Ocelot latrines: communication centers for Neotropical mammals

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1 Journal of Mammalogy, 98(1): , 017 DOI: /jmammal/gyw174 Published online November, 016 Ocelot latrines: communication centers for Neotropical mammals Travis W. King, * Roberto Salom-Pérez, Lisa A. Shipley, Howard B. Quigley, and Daniel H. Thornton School of the Environment, Washington State University, 53 Heald Hall, Pullman, WA , USA (TWK, LAS, DHT) Panthera-Costa Rica, P.O. Box , Curridabat, San José, Costa Rica (RS-P) Panthera, 8 West 40th Street, 18th Floor, NY 10018, USA (HBQ) * Correspondent: Travis.W.King@wsu.edu Olfactory communication among mammals remains poorly studied yet may be key to understanding their ecology. This is particularly true for mammalian carnivores, which rely extensively on scent marking for communication. Previous research suggests that carnivore latrines play a large role in both intra- and interspecific communication. Despite the apparent complexity of mammal use of latrines, little work has examined behavior patterns of species that visit latrines. We used motion-triggered video cameras to study use and behavior of mammals at ocelot (Leopardus pardalis) latrines in Costa Rica. We documented temporal patterns of use by the focal species (ocelots), diversity of mammalian species using latrines, and behaviors that occurred at these sites. Ocelots showed peaks in visitation every days and a shorter gap between intersexual versus intrasexual visits, supporting the idea that ocelot latrines are used to communicate information about reproductive status. Fourteen terrestrial mammal species visited the latrines, and these species engaged in a variety of behaviors, including mark investigation, scent marking, and acceptance of scent marks. The complexity and frequency of behaviors by nonfocal species suggest that latrines may play as important a role in communication for these other species as they do for ocelots. La comunicación olfativa entre mamíferos ha sido poco estudiada, sin embargo puede ser la clave para el entendimiento de su ecología. Esto es particularmente cierto para mamíferos carnívoros quienes dependen extensivamente de el marcaje de rastros olfativos para su comunicación. Estudios previos siguieren que las letrinas juegan un rol importante en la comunicación intra- e inter-específica. A pesar de la aparente complejidad del uso de letrinas por parte de mamíferos, pocos trabajos han examinado los patrones de comportamiento de las especies que las visitan. Utilizamos cámaras de videos sensibles al movimiento para estudiar el uso y comportamiento de mamíferos en letrinas de ocelotes (Leopardus pardalis) en Costa Rica. Documentamos los patrones de uso temporal por parte de especies focales (ocelotes), la diversidad de especies de mamíferos y los comportamientos que ocurrieron en estos lugares. Los resultados indican que el uso por parte de los ocelotes muestran picos de visitas cada 60 a 67 días y un intervalo más corto para visitas inter-sexuales que para visitas intra-sexuales. Esto apoya la idea de que las letrinas de los ocelotes son utilizadas para comunicar estatus reproductivos. Un total de catorce mamíferos terrestres visitaron las letrinas y estas especies mostraron una variedad de comportamientos, incluyendo investigación, marcaje y aceptación de rastros olfativos. La complejidad y frecuencia de comportamientos por especies no focales, sugieren que las letrinas pueden jugar un rol importante en la comunicación de otras especies como lo es para los ocelotes. Key words: Eira barbara, interspecific communication, latrines, Leopardus pardalis, ocelot, scent marking In mammals, odors are a primary medium for the transmission of signals that play an important role in their daily lives (Brown and Macdonald 1985; Jordan 005). These olfactory signals, which form the basis of intra- and interspecific communication in many mammals, are an important component of the study of animal behavior. However, observation of these transmissions can often be difficult for cryptic species (Allen et al. 015). This is especially true for mammalian carnivores where the use of scent marking, the active deposition of feces or glandular secretions, is quite common (e.g., Felidae Mellen 1993; 016 American Society of Mammalogists, 106

2 KING ET AL. USE OF AND BEHAVIOR AT OCELOT LATRINES 107 raccoons, Procyon lotor Page et al. 1999; Eurasian badgers, Meles meles Palphramand and White 007; giant river otters, Pteronura brasiliensis Leuchtenberger et al. 01) but difficult to observe (Jordan 005). The secretions from each individual and species carry a characteristic odor supplying intra- and interspecific information of an individual s sex, diet, territory, reproductive state, and movements (Gorman and Trowbridge 1989), and thus are an invaluable source of information about the ecology of these species. Some species create collections of scent marks, typically feces and at least 1 additional secretion, at specific locations, forming what are known as latrines (Jordan 005; Darden et al. 008). Traditionally, the latrine s primary role was assumed to be territorial defense with individuals marking the edges of home ranges or asserting one s dominance (Emmons 1988; Jordan 005). However, more recent work suggests this may be an oversimplification, because these latrines often are used by multiple individuals for territorial maintenance and a host of other behaviors (e.g., mate attraction Gorman and Trowbridge 1989; Roper et al. 1993). Moreover, multiple species may use latrines, suggesting these sites are complex information centers. Despite the likely importance of latrines in communication, patterns of latrine use and behavior at latrine sites for many mammals is poorly known. Additional research on latrine use will therefore improve our understanding of olfactory communication among Mammalia. Although there is relatively low occurrence of latrine behavior in the Felidae, some species such as ocelots (Leopardus pardalis) do create and maintain latrines (Jordan 005; Moreno and Giacalone 006). Ocelots leave scent marks throughout their home ranges, but ocelot latrines are found in prominent places on the landscape (e.g., buttresses of large trees Moreno and Giacalone 014; Rodgers et al. 014; Hunter 015). Ocelots are not the only species to frequent these latrines. White-nosed coatis (Nasua narica), common opossums (Didelphis marsupialis), tayras (Eira barbara), striped hog-nosed skunks (Conepatus semistriatus), and northern tamanduas (Tamandua mexicana) have all been observed visiting these sites (Bustamante 008; Moreno and Giacalone 014). It has been hypothesized that these other species frequenting a latrine may be attracted to feed on the ocelot feces or to engage in some form of antipredator behavior (Bustamante 008; Moreno and Giacalone 014). Though some research has been conducted focusing on the importance of latrine behavior for ocelots (Moreno and Giacalone 006; Bustamante 008; Rodgers et al. 015), an extensive ethnographic analysis of use and behavior of the whole community of mammals that visit latrine sites has not been conducted. In this study, we used motion-triggered video cameras to document activity and behavior of ocelots and other mammals at known ocelot latrine sites in central Costa Rica. We determined 1) the temporal patterns of use by ocelots, ) the composition of mammalian species using these ocelot latrines, and 3) the behaviors that occurred at these sites for ocelots and other species. We hypothesized that latrines would serve as communication centers regarding reproductive status of ocelots, as suggested by Moreno and Giacalone (006) and Rodgers et al. (015). Thus, we predicted that 1) visitation frequencies of ocelots would be highly variable with notable peaks in activity and ) male female or female male visitation patterns would occur more frequently than male male or female female visitation patterns. Given the potential role of latrines as communication centers for multiple species, we also predicted that 3) multiple mammalian species will visit the sites and 4) other mammals in addition to ocelots will use the sites to deposit or accept (e.g., by body rubbing) scent marks. Materials and Methods Study area. This study took place in the Barbilla-Destierro Biological Sub-Corridor (BBS) region of the Central Caribbean slope of Costa Rica (Fig. 1). This area consists of a patchwork of forest, pasture, and agriculture. Approximately 360 km in area, the BBS acts as natural link between major mountain ranges of the country, the Cordillera Volcánica Central and the Cordillera Talamanca ranges. Following the bioclimatic classification of Holdridge (1967), this region can best be depicted as 3 different life zones: tropical wet forest (which predominates), premontane wet forest, and premontane rainforest, with annual precipitation varying between 4,000 and 6,000 mm, mean daily temperature ranging from 17 C to 4 C, and a dry season lasting between 0 and 5 months (Bolaños and Watson 1993). Common Neotropical species at latrine sites. The ocelot is a medium-sized spotted cat, weighing on average 8 10 kg (Hunter 015). Ocelots have a varied diet, with more than 50 documented prey species (Emmons 1987; Bustamante 008; Hunter 015). They are believed to be polyestrous though peaks in mating and parturition are seen in some populations with the timing of such peaks dependent on geographic location (Murray and Gardner 1997). Furthermore, like most felids, ocelots base their social system on the defense of a defined home range with territorial maintenance thought to be one of the main uses of their scent marking (Emmons 1988; Moreno and Giacalone 006). The tayra is one of the largest mustelids in the Neotropics and is a diurnal carnivore and opportunistic omnivore, consuming a variety of small vertebrates, fruits, carrion, and insects (Presley 000). Tayras are typically observed singly, in male female pairs, or as small groups of mother and subadult offspring (Defler 1980). This was the case in our study, with lone males and groups of females or a female with young males being observed. There is no reported predation on tayra by larger carnivores (Presley 000). White-nosed coatis are omnivorous members of the family Procyonidae and are a known prey item for ocelots and other large Neotropical carnivores (Emmons 1987; Moreno et al. 006). They can be seen in bands of several adult females with subadults or solitary adult males, and coatis, like tayras, are primarily diurnal (Gompper 1995). The common opossum is a nocturnal marsupial found throughout much of the Neotropics and is a prey item of ocelots and other Neotropical carnivores (Emmons 1987; Moreno

3 108 JOURNAL OF MAMMALOGY Fig. 1. The Barbilla-Destierro Biological Sub-Corridor region of Costa Rica displayed as a solid with the dots of the enlarged image indicating the locations of ocelot (Leopardus pardalis) latrines used in this study (each latrine used being > 0.5 km apart). et al. 006). Members of the genus Didelphis have been known to engage in coprophagy as an opportunistic food source (e.g., Didelphis virginiana Livingston et al. 005) and are omnivorous in nature. Field methods. We identified 4 ocelot latrines for this study (Fig. 1). Latrines were encountered in an ad hoc fashion during a previous study of ocelots conducted throughout the BBS. Land use within a 5-km buffer around each site was dominated by primary secondary growth forest (53 99%) and agricultural land ( 47% Environmental Systems Research Institute 011; Instituto Costarricense de Electricidad 014). Three of the 4 latrines were located between the buttresses of large trees with the final being in the nd story of an abandoned humanmade structure. The primary cover immediately surrounding these sites was forest though all were less than 0.5 km from agricultural land. We used video camera-traps (Bushnell TrophyCam HD Max Black, Overland Park, Kansas) triggered by infrared-sensitive motion detectors to monitor the 4 ocelot latrines 4 h per day for a 6-month period. The camera was set to film for 30 s at a time and would trigger again if the animal was still in view with a 1-s delay between captures. Each of the 4 latrines was monitored from June to December of 014, recording visitation and species composition at these sites. All mammal species visiting the sites were identified and behaviors recorded. Behavioral classifications were developed through the work of Mellen (1993), Bustamante (008), Leuchtenberger et al. (01), Moreno and Giacalone (014), Vogt et al. (014), and Allen et al. (015). Additionally, all ocelots recorded at the latrines were subsequently identified using their individual spot patterns. Sex was determined by the visual presence or absence of male gonads, and any unique features or unusual behaviors per visit were noted. Statistical analysis. To test predictions regarding the temporal patterns of use by ocelots (predictions 1 and ), we used log-survivorship bout analysis to examine the existence and extent of peaks in ocelot visitation and analysis of mean visitation lengths between and within ocelot sexes to examine the potential for reproductive communication. Data on visitations were summed across all 4 latrines for these analyses. To conduct the log-survivorship analysis, the cumulative frequency of gap lengths (on a logarithmic scale) was plotted against gap length in days (Sibly et al. 1990). The log-survivorship analysis is used to create a criterion to separate a time series of events into bouts of behavior with accumulation starting with the longest gap length. A broken-stick model was then fitted by eye to the data, and the interception point of the straight lines was taken to be the bout interval (Sibly et al. 1990). This bout interval was then used to divide the visitation time line to examine if peaks in ocelot visitation occurred. For each latrine site, we also determined the length (in days) between visits, i.e., the gap between visits by ocelots of the same, and the opposite, sex. We used program R version 3.1. (R Development Core Team 015) to determine if there was a significant difference in mean gap length between intersexual visitations (male female or female male visitation) and intrasexual visitations (male male or female female visitation) for ocelots.

4 KING ET AL. USE OF AND BEHAVIOR AT OCELOT LATRINES 109 To test predictions regarding use of latrine sites by other species and their behaviors at latrines (predictions 3 and 4), we identified all species that visited each latrine site, the number of independent visits (visits separated by more than 4 h), and categorized the behaviors displayed (Table 1; Supplementary Data SD1 SD6). Data were combined across all 4 latrines for the analysis. We restricted our behavioral analysis to only the 4 species that visited the ocelot latrines most frequently (ocelot, tayra, coati, and opossum). We then created an ethogram to describe the visitations and observed behaviors for the 4 species using the ocelot latrines most frequently (Table 1). Each visit by a species to a latrine was scored according to whether or not they displayed the behaviors listed in the ethogram. In a given visit, an individual could display multiple behaviors, e.g., sniffing and scent rubbing. We acknowledge that comparisons between behaviors are not statically independent and therefore, each behavior was analyzed separately. For each behavior, differences in frequency of occurrence for each species was compared using a Pearson s chi-square test for independence (α = 0.05) with significant values determined via analysis of the Pearson residuals (where values greater than were considered to be significantly different in comparison to the other species Gardener 01). Species were excluded from a test if no behavior was recorded. Where more than species were analyzed for a given behavior, Pearson residuals were used to determine direction (Gardener 01). In total, we performed 7 tests and Hommel s method was used to control overall false positive rate (Hommel 1988). Results Sixteen individual ocelots (identified by spot patterns) were identified using the latrines with multiple male and female ocelots visiting each latrine with the exception of the latrine within the artificial structure, where only a single male and female were observed. Of these ocelots, several females had visual signs of being in estrus as described in Moreno and Giacalone (006). Furthermore, several mother juvenile pairs and mother subadult pairs (individuals observed with mother several months prior to the start of this study) were seen visiting the latrines. The log-survivorship bout analysis for ocelot visitations revealed that peaks in ocelot activity can be classified into bouts (single temporal events) of approximately 15 days in length (Fig. ). During the study period, use of latrines by ocelots peaked at regular intervals (approximately every days) with 3 peaks observed (Fig. 3). When examining the timing of visitations by sex, the mean gap length was significantly shorter for intersexual visitations (X = 6.86 days) than for intrasexual visitations (X = days; t 81 =.988, P = 0.004; Fig. 4). A total of 14 terrestrial mammal species was identified visiting and using the ocelot latrines. These included 1 species of marsupial the common opossum (50 visits), 3 species of xenarthrans the northern tamandua (5 visits), 9-banded armadillo (Dasypus novemcinctus; 4 visits), and northern naked-tailed armadillo (Cabassous centralis; 19 visits); 5 species of carnivorans coyote (Canis latrans; visits), ocelot (63 visits), striped hog-nosed skunk (5 visits), tayra (88 visits), and white-nosed coati (44 visits); and 5 species of rodents agouti (Dasyprocta punctata; 7 visits), variegated squirrel (Sciurus variegatoides; visits), red-tailed squirrel (Sciurus granatensis; 5 visits), and unidentified rat species with 34 visitations. Tayras, ocelots, common opossums, and white-nosed coatis were the 4 most frequent users of the latrines. Table 1. An ethogram and frequency of behaviors by the 4 most frequent species visiting ocelot latrines in Costa Rica s Barbilla-Destierro Biological Sub-Corridor region from June to December 014. Asterisks indicate a statistically significant behavior as determined by the Pearson s chi-square test for independence on frequency of behaviors among species (α = 0.05) and analysis of Pearson residuals (where values greater than were considered significant). See Supplementary Data SD1 SD6 for examples. Behavior Description Occurrence of behavior by species (%) Scent marking defecation Scent marking urination Scent marking anal rubbing Facial rubbing Body rubbing Sniffing Foraging Ocelot (Leopardus pardalis) Tayra (Eira barbara) White-nosed coati (Nasua narica) Common opossum (Didelphis marsupialis) An individual defecated at the latrine. 35* 10 0 An individual squatted against the ground and urinated or sprayed the latrine with urine. An individual rubbed its anal region on the ground or against an object. An individual rubbed its cheeks or face against the ground or latrine (often in a repeated fashion). An individual rubbed its shoulder on the ground or latrine, rolled back and forth on the ground, or used its forepaws to rub material into its fur. An individual used its nasal region to investigate the latrine. An individual appeared to be searching for and possibly consumed material at the latrine. 56* * * * * Total number of visitations

5 110 JOURNAL OF MAMMALOGY Fig.. A log-survivorship curve of the cumulative frequency of gap lengths (the amount of days between visits) was plotted against gap length for ocelot (Leopardus pardalis) visitations to latrines in the Barbilla-Destierro Biological Sub-Corridor region of Costa Rica from June to December 014. A broken-stick model (the bisecting lines) was fitted to the curve, indicating the area on the graph with the greatest change in slope. The point at which these lines intersect is taken to be the bout length (@ 15 days). Fig. 4. The mean gap length (the time between visitations) given in days for intrasexual and intersexual visitations to latrines by ocelots (Leopardus pardalis) in the Barbilla-Destierro Biological Sub- Corridor region of Costa Rica from June to December 014. Error bars indicate 95% confidence intervals. A significant difference in gap length between intersexual and intrasexual visitations was detected (t 81 =.988, P = 0.004). level (with the exception of foraging; Table 1). There were substantial differences in the frequency of behavioral types among species. Ocelots engaged in a significantly higher amount of scent marking than the other 3 species (defecation 35%, χ = 13.7, P = 0.000, Residual =.64, and uri- nation 56%, χ = 59.3, P < , Residual = 5.3; see Table 1). In contrast, ocelots engaged in significantly less accepting of scent marks at latrines than the other 3 species (5%, χ 3 = 9.7, P < , Residual = 4.754). Tayras engaged in significantly more anal marking (3%, χ = 9.761, P = , Residual =.01) and facial rubbing (86%, χ 3 = 14.4, P < , Residual = 6.41) than the other 3 species, and significantly less sniffing (33%, χ 3 = 6.8, P < , Residual = 3.54; Table 1). Fig. 3. A plot of ocelot (Leopardus pardalis) visitation frequency to latrines (the number of visitations in a given 15 days) against the days since trapping started in the Barbilla-Destierro Biological Sub-Corridor region of Costa Rica from 4 June to 1 December 014. A total of 13 bouts were created from the 19 days of trapping. Because 19 days did not divide equally into 13 bouts of 15 days, a bout of 1 days was made and placed at the beginning of the graph. Whether this shortened bout was placed at the beginning or end of the graph did not change the observed pattern. For the graph, a spline was used to create a smooth line connecting the dots, which represent the actual data. Species that visited latrine sites engaged in a wide variety of behaviors. As predicted, these behaviors included scent marking and acceptance of scent marks. Three of the 4 most frequent visitors (tayras, ocelots, common opossums, and white-nosed coatis) engaged in all of the behaviors at some Discussion Our study is one of the first to obtain detailed behavioral data from multiple species at latrine sites and demonstrates that ocelot latrines act as information centers for multiple mammalian species. The complexity and frequency of behaviors by the nonfocal species (particularly tayras) suggest that these latrines may play as important a role in communication for these other species as they do for ocelots, and that latrines may have greater functionality beyond their role as places to leave, or explore, intraspecific scent marks. Our study also reinforces suggestions that ocelot latrines have important roles to play in reproduction, as we documented frequent cross-sex visitation and bouts of activity that may be related to the reproductive cycle. Taken together, our findings indicate that latrine sites serve as focal points for olfactory communication in Neotropical mammals.

6 KING ET AL. USE OF AND BEHAVIOR AT OCELOT LATRINES 111 Our observation that 14 species of terrestrial mammals visited the 4 ocelot latrines during this study supports the idea that the latrines act as important communication hubs for the local mammalian communities (Bustamante 008; Moreno and Giacalone 014; Rodgers et al. 015). In addition to the 4 most frequent visiting species in our ethnographic analysis, several other species engaged in obvious behaviors at latrines sites, including foraging (i.e., 9-banded armadillo), sniffing (i.e., agouti), and rubbing (i.e., rat species). Ocelot latrines may act as a food resource for some visiting species, which has been observed as a driving force in the visitation of both raccoon (Page et al. 1999) and giant river otter (Leuchtenberger et al. 01) latrines. Species such as common opossums and rodents were seen directly feeding on feces at the latrines, whereas both species of armadillo and striped hog-nosed skunks seem to be foraging on the abundant insects found at the sites. The differing frequencies of behavior that were displayed by the most common visitors to latrines (ocelots, tayras, opossums, and coatis) may indicate fundamental differences in how the latrines are used by a diverse mammal fauna. Common opossums and white-nosed coatis displayed a high frequency of scent acceptance behavior (e.g., body rubbing), but limited or no scent marking behavior at the latrine sites (similar behaviors were also seen in the unidentified rat species). This behavior, which makes an individual smell like an ocelot, may act as an antipredator or predator avoidance technique because smelling like an ocelot may reduce risk of predation from carnivores (Bustamante 008); both opossums and coatis are occasional prey items for ocelots and other larger carnivores (Moreno et al. 006; Bustamante 008; Hunter 015). Tayras visited the putative ocelot latrines more frequently than ocelots, and both species engaged in scent marking and scent acceptance behavior quite frequently, although scent acceptance was more common than scent marking for tayras, and vice versa for ocelots. This suggests a strong role for these latrines in intraspecific communication for both of these species. These patterns of behavior also suggest some degree of interspecific communication may be occurring. One possible explanation for the high degree of marking and rubbing by these species at the same latrines could be that announcing your presence to other mesopredators at ocelot latrines helps to reduce acts of aggression between neighbors known as the dear enemy phenomenon (Palphramand and White 007). The dear enemy phenomenon is the recognition and tolerance of neighbors and aggression and intolerance of strangers within ones landscape (Fisher 1954). This effect has been noted in a number of mammalian species, including intraspecific occurrences in badgers (Müller and Manser 007; Palphramand and White 007) and interspecific occurrences in species of Peromyscus (Wolff et al. 1983). This recognition may allow these neighbors to focus aggressive action on newcomers to the area rather than possibly wasting efforts on already established members of their region. However, further studies are needed to make this connection, as this phenomenon is not well studied; the suggestion that territoriality may be based on familiarity with neighbors has been noted for bobcats (Lynx rufus Allen et al. 015). The temporal patterns in latrine use by ocelots show peaks in visitation. This may suggest that latrines play a role in mating, as multiple males and females mark and examine these sites during bouts of activity. This idea is reinforced by the fact that intersexual visitation rates were more frequent than intrasexual visitation. Sex and reproductive status of individual felids can be obtained via their scent marks (Jordan 005). Heightened use of latrine sites has been noted as a good indication of sexual reproductive activity in small felids (and specifically ocelots Mellen 1993; Murray and Gardner 1997) and this pattern of greater latrine use during probable mating periods has been noted for badgers (Roper et al. 1993), river otters (Lontra canadensis Stevens and Serfass 008), and bobcats (Allen et al. 015). These results are supported by the work of Moreno and Giacalone (006), who found a rise in number of ocelots visiting a latrine when a female had visual signs of being in estrus (a visibly pink vulva). Within our study, similar visual evidence of estrus was noted on 3 occasions. However, use of black-and-white nighttime videos in our study limited our ability to identify estrus during latrine visitation. Furthermore, the frequent acts of both facial rubbing and urination observed by female ocelots also can be indicators of estrus or reproductive activity (Verberne and Leyhausen 1976; Mellen 1993). The peaks in ocelot visitation occurred across all the latrines in the study within the same bouts of activity (roughly at the same time), which may suggest these visits are trigged by external stimuli. For example, these peaks which are separated by approximately days are near the length of lunar cycles. However, a longer study would be needed to test this hypothesis. Furthermore, the shorter-duration gap lengths between intersexual visitations than between intrasexual visitations also may indicate that latrines play a role in reproductive activities because females may not simply appear once when ready to mate. Instead, visitations from both sexes may act as a progression from initial investigation and scent marking to indicate availability of a female, to a male s response, which may lead to further response or investigation by the female or eventual mating (Moreno and Giacalone 006; Allen et al. 014). From this possible progression of actions, one would expect a rapid turnaround between intersexual visitations, which we observed. Note that not all ocelot visitations can be explained as mating orientated, as both mother cub pairs and subadults who are likely nonreproductive can be seen at the sites. This supports the idea of ocelot latrines playing multiple roles in mammal communication (Gorman and Trowbridge 1989; Roper et al. 1993; Moreno and Giacalone 006; Moreno and Giacalone 014; Rodgers et al. 015). This investigation provides novel information on the importance of ocelot latrines for Neotropical mammals. Future studies should be conducted on a longer temporal scale with multiple latrines, and perhaps involving manipulation of latrines, to test the validity of the general and seasonal patterns documented here, as well as testing for the ultimate causes of many of the behaviors witnessed at the latrines in this study. Nevertheless, our results provide valuable species composition and behavioral data for several elusive species, and key details on the

7 11 JOURNAL OF MAMMALOGY complicated communication networks that remain poorly known for Neotropical mammals. Acknowledgments Funding was provided by the Washington State University College of Arts and Sciences Undergraduate Summer Minigrant for Research, Honors College Jacklin Family Honors Education Abroad Scholarship, Auvil Scholars Fellowship, and Panthera Small Grant for Summer Research. We wish to thank Panthera and Panthera-Costa Rica for their collaboration, and field support in Costa Rica, and we especially thank E. Urbina Ruiz for continuing to monitor the cameras. We thank P. Verrell for his input on behavioral analysis and possible explanations of observed behavioral patterns. We thank T. Coffey of Washington State University CISER for review of our statistical analysis. Supplementary Data Supplementary Data SD1. Video of a female ocelot investigating (sniffing)/scent marking (urination) a latrine in the Barbilla-Destierro Biological Sub-Corridor region of Costa Rica Supplementary Data SD. Video of a male tayra investigating (sniffing), body rubbing, and anal marking at a latrine in the Barbilla-Destierro Biological Sub-Corridor region of Costa Rica Supplementary Data SD3. Video of a family group of white-nosed coatis investigating (sniffing) and engaged in body rubbing at an ocelot latrine in the Barbilla-Destierro Biological Sub-Corridor region of Costa Rica Supplementary Data SD4. Video of a common opossum investigating (sniffing) and foraging at an ocelot latrine in the Barbilla-Destierro Biological Sub-Corridor region of Costa Rica Supplementary Data SD5. Video of a female ocelot investigating (sniffing), facial rubbing, and scent marking (defecation) at a latrine in the Barbilla-Destierro Biological Sub-Corridor region of Costa Rica Supplementary Data SD6. 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