INTRODUCTION

The oncilla also called “northern tiger cat” (Leopardus tigrinus oncilla) is a little known and poorly studied felid subspecies from Central America, only reported on two sites in Panama (de Oliveira et al., 2020; Meyer et al., 2015; Rodgers & Kapheim, 2017). It is currently considered Vulnerable (VU) according to the latest IUCN Red List assessment (Payan & Oliveira, 2016; Pineda-Guerrero, 2018). This felid used to be considered a single unique subspecies until genetic studies split them between L. t. oncilla (Northern tiger cat) and L. gutulus (Southern tiger cat) (O’Brien & Koepfli, 2013; Payan & Oliveira, 2016). This northern subspecies has been reported from Costa Rica (3,625 m a.s.l.), Panama (3,475 m a.s.l.), and Colombia (4,800 m a.s.l.), confirming a preference for cloud forest (de Oliveira et al., 2008; Pineda-Guerrero, 2018; Rodgers & Kapheim, 2017; Rogan, 2021). In Panama, it has been genetically detected via scat from Chiriqui highlands, Barú Volcano National Park (Rodgers & Kapheim, 2017), and via camera traps studies from Darien National Park (Meyer et al., 2015). Despite several biodiversity studies throughout the country, including the Panama Canal Watershed (Méndez-Carvajal, 2012), Cerro Hoya National Park (Fort et al., 2014), El Montuoso Forest Reserve (Méndez-Carvajal et al., 2020), Central Panama (Meyer et al., 2015), and southeastern Darien (Méndez-Carvajal et al., 2021), where the oncilla have not been detected. Although it has been difficult to assess L. t. oncilla due to its cryptic behaviour, some population densities have been published, encouraging additional studies to better understand the species’ conservation status, distribution, and threats (de Oliveira et al., 2020). In this study, we evaluate the presence of L. t. oncilla in Darien-Panama at the Chucantí Private Natural Reserve (CPNR), considered a hotspot for endemism and diversity.

METHODS

Study area

Study area

The Chucantí Private Nature Reserve (CPNR; N 08°47 ́16”, W 078°27 ́01”) is located in the Darién province, Republic of Panama (Laurance, 2008). The average temperature falls between 24.0-27.2 ºC, with approximately 1,940.5 mm of precipitation annually (Gutiérrez- Pineda et al., 2021). The site is a mixture of montane and submontane forest (Méndez-

Carvajal, 2012). It has a high degree of plant diversity and endemism, largely dominated by the families Rosaceae, Magnoliaceae, Gentianaceae and Fabaceae (Ortiz et al., 2016; Flores et al., 2017; Mijango-Ramos et al., 2020) (Fig. 1). This study was carried out under scientific permission No. SE/A-53-18 and ARB-0028-2021.

Data collection for terrestrial mammals

Data were generated by a long-term monitoring project that utilized three camera traps (Cuddeback and Bushnell Trophy Cam models) at the ground-level of the forest, which operated from December 2012 to May 2014 (Méndez-Carvajal in prep.). Camera traps were positioned along the first two kilometers of the trail on the left-hand side. The cameras were located as follows: station 1 "Tronco” (800 m.a.s.l; N 08º47'36.69'' W 78º27'33''), station 2 "Filo 1” and station 3 "Filo 2" (1,375 m.a.s.l; N 08º48' 05.01'' or W 78º27'42.038''). The cameras were spaced 1 km apart along the 3 km trail (Gutiérrez-Pineda et al., 2021) (Figure. 1).

Figure 1.
Map of the study area and location of Chucantí Private Nature Reserve, Agua Fría, Chepigana, Darién province, Republic of Panama.
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Data analysis

We calculated the frequency of detection of the number of species per month. Having acquired more than 11 photographic events with intervals ≥30 min apart, we were able to generate circadian activity graphs (Mosquera-Muñoz, 2014). All statistics were analyzed using PALeontological STatistics software (PAST 4.02).

RESULTS

We detected Leopardus tigrinus oncilla between 19 terrestrial mammal species across 38, 592 trapnight/hours. Particularly, L. t. oncilla appeared at the three sites sampled (Tronco,

Filo1 and Filo 2) from 800 to 1,375 m.a.s.l. in the cloud forest. L. t. oncilla appeared to be mostly active between 17:00 to 04:00, confirming a marked nocturnal activity (Table 1).

Figure 2.
Circadian activity of L. t. oncilla at Chucanti Private Natural Reserve, Agua Fría, Chepigana, Darien, Republic of Panama.
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Identification of L. t. oncilla was based on tail position, orbital distance between eyes, small body size compared at the same position using a natural reference for different felines photographed at the same station, frontal extension between ears and eyes are short (Figure 3). IDs were confirmed by Tadeu de Oliveira, a specialist on this taxon.

Figure 3.
Photo-capture of L. t. oncilla at Chucanti Private Natural Reserve, Darien province, Panama.
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DISCUSSION

This study reports the eastern-most distribution for L. t. oncilla in Panama. This species may have been missed on previous mammal diversity studies, possibly due to short study periods and reduced effort within the central provinces of Panama, as noted by Fort et al. (2014). Updating biogeographical information helps improve distribution knowledge and identify potential conservation areas as priorities (Loyola et al., 2008; Méndez-Carvajal et al., 2021). Our results showed that L. t. oncilla is sympatric with five other wild feline species as Panthera onca, Puma concolor, Leopardus pardalis, Leopardus wiedii, and Herpailurus yagouaroundi in Chucanti, making this area a unique zone in Panama. Furthermore, due to this high felid sympatry, this is one of the few areas in L. t. oncilla`s entire distribution where such an interspecific ecological niche study could be undertaken. Chucanti Private Natural Reserve is an ideal place to examine ecological problems related to the co-existence of species with similar morphological adaptations and diet (Tokeshi, 2009). The number of prey species at Chucanti are well defined for big cats such as Panthera onca and Puma concolor, which also made few appearances on our camera traps over two consecutive years. In fact, these large cats were captured less than their regular prey, including Cuniculus paca, Mazama temama, Odocoileus virginianus, Pecari tajacu, and Crax rubra, as reported in Darien by Arosemena (2017) and Moreno (2006).

Interestingly, L. t. oncilla may maintain a different activity pattern compared to sympatric margay (L. wiedii) and ocelots (L. pardalis). According to the principle of intraguild predation proposed in Polis et al. (1989), the felids of Chucanti could be segregating themselves by habitat, where small felids are expected to have smaller ranges and large cats increase their habitat range, as reported for de Oliveira & Pereira (2014). It is important to note that there are reports of social predators such as coyotes (Canis latrans) near Chucanti (Bermúdez et al., 2012), though we did not detect this species within the main reserve possibly due to the presence of a P. onca and P. concolor. Social predators, such as coyotes, tend to put more pressure on small mammals in habitats where they dominate, impacting mammal diversity. The L. t. oncilla are considered victims of intraguild predators, and for areas fragmented or with human presence, pressures may increase if hunting dogs, and feral cats are nearby. In Chucanti, hunting dogs were detected on the same trails, while there has also been evidence of domestic cats infected with Ricketssia felis around the scientific station, which could lead to zoonotic disease transmission and threaten wildlife populations (Bermúdez et al., 2012; Vieira et al., 2018).

CONCLUSION

This study confirmed the presence of one of the least known Neotropical felid species in eastern Panama. The presence of L. t. oncilla in the area between Chiriquí to Darien, supports the potential for this species having a broader distribution in Panama. This study also supports Chucanti Private Natural Reserve as an important site to study the effects of sympatric felines on mammal populations as it is the only known habitat in Panama with six feline species. Although previously reported from other habitats, both hunting pressure and the presence of domestic canids and felines can cause threaten population stability. Finally, our study reinforces the importance of long-term biodiversity monitoring for expanding our understanding of cryptic and elusive species with low densities across landscapes.

Acknowledgments

ACKNOWLEDGEMENTS

We thank the Fundación Pro-Conservación de los Primates Panameños (FCPP), Primatology Research Group of the University of Panama (GIP-UP), Yaguará Panamá S.A., ADOPTA Bosque Panamá for supporting this study at a logistical level. This long-term study was funded by Idea Wild, Mohamed bin-Zayed Species Conservation Fund (project # 1025476 and 12055182); II Rufford Small Grant for Nature Conservation (project #16021) and Re:wild (2023). Our thanks to the students of the different primatology courses carried out by FCPP-GIP-UP, who kept the monitoring project year after year in the reserve to the present. To the park rangers of the reserve Juan Camaño, Arcelio Castillo. Special thanks to Tadeu de Oliveira, wildcat specialist from Maranhão State University (UEMA), Brazil and member of the IUCN Red List of Threatened Species, who kindly helped to confirm the ID of some of the L.t.oncilla pictures. We thank Timothy M. Eppley, for English edition and review on previous version of this manuscript as well editors from Tecnociencia. We thank Ricardo Moreno and Guido Berguido as colaborators in this project.

References

Arosemena, M. (2017). El uso de trampas cámaras como herramienta de evaluación y conservación de mamíferos silvestres. Visión Antataura. 1(2): 89-90.

Flores, R., C. Black, C. & A. Ibáñez (2017). A new species of Heliconia (Heliconiaceae) with pendent inflorescence, from Chucantí Private Nature Reserve, eastern Panama. PhytoKeys. (77): 21.

Fort, J. L., C.K. Nielsen, E.R. I.C., Donoso, R. Samudio & G. A. Durán (2014). First camera survey of wild felids in Cerro Hoya National Park, Panama. Cat News. 60: 36-37.

Gutiérrez-Pineda, K. M., G. Berguido & P.G. Méndez-Carvajal (2021). Diversidad ecológica de aves caminadoras en la Reserva Natural Privada Cerro Chucantí, Darién, Panamá. Mesoamericana. 25(1): 1-14.

de Oliveira, T.G., J. Schipper & J.F. Gonzalez-Maya (2008). Leopardus tigrinus ssp. oncilla. In: IUCN 2009. IUCN Red List of Threatened Species. Version 2009.2. . Downloaded on 09 January 2010.

de Oliveira, T.G. & J.A. Pereira (2014). Intraguild predation and interspecific killing as structuring forces of carnivoran communities in South America. Journal of Mammalian Evolution. 21: 427-436.

de Oliveira, T.G., B.C. Lima, L. Fox-Rosales, R.S. Pereira, E. Pontes-Araujo & A.L. de Sousa (2020). A refined population and conservation assessment of the elusive and endangered northern tiger cat (Leopardus tigrinus) in its key worldwide conservation area in Brazil. Global Ecology and Conservation. 22: e00927.

Bermúdez, S., R. Miranda, Y. Zaldívar, P. González, G. Berguido, D. Trejos, ... & M. Labruna (2012). Detección de Rickettsia spp. en ectoparásitos de animales domésticos y silvestres de la Reserva Natural Privada Cerro Chucantí y comunidades aledañas, Panamá, 2007-2010. Biomédica. 32(2): 189-195.

Rodgers, T. W. & K.M. Kapheim (2017). A High-Elevation Record of the Little Spotted Cat (Leopardus tigrinus oncilla) from Western Panama. The Southwestern Naturalist. 62(3): 225-227.

Rogan, J. E. (2021). First Locality Record of Melanistic Oncilla (Leopardus tigrinus) in Monteverde, Costa Rica. Neotropical Biology and Conservation. 16(3): 427-434.

Laurance, W. F. (2008) Adopt a forest. Biotropica. 40(1): 3-6.

Loyola, R. D., G. de Oliveira, J.A.F. Diniz‐Filho & T.M. Lewinsohn (2008). Conservation of Neotropical carnivores under different prioritization scenarios: mapping species traits to minimize conservation conflicts. Diversity and Distributions. 14(6): 949-960.

Méndez-Carvajal, P.G. (2012). Censo preliminar de primates en la Reserva Natural Chucantí, provincia de Darién, República de Panamá. Mesoamericana. 16(3): 22-29.

Méndez-Carvajal, P. G., I.G. de Huertas, K.M. Gutiérrez-Pineda, R.S. Moreno, M.A. Peñafiel, A.S. Girón-Rengifo… & P.A. González-Hernández (2020). Potencial regenerativo de bosques de galería en base a diversidad y abundancia de mamíferos en la Reserva Forestal El Montuoso y afluentes del río La Villa, Herrera, Panamá. Mesoamericana. 24(1): 58-76.

Méndez-Carvajal, P. G., A. Batista, O. Jaramillo, B. Rodríguez-Beitía & K.M. Gutiérrez- Pineda (2021). Biodiversity assessment at the southeastern side of Darien National Park, Panama: Diversity of southwestern Darien. Mesoamericana. 25(1): 22-36.

Meyer, N. F., H.J. Esser, R. Moreno, F. van Langevelde, Y. Liefting, D.R. Oller, ... & P.A. Jansen (2015). An assessment of the terrestrial mammal communities in forests of Central Panama, using camera-trap surveys. Journal for Nature Conservation. 26: 28-35.

Mijango-Ramos, Z., M.S. de Stapf, C. Vergara & J. Mendieta (2020). Diversidad de Árboles y Arbustos en la Reserva Privada Cerro Chucantí en Darién, Panamá. Tecnociencia. 22(1): 17-36.

Moreno, R. (2006). Parámetros poblacionales y aspectos ecológicos de los felinos y sus presas en Cana, Parque Nacional Darien, Panamá. Tesis de Maestría. Instituto Internacional en Conservación y manejo de Vida Silvestre, Heredia, Costa Rica. 136 pp.

Mosquera-Muñoz, D. M., G. Corredor, P. Cardona & I. Armbrecht (2014). Fototrampeo de aves caminadoras y mamíferos asociados en el piedemonte de Farallones de Cali. Bol. Cient. Mus. Hist. Nat. U. de Caldas. 18 (2): 144-156.

O'Brien, S. J. & K.P. Koepfli (2013). Evolution: a new cat species emerges. Current Biology. 23(24): R1103-R1105.

Ortiz, O. O., R.M. Baldini, G. Berguido & T.B. Croat (2016). New species of Anthurium (Araceae) from Chucantí nature reserve, eastern Panama. Phytotaxa. 255(1): 47-56.

Payan, E. & T.G. de Oliveira (2016). Leopardus tigrinus. The IUCN Red List of Threatened Species 2016: e.T54012637A50653881. https://dx.doi.org/10.2305/IUCN.UK.2016- 2.RLTS.T54012637A50653881.en. Accessed on 10 April 2023.

Pineda-Guerrero, A. (2018). Nuevo registro de Leopardus tigrinus (Carnivora: Felidae) en la Reserva Forestal Protectora Bosque Oriental de Bogotá, Colombia. Mammalogy Notes. 5(1-2): 15-17.

Polis, G.A, C.A. Myers, R.D. Holt. (1989). The ecology and evolution of intraguild predation: Potential competitors that eat each other. Annu. Rev Ecol Systemat. 20:297- 330.

Tokeshi, M. (2009). Species coexistence: ecological and evolutionary perspectives. John Wiley & Sons.

Vieira, F. E. G., D.L.F. Caetano, F.R. Barbosa, G.S. de Oliveira, J.N.K. de Souza-Serafim, J. N. K. & J.P. Sasse. (2018). Parasitismo gastrointestinal em Leopardus trigrinus (Schreber, 1775) (Carnivora; Felidae) e carnívoros domésticos no norte do estado do Paraná, Brasil. Brazilian Journal of Animal and Environmental Research. 1(2): 373- 385.