ferreira, and R M. Taylor. 1946. Investigation of vertebrate hosts and arthropod vectors. Amen 1 Trop. Med. 26(suppl):23-69. 186 BULLEnN FLORIDA SrATE MUSEUM 34(4) Ispes, A. D. 1982. Proposta de piano diretor do Parque Estadual Rio Doce. Instituto Estadual de Florestas, M. G., Brazil. Mares, M. A., M. A Willig, K E Streilein, and T. E. Lacher. 1981. The mammals of northeastern Brazil: A preliminary assessment. Ann. Carnegie Mus. 50:80-137. Miles, M. A., A. A. de Souza, and M. M. Povoa. 1981. Mammal tracking and nest location in Brazilian forest with an improved spool and line device. J. Zool. London. 195:331-347. Mittermeier, R. A., A. F. Coimbra-Filho, I. D. Constable, A. B. Rylands, and C. Valle. 1982. Conservation of primates in the Atlantic forests of Brazil. Int. Zoo. Yearbook. Moojen, J. 1952. Os Rdtdores do Brasil. Instituto Nacional do Livro, Rio de Janeiro. Mori, S. A., B. M., Boom, and G. T. Prance. 1981. Distribution patterns and conservation of eastern Brazilian coastal forest tree species. Brittonia 33(2):233-245. Miiller, P. 1973. The dispersal centers of terrestrial vertebrates in the Neotropical realm. W. Junk Publishers, The Hague, Netherlands. 244 pp. Nitikman, L., and M. Mares. 1987. Ecology of small mammals in a gallery forest of central Brazil. Ann. Carnegie Mus. 56(2):75-95. Nowak, R M., and J. L, Paradiso. 1983. Walker's Mammals of the World. 4th Ed. Johns Hopkins Univ. Press, Baltimore, Maryland. 1362 pp. O'Connell, M. A. 1979. The ecology of didelphid marsupials in northern Venezuela. Pp. 73- 87 in J.F. Eisenberg (ed.). Vertebrate Ecology in the Northern Neotropics. Smithsonian Inst. Press, Washington, D.C. - 1981. Population biology of North and South American grassland rodents: A comparative review. Pp. 167-185 in M. M. Mares and H. H. Genoways (eds.). Mammalian Biology in South America. Spec. Publ. Ser., Pymatuning Lab. Ecol. Univ. Pittsburgh, Pennsylvania. Peterson, N. E. In press. Short term succession of mammal species after clearing and burning the Amazon forest. Rev. Bras. Pesq. Med. Biol. Redford, K H., and G. A. B. Fonseca. 1986. The role of gallery forests in the zoogeography of the Cerrado's non-volant mammalian fauna. Biotropica 18(2):126-135. Robinson, J. G., and K H. Redford, 1986. Body size, diet, and population density of Neotropical forest mammals. Amen Nat. 128(5):665-680. / Saijo, Y., and J. G. Tundisi (eds.). 1985. Limnological studies in centiat Brazil, Rio Doce valley lakes and Pantanal wetland. 1st Report. Lab. Chem. Biol.fWater Research Inst., Nagoya Univ., Nagoya, Japan. Silva-Neto, H. F. 1984. Infuencia dos incendios florestais no aparecimento de embaubas (Cercropia holoteuca) na floresta do Parque Florestal Estadual do Rio Doce. Monograph presented to the Department of Forest Engineering, Centro de Ciencias Agrarias, Universidade Federal de Vicosa, Vicosa, Brasil. 15 pp. Stallings, J. R 1988. Small mammal communities in an eastern Brazilian park. Ph. D. dissertation, Univ. Florida, Gainesville. Streikin, K. E. 1982. Behavior, ecology and distribution of the South American marsupials. Pp. 231-250 in MA. Mares and H. H. Genoways (eds.). Mammalian Biology in South America. Spec. Publ. Ser. Pymantuning Lab. Ecol., Univ. Pittsburgh, Pennsylvania. Tate, G. H. H. 1933. A systematic revision of the marsupial genus Marmosa. Bull. Amer. Mus. Nat. Hist. 66(1):1-246. Terborgh, J. W., J. W. Fitzpatrick, and L. Emmons. 1986. Annotated checklist of bird and mammal species of Cocha Cashu Biological Station, Manu National Park. Fieldiana 21:1-29. Valle, C. M. de C., and J. B. M. Varejao. 1981. Nota previa sobre o padrao de dispersao das especies do genero Didelphis (Marsupialia) no estado de Minas Gerais (Brasil)· Resumos do VIII Congr. Bras. Zool. Brasilia, DF: 199. Vieira, C. 1955. Lista remissiva dos mamiferos do Brasil. Arq. Zool. de Sao Paulo. 8(2):341- 464. STALLINGS: BRAZILIAN SMALL MAMMAL INVENTORIES 187 Table 1. Forested and open/field habitats in the Rio Doce State Forestry Park, Minas Gerais, Brazil. Habitats follow Gilhaus (1986). Habitat Type % Total FORESTED HABITATS Tall Primary Forest with Epiphytes 8.4 Tall Forest 30.0 Medium to tall Forest with Bamboos and Graminoids 30.6 Medium Secondary Foreit with Bamboos and Graminoids 17.2 Low Secondaty Forest 0.1 OPEN/FIELD HABITATS Low Woodland 1.1 Low Tree and Scrub Tallgrass Savanna 0.6 Tallfem Field 0.1 Evergreen Tallgrass Field with 1*ha sp. 3.0 Partially Submerged Shortherb held and Aquatic Habitat 8.9 T66~6 Table 2. Capture results from native forested plots in Rio Doce State Forestry Park, Minas Gerais, Brazil. RECAP INDEX = total captures/first captures, and represents the average number of times that an individual of species X is captured. Numbers in parentheses represent percent of contribution of capture per species per taxonomic group. Total First Recap Species Captures % Total Captures % Total Index MARSUPIALS Didelphis marsupialis 35 F3-~.~ 32 7.8 ( 9.9) 1.1 Metachirus nudicaudatus 140 18.0 (11 .6 91 22.3 (28. 11) 13 Marmosa incana 154 19 .8 (23.8 90 22 .1 (27.81 1 .7 Marmosa cinerea 283 36.4 (43. 92 22.5 (28 .4) 3.1 Marmosa microtarsus 1 0. 1 0.2 1 0.2 ( 0.31 1 .0 Caluromys philander 34 0.4 5 .3 18 _-4.4£M) 1 .9 777 RODENTS Nectomys squamipes 15 1.9 (113 9 2.2 (10. 1.7 Rilidpidomys mastacalis 7 0 . 9 ( 5 .4 3 0.7 ~16 2 . 3Akodon cursor 52 6.7 (40.0 27 6.6 ' 2.1 1.9 Oecomys #initans 21 2.7 (13.8 19 4.7 (22.6 1.1 Ogzomys capito 18 2.3 (13.8 15 3.7 (17.9 1.2 Oozomys sutflavus 13 1.7 (10.0 7 1.7 8.3 1.9 Oxyn,ycterus roberti 3 0.4 2.3 3 0.7 3.6 1.0 Abrawayomys ruschii 1 0. 1 0.8 1 0.2 1 .2 1 .0 T36 EL 188 BULLEI'IN FLORIDA STATE MUSEUM 34(4) Table 3. Capture results from eucalypt forest with native forest subcanopy. RECAP INDEX= total captures/first captures and represents the average number of times that an individual of species X is captured. Numbers in parentheses represent percent of contribution of capture per species per taxonomic group. Total First Recap Species Captures 9 Total Captures % Total Index MARSUPIALS Didelphis marsupialis 7 4.4 (58) 3 5 .6 ( 81) 2.3 Metachirus nudicaudatus 18 11 .4 (13.'71 9 16.7 05:0) 2.0 Marmosa incana 14 8.7 (10.71 8 14.8 (222) 1.8 Marmosa cinerea 77 48.7 88.81 10 185 (27.8) 7.7 Caluromys philander 15 9.6 (11 .51 6 11 .1 (16 .71 25 ~[3T -EZE[m~I --36 -66~Mbl RODENTS Akodon cursor 25 15 .8 (92 .6) 16 29.6 (88.9) 1 .6 Oryzomys capito 2 1 .4 ( 7.4) 2 -3.7 (11 ·1) 1 .0 77 773fTW~ .L. . Table 4. Capture results from wet meadow site in Rio Doce State Forestry Park, Minas Gerais, Brazil. RECAP INDEX= total captures/first captures and represents the average number of times an individual of species X was captured. Numbers in parentheses represent percent contribution of captures per species per taxonomic group. Total First Recap Species Captures % Total Captures % Total Index MARSUPIALS Mannosa incana 3 0.8 (50.01 2 1.3 (50.0) 1.5 Marmosa cinerea 1 0.3 (16.n 1 0.7 (25 .03 1 .0 Caluromys philander 2 03 (33.3) 1 0.7 (25 .0) 2.0 1.6(100.0) -1 -HmmRODENTS Necton:ys squamipes 5 1.3(1.4 4 2.7(2.7 1.3 Akodon cursor 315 843 (85 .8 105 70.0 (71 .9' 3.0 On)zomys capito 1 0.3 0.3 1 0.7 ( 0.7 1 .0 Ogzon,ys sub/lavus 31 8.3 8.4 22 14.7 (15.1' 1.4 Oryzomys nigripes 4 1 .0 1 .1 4 2.7 ( 2.7 1 .0 Calomys kiucha 4 1 .0 1 . 1 4 2.7 < 2 .7 1 .0 Oxymycterus roberti 4 1 .0 1 . 1 4 2.7 ( 2 .7 1 .0 Euryzygomatomys spinosus 2 03 03 2 1 .3 < 1 .4< 1 .0 373 -136 -9738Wl- STALLINGS: BRAZILIAN SMALL MAMMAL INVENTORIES 189 Table 5. Trapping success of small mammals calculated bs, habitat category in Rio Doce State Forestry Park, Minas Gerais, Brazil. Habitat Number of Number of Category Trap Nights Captures % Success Native forest (excluding platforms) 30,960 710 2.3 Native forest, platforms only 1,050 66 6.3 Wet meadow 1,980 373 18.8 Eucalxpt forest w/native species Euca~~cta~yfs~t w/no subcanopy 6,000 158 2,6 500 1 0.0 TOTALS 40,490 1,308 Table 6. Trapping success by trap type for all species in all habitat types. Trap types are arranged according to trapping location: terrestnal or arboreal. Trap types are as iollows: 1Sr= small terrestrial Sherman live trap; 3MT= medium sized terrestrial live trap; 5LT= large terrestrial live trap; 2SA= small arboreal Sherman live trap; 4MA= medium sized arboreal live trap; 6PA= arboreal platform trap. No. No. Percent Trap Type Captures Trap Nights Success TERRESTRIAL 1ST 370 4,080 9.1 3MT 553 18,000 3.1 SLT 42 3~'~ 0.7 , 3.2 ARBOREAL 2SA 48 2,640 1.8 4MA 228 8,640 2.6 6PA 66 1.050 6.3 --m 12,330 2.8 190 BULLETIN FLORIDA SrATE MUSEUM 34(4) Table 7. Trap response by species across all habitat types. Trap types are explained.in Table 6. Trap Types Species Total 1ST JA 3MT 4MA 5LT 6AP Didelphis marsupialis 42 0 0 24 2 15 1Metachints.nudicaudatus 158 0 0 129 3 26 0 Marmosa incana 171 13 14 106 38 0 0Nfarmosa cinerea 361 2 29 145 150 0 35 Caluromys philander 51 0 0 9 13 0 29 Marmosa microtarsus 1000100Nectomys squamipes 19 4 0 13 1 1 0Ripidomys mastacalis 7003301 Alcotton cursor 392 312 2 78 0 0 0Oecomys vinitatis 21 1 3 9 8 0 0Oryzomys capito 21 8 0 11 2 0 0Oxymycterus roberti 10 7 0 3 0 0 0 Abrawayaomys nischii 1000100Oryzomys subflavus 44 18 0 20 6 0 0Calomys laucha 2 2 0 0 0 0 0 Oryzomys nigripes 4301000 Eurmgomatomys spinosus 2002000Tofars Fm im 75 9 28 12-66 Percentages 28.3 3.7 42.3 17.4 3.2 5.0 Table 8. Ecological place of each species captured during this studv in the Rio Doce State Forestry Park. GM = grasslands and wet meadows, B= brushy areas, 5 = secondary forests, P= primary forests, F= fossorial or semifossorial, SA= semiaquatic T= terrestrial, S= scansorial, A = arboreal, HG = herbivore-grazer; FG = frugivore-granivore; ~0 - frugivore-omnivore; IO = insectivore-omnivore. Spatial Dietary Species Habitat Adaptation Classification MARSUPIALS Didelphis marsupialis B, S, P T, S FO Metachirus nudicaudatus S, P T IO/FO Marmosa incana* B, S , P S IO M. cinerea B, S, P A IO M. microtarsus' S, P A IO Caluromys philander S, P A FO RODENTS Oecomys trinitatis S, P S FG Oryzomvs ca 51 mm MALES MASS 9 578.00 1300.00 938.89 1295.32 BODY 9 292.00 415.00 354.78 1 39.98 TAIL 9 312.00 393.00 355.33 1 28.41 EAR 9 47.00 55.00 50.671 2.78 FOOT 9 51.00 63.00 57.891 4.20 FEMALES MASS 8 568.00 1855.00 1158.62 +369.98 BODY 8 335.00 400.00 372.87 + 24.82 TAIL 8 345.00 400.00 377.25 1 18.01 EAR 8 46.00 58:00 51.371 3.93 FOOT 8 51.00 64.00 57.87 + 4.97 Memchirus nudicaudatus -- ADULTS, MASS > 90 g MALES MASS* 35 102.00 480.00 28131 +117.00 BODY 35 170.00 300.00 233.89 + 36.76 TAIL 35 227.00 373.00 307.66 1 41.67 EAR 35 28.00 40.00 35.40 + 2.66 FOOT** 35 35.00 52.00 43.71 1 3.86 FEMALES MASS 51 91.00 345.00 235.88168.93 BODY 50 150.00 265.00 222.931 2836 TAIL 51 178.00 363.00 297.961 43.19 EAR 50 31.00 43.00 35.721 2.62 FOOT 51 34.00 47.00 41.35 + 2.96 Marmosa incana -- ADULXS, MASS > 35 g MALES MASS 46 35.00 130.00 66.041 27.74 BODY 46 95.00 192.00 138.00 + 21.96 STALUNGS: BRAZILIAN SMALL MAMMAL INVENTORIES 195 N MIN MEAN + SD TAIL 45 162.00 296.00 195.091 2132 EAR** 46 24.00 32.00 27.52 + 1.96 FOOT ** 46 18.00 26.00 2132 + 1.72 FEMALES MASS 22 44.00 73.00 58.641 - 8.32 BODY 22 110.00 162.00 13730 1 11.90 TAIL 22 163.00 199.00 183.411 8.62 EAR 22 23.00 29.00 25.861 1.49 FOOT 22 16.00 24.00 19.821 1.79 Marmosa cinerea -- ADULTS, MASS > 50 g MALES MASS 36 56.00 194.00 109.94 1 28.75 BODY 36 146.00 210.00 176.89 1 16.08 TAIL* 35 200.00 291.00 259.63 1 20.42 EAR* 36 24.00 35.00 30.811 2.49 FOOT* 36 24.00 31.00 28.19 + 1.62 FEMALES MASS 28 53.00 230.00 99.071 41.40 BODY 28 125.00 205.00 165.68 1 23.09 TAIL 28 192.00 29100 248.68 + 26.41 EAR 28 24.00 34.00 29.14 1 2.77 FOOT 28 22.00 35.00 26.64 + 2.57 Marmosa microtarsus -- ADULTS MALES MASS 1 31.00 31.00 31.00 BODY 1 106.00 106.00 106.00 TAIL 1 148.00 148.00 148.00 EAR 1 14.00 14.00 14.00 FOOT 1 17.00 17.00 17.00 Caluromys philander-- ADULTS MALES MASS 18 123.00 261.00 189.83 1 35.72 BODY 18 180.00 245.00 215.33 1 17.08 TAIL 18 225.00 322.00 295.721 22.15 EAR 18 31.00 38.00 33.671 1.94 FOOT 18 32.00 41.00 35.611 2.00 196 BULLEI'IN FLORIDA SrATE MUSEUM 34(4) N MIN MAX MEAN 1 SD FEMALES MASS 12 115.00 286.00 186.75 i 44.67 BODY 12 200.00 240.00 216.75 1 12.27 TAIL 12 283.00 313.00 299.831 9.68 EAR 12 30.00 37.00 33.42 + 2.02 FOOT 12 30.00 42.00 35.00+ 2.92 Nectomys squamipes -- ADULrS, MASS > 60 g MALES MASS 8 75.00 235.00 146.12 1 64.97 BODY 8 138.00 203.00 173.00 + 24.63 TAIL 8 160.00 302.00 199.75 + 47.06 EAR 8 20.00 24.00 22.001 1.51 FOOT 8 42.00 53.00 47.87+ 332 FEMALES MASS 4 98.00 217.00 16030 + 57.97 BODY 4 165.00 200.00 183.001 17.11 TAIL 4 188.00 231.00 206.75120.35 EAR 4 21.00 24.00 23.00 1 1.41 FOOT 4 46.00 50.00 48.00 + 1.83 Rh 25 g MALES MASS 73 25.00 66.00 41.181 1035 BODY** 73 83.00 123.00 104.82 + 9.24 TAIL* 65 75.00 108.00 93.49 + 7.65 EAR 71 14.00 22.00 18.62 + 1.46 FOOT 72 21.00 30.00 26.061 1.38 FEMALES MASS 38 26.00 66.00 38.53 1 10.04 BODY 38 76.00 120.00 9933 1 10.99 TAIL 38 76.00 103.00 89.58 + 7.02 SIALLINGS: BRAZILIAN SMALL MAMMAL INVENTORIES 197 N MIN MEAN + SD EAR 37 16.00 20.00 18.30 + 1.02 FOOT 38 23.00 30.00 26.00 + 1.47 Oryzomys capito -- ADULTS, MASS > 35 g MALES MASS 5 38.00 72.00 59.60 + 13.81 BODY 5 118.00 135.00 127.601 8.14 TAIL 5 124.00 145.00 132.00 + 8.69 EAR 5 16.00 24.00 21.00 1 3.16 FOOT 5 23.00 35.00 30.801 4.55 FEMALES MASS 10 50.00 65.00 59.10 1 5.22 BODY 10 110.00 147.00 124.401 10.86 TAIL 10 115.00 135.00 125.40 + 6.15 EAR 9 21.00 22.00 21.44 + 033 FOOT 10 30.00 34.00 31.80 + 1.32 Oecomys trinitatis -- ADULTS MALES MASS 10 42.00 100.00 6530 + 16.95 BODY 10 114.00 135.00 124.60 + 7.53 TAIL 10 133.00 155.00 147.40 + 7.99 EAR 9 16.00 25.00 18.67 1 2.78 FOOT 10 13.00 39.00 2930 + 8.66 FEMALES MASS 5 61.00 95.00 76.20 1 12.87 BODY 5 124.00 143.00 131.80 1 7.79 TAIL 5 129.00 236.00 166.60 1 42.07 EAR 4 18.00 21.00 19.00 1 1.41 FOOT 5 28.00 33.00 30.60 + 2.07 Ofomys nigripes - ADULTS MALES MASS 2 25.00 28.00 26301 2.12 BODY 2 83.00 100.00 91301 12.02 TAIL 2 122.00 132.00 127.00 + 7.07 EAR 2 16.00 17.00 16.50 1 0.71 FOOT 2 22.00 26.00 24.00 + 2.83 198 BULLETIN FLORIDA SI'ATE MUSEUM 34(4) N MIN MAX MEAN 1 SD FEMALES MASS 2 7.00 22.00 14301 10.61 BODY 2 47.00 85.00 66.001-26.87 TAIL 2 83.00 115.00 99.001 22.63 EAR 2 15.00 15.00 15.001 0.00 FOOT 2 20.00 24.00 22.00 1 .2.83 Oiyzomys subj?avus -- ADULTS, MASS > 40 g MALES MASS 10 54.00 101.00 77.90 + 17.64 BODY 9 118.00 170.00 144.00 1 16.15 TAIL 9 147.00 193.00 172.671 16.40 EAR 9 22.00 26.00 23.78 1 1.30 FOOT 9 34.00 38.00 35.331 1.22 FEMALES MASS 12 58.00 135.00 8838 1 22.22 BODY 11 114.00 170.00 146.00 1 19.28 TAIL 11 140.00 200.00 178.09 1 17.48 EAR 11 20.00 27.00 23.82 + 2.32 FOOT 11 27.00 37.00 34.451 2.77 Abrawayaomys ruschii -- ADULTS MALES MASS 1 63.00 63.00 63.00 BODY 1 128.00 128.00 125.00 TAIL 1 146.00 146.00 146.00 EAR 1 20.00 20.00 20.00 FOOT 1 31.00 31.00 31.00 Catomys laucha -- ADULTS FEMALES MASS 1 20.00 20.00 20.00 BODY 0 - - - TAIL 1 74.00 74.00 74.00 EAR 1 17.CIO 17.00 17.00 FOOT . 1 19.00 19.00 19.00 STALLINGS: BRAZILIAN SMALL MAMMAL INVENTORIES 199 N MIN MAX MEAN 1 SD Oxymyctents roberti -- ADULTS MALES MASS 5 45.00 120.00 73.80 1 29.19 BODY 5 94.00 150.00 125.801 20.20 TAIL 3 105.00 125.00 118.00 + 11.27 EAR 5 19.00 25.00 21.40 + 2.19 FOOT 5 33.00 39.00 35.00 + 2.35 FEMALES MASS 4 82.00 110.00 93.00 1 12.62 BODY 4 145.00 245.00 173.00 1 48.19 TAIL 2 121.00 122.00 121.50 1 0.71 EAR 4 22.00 23.00 22.251 0.50 FOOT 4 30.00 36.00 32.501. 2.65 Euryzgomatomys spinosus -- ADULTS MALES MASS 2 165.00 210.00 18750 1 31.82 BODY 2 185.00 188.00 18630 + 1.14 TAIL 2 61.00 65.00 63.00 + 2.83 EAR 2 17.00 18.00 1730 + 0.71 FOOT 2 35.00 35.00 35.00 + 0.00 Cavia fulgida -- ADULTS MALES MASS 1 285.00 285.00 285.00 BODY 1 223.00 22100 223.00 TAIL 1 - - - EAR 1 22.00 22.00 22.00 FOOT 1 46.00 46.00 46.00 FEMALES MASS 1 280.00 280.00 280.00 BODY 1 234.00 234.00 234.00 TAIL 1 17.00 17.00 17.00 EAR 1 21.00 21.00 21.00 FOOT 1 47.00 47.00 47.00 200 BULLEUN FLORIDA STATE MUSEUM 34(4) N MIN MEAN + SD Dasyprocta azarae -- ADULTS MALES MASS 1 2560.00 2560.00 2560.00 BODY 1 450.00 450.00 450.00 TAIL 1 28.00 28.00 28.00 EAR 1 45.00 45.00 45.00 FOOT 1 130.00 130.00 130.00 FEMALES MASS 1 2056.00 2056.00 2056.00 BODY 1 460.00 460.00 460.00 TAIL 1 19.00 19.00 19.00 EAR 1 38.00 38.00 38.00 FOOT 1 120.00 120.00 120.60