Circadian rhythm overlap between felids (Carnivora: Felidae) and their prey in a Subtropical Atlantic Forest protected area

ABSTRACT

When morphologically similar species, such as felids, coexist, resource sharing occurs across various niche dimensions-temporal, spatial, and trophic-to reduce competition. This study evaluated the circadian rhythm overlap between five Neotropical felid species and their most frequently recorded prey. Circadian rhythms were determined using Kernel density estimation based on camera-trap records. Pairwise comparisons between each felid and its prey were performed using overlap coefficients ($\Delta$), and differences in activity patterns were tested using the Mardia-Watson-Wheeler test. Overlap values between felids and their main prey were high for Puma concolor, Leopardus pardalis, and Leopardus wiedii. In contrast, Leopardus guttulus did not exhibit high overlap with its prey, and Herpailurus yagouaroundi did not yield the minimum records required for overlap analysis. The three larger felids synchronize their activity with the peak periods of their prey, suggesting that the benefits of successful foraging outweigh the costs of potential antagonistic encounters. The two larger species were influenced by the bottom-up effect, as expected. However, only one of the smaller species was influenced by the top-down effect, which may be related to the broader use of forest strata by L. wiedii, reducing its vulnerability to predation by larger felids.

KEYWORDS:
bottom-up effect; ecological competition; ecological niche; optimal foraging; tropical Forest

INTRODUCTION

The phenomenon of segregation into different niche dimensions is expected to reduce competition among morphologically similar species (Kiltie 1984). Felids possess a conservative morphology with discrete modifications that enable this species complex to adapt to diverse environments (Bellani 2020). Across the tropics, the co-occurrence of felid species can result in adaptations in their spatiotemporal habi tat use or promote shifts in feeding habits (Schoener 1974, Sunquist and Sunquist 1989, Moreno et al. 2006). Therefore, mechanisms that reduce niche overlap in at least one of the primary ecological dimensions are advantageous for co-occurring species (Di Bitetti et al. 2010, Nagy-Reis et al. 2019).

Although some felids share areas, their use of space is characterized by habitat heterogeneity and complexity (sensu August 1983), with horizontal displacement often occurring as a function of body size. For instance, large-bodied species typically occupy wider territories (Goulart et al. 2009), whereas small-bodied felids-such as Leopardus wiedii (Schinz, 1821)-are adapted to the vertical strata of tropical forests (Oliveira et al. 2010). Moreover, evidence suggests that Neotropical felids exhibit temporal segregation in their circadian rhythms, facilitating coexistence and reducing the probability of intraguild predation (Oliveira-Santos et al. 2012, Nagy-Reis et al. 2019).

Regarding dietary niches, felid species often have overlapping diets, a primary determinant of interspecific competition (Polis et al. 1989, Palomares and Caro 1999). Food availability is a key factor influencing felid activity patterns, leading to the synchronization of their rhythms with those of their major prey (Monterroso et al. 2013). Optimal foraging theory predicts that predators synchronize their activities with their prey to minimize energetic costs and increase capture success (Frey et al. 2017, Botts et al. 2020). Conversely, prey strive to reduce exposure to predators (Lima 2002), while predators must overlap their foraging schedules with prey activity. Simultaneously, smaller felids may avoid competition and decrease predation risks from larger species (Foster et al. 2013, Monterroso et al. 2013). The nocturnal, solitary, and territorial habits of most felids are influenced by the balance of these factors (Emmons 1988). In this context, large-bodied carnivores often follow a bottom-up pattern influenced by prey activity, whereas small-bodied carnivores may follow a top-down pattern influenced by the presence of larger predators (Vallejo-Vargas et al. 2022).

We aimed to evaluate the temporal niche overlap and activity patterns of five felid species and their prey in a protected area of the Southern Atlantic Forest, Brazil. We hypothesized that the circadian rhythms of larger felids-Puma concolor (Linnaeus, 1771) and Leopardus pardalis (Linnaeus, 1758))-overlap with those of their most common prey (bottom-up effect). In contrast, we hypothesized that smaller felids-L. wiedii, Leopardus guttulus (Hensel, 1872), and Herpailurus yagouaroundi (É. Geoffroy, 1803)-do not exhibit a strictly prey-synchronized pattern, as they are primarily controlled by the top-down effect of larger intraguild predators.

MATERIAL AND METHODS

Our study area is located in the buffer zone of the Aguaí State Biological Reserve, on the Serra Geral slopes of Treviso municipality (28°29’23”S and 49°31’12”W), Santa Catarina, Brazil (Fig. 1). The Aguaí State Biological Reserve was founded in 1983, occupies 76.72 km² in the core zone of the Atlantic Forest Biosphere Reserve, and is the second largest fully protected area in Santa Catarina, with high connectivity to other protected areas in Serra Geral (FATMA 2009).

Figure 1
Location of the study area (green dot), in the buffer zone of the Aguaí State Biological Reserve (yellow polygon), in the Treviso municipality, southern Santa Catarina state, Brazil.

According to the Köppen-Geiger climate classification, the study area has a humid subtropical climate, with mean annual precipitation of 1,200-1,600 mm and mean annual temperatures of 17.0-19.3 °C (Alvares et al. 2014). Mean hottest temperatures range from 23.4-25.9 °C in January, and mean coldest temperatures from 12.0-15.0 °C in July (Alvares et al. 2014). The relief is highly rugged, characterized by deep valleys with drops exceeding 1,000 m (Rocha 2016). The phytophysiognomy is typical of the southern Atlantic Forest, marked by Submontane Dense Ombrophylous Forest (Vibrans et al. 2013), with varying conservation levels due to historical exploitation and logging. The landscape consists of forest remnants on the slopes of Serra Geral and a mosaic of human-dominated areas such as urban zones, pastures, and agricultural fields.

Data were obtained from the Laboratory of Zoology and Ecology of Vertebrates (LABZEV) database at Universidade do Extremo Sul Catarinense (UNESC). We recorded felid and prey activity from February 2018 to July 2020 using 18 camera-traps programmed with a 2-second interval bet ween photos, operating 24 hours a day for 30 months (18 traps × 911 days). Effective sampling effort, calculated by subtracting periods with interruptions from the total days in the field, resulted in 16,398 camera-trap-days. Sampling points were at least 300 m apart, placed on trails, abandoned roads, or near watercourses at 0.40 m above ground, without olfactory attractants or baits.

We conducted a literature review via Scopus platform using the terms “diet”, “Felidae”, and “Atlantic Forest” to characterize felid diets across the Atlantic Forest domain, composing an adjacency matrix or grouping prey into taxa (e.g., lizards, snakes, birds, primates, and mammals < 1 kg) with the highest consumption frequency for each felid species. Frequency values were used to identify the most important prey in each felid’s diet. Prey ranked from first to fifth in consumption frequency within each article were assigned scores on a declining scale from 5 to 1, respectively. Points for each prey were summed across articles to provide an overall ranking of the most consumed prey per felid (Table 1).

Table 1
List of probable prey consumed by felids in the Atlantic Forest. This list focused on the study species, Leopardus guttulus, L. pardalis, L. wiedii and Puma concolor, and was created based on scientific articles available on the Scopus platform when searching with the terms “diet”, “Felidae” and “Atlantic Forest”.

Felines	Prey preference rank
	1st	2nd	3rd
Puma concolor	Cuniculus paca	Dasypus spp.	Nasua nasua
Leopardus pardalis	Mammals < 1 kg	Birds	Salvator merianae
L. wiedii	Mammals < 1 kg	Birds	-
L. guttulus	Mammals < 1 kg	Birds	S. merianae

Once prey rankings were determined, we verified which taxa and felid species had sufficient independent records (minimum of 20) obtained via camera-trapping (one record/species/hour/sampling point, unless multiple individuals or species appeared in the same photo). The three highest-ranking taxa were defined as having potential circadian synchronization with their predators.

To determine circadian activity overlap, we used kernel density estimation to obtain probability densities. The overlap coefficient (∆) varies from 0 (no overlap) to 1 (complete overlap) (Meredith and Ridout 2014), with overlaps classified as low (≤0.49), moderate (0.50-0.74), and high (≥0.75) (adapted from Massara et al. 2018). Finally, we used the Mardia-Watson-Wheeler test to evaluate homogeneity between rhythms, defining differences at p < 0.05. Using R-code (R Core Team 2020) and the Overlap R-package (Meredith and Ridout 2014), we calculated the circadian overlap between felids and their prey and tested rhythm homogeneity.

RESULTS

We recorded five felid species across the study area: H. yagouaroundi, L. guttulus, L. pardalis, L. wiedii, and P. concolor. Only H. yagouaroundi failed to reach the minimum 20 records required for overlap analyses (Table 2). According to the frequency values from the literature review (Table 1), the most important prey in the small felid diet were mammals < 1 kg, birds, and Salvator merianae (Duméril & Bibron, 1839), while the P. concolor diet was dominated by Cuniculus paca (Linnaeus, 1766), Dasypus spp., and Nasua nasua (Linnaeus, 1766).

Table 2
Total number of records of Herpailurus yagouaroundi, Leopardus guttulus, L. pardalis, L. wiedii and Puma concolor, and their probable prey taxa, obtained from May 2018-July 2019, with camera traps in an Atlantic Forest area located in the Treviso municipality, southern Santa Catarina state, Brazil.

Species of felines and prey taxa	Number of records
	Felines	Prey
Leopardus guttulus	89
L. wiedii	73
Puma concolor	62
L. pardalis	28
Herpailurus yagouaroundi	3
Birds		2,502
Salvator merianae		250
Dasypus spp.		240
Cuniculus paca		83
Mammals < 1 kg		70
Nasua nasua		46

Puma concolor, L. pardalis, and L. wiedii showed high circadian rhythm overlap with their respective main prey, with no significant differences observed (p > 0.05) between predator and prey activity rhythms (Table 3, Fig. 2): P. concolor versus C. paca (∆ OT = 0.8; BOOTStrap = 0.68-0.89; W = 2.29, p = 0.32), L. pardalis versus small mammals (∆ OT = 0.84; BOOTStrap = 0.63-0.85; W = 2.83, p = 0.24), and L. wiedii versus small mammals (∆ OT = 0.83; BOOTStrap = 0.69-0.89; W = 3.64, p = 0.16). Conversely, overlap values for the other prey of these three felids were moderate or low, with significant differences in activity rhythms (p < 0.05; Table 3, Fig. 2). Leopardus guttulus showed moderate overlap with its first and second ranked prey (mammals < 1 kg and birds, respectively) and low overlap with the third (S. merianae), with significant differences observed in all cases (p < 0.05; Table 3, Fig. 2).

Table 3
Overlap of the circadian rhythms of Leopardus guttulus, L. pardalis, L. wiedii and Puma concolor, and their preferential prey, in an Atlantic Forest area located in the Treviso municipality, southern Santa Catarina state, Brazil. Where: Δ OT = the most robust overlap coefficient values calculated by Overlap True, BOOTStrap = range of overlap values, W = Mardia-Watson-Wheeler test. p < 0.05 = different circadian rhythms. Values in bold demonstrate the absence of a significant difference using the Mardia-Watson-Wheeler test.

Preferential prey	Statistical parameters / Feline species
	∆ OT				BOOTStrap				W (p)
	P. concolor	L. pardalis	L. wiedii	L. guttulus	P. concolor	L. pardalis	L. wiedii	L. guttulus	P. concolor	L. pardalis	L. wiedii	L. guttulus
Cuniculus paca	0.8				0.68-0.89				2.29 (=0.32)
Dasypus spp.	0.73				0.61-0.81				12.24 (<0.05)
Nasua nasua	0.64				0.46-0.75				24.33 (<0.05)
Mammals < 1 kg		0.84	0.83	0.61		0.63-0.85	0.69-0.89	0.47-0.67		2.83 (=0.24)	3.64 (=0.16)	45.70 (<0.05)
Birds		0.55	0.47	0.76		0.36-0.65	0.29-0.48	0.66-0.81		22.47 (<0.05)	92.19 (<0.05)	22.04 (<0.05)
Salvator merianae		0.30		0.46		0.08-0.40		0.29-0.53		22.85 (<0.05)		35.58 (<0.05)

Figure 2
Overlay of circadian rhythms of Puma concolor, Leopardus pardalis, L. wiedii and L. guttulus, and their preferential prey (shaded polygons under the curves), in an Atlantic Forest area located in the Treviso municipality, southern Santa Catarina state, Brazil. ∆ = Overlap True (p < 0.05); * = absence of significant overlap (p > 0.05).

DISCUSSION

For the larger species-P. concolor and L. pardalis-the hypothesis was confirmed: their circadian rhythms overlap with those of their most common prey, defined by the bottom-up effect. This corroborates expected patterns for large Neotropical carnivores (Vallejo-Vargas et al. 2022) and provides a double advantage: increasing food acquisition while reducing foraging energy expenditure (Jenny and Zuberbuhler 2005).

The behavior of the smallest species, L. guttulus, also followed the predicted hypothesis, as it lacked high overlap with analyzed prey-a characteristic of the top-down effect (Vallejo-Vargas et al. 2022). In this scenario, the presence of larger predators forces smaller species to use less advantageous periods, resulting in the cathemeral behavior recorded here and in previous studies (Pratas-Santiago et al. 2016, Dias et al. 2018, Marinho et al. 2018, 2020, García-R et al. 2019). Notably, many predation events occur while prey are at rest (Harmsen et al. 2011, Emsens et al. 2013), outside their peak activity hours, as felids are both generalists and opportunists (Pratas-Santiago et al. 2016, Marinho et al. 2018, 2020, Santos et al. 2019).

Lastly, L. wiedii exhibited unexpected behavior; despite its small size, it showed patterns similar to larger species, with high overlap with its most common prey. The margay’s primary use of the arboreal stratum (Oliveira et al. 2010) may allow spatial segregation, avoiding encounters with larger terrestrial species. Thus, L. wiedii deviates from the expected top-down effect, following a bottom-up pattern that warrants more in-depth investigation.

ACKNOWLEDGMENTS

We thank Pousada Santo Antônio and Instituto Alouatta for study site access. We are also grateful to the two anonymous reviewers, whose suggestions helped refine this manuscript.

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ADDITIONAL NOTES

• ZooBank register
  https://zoobank.org/93308468-C851-4067-AB95-0D11F3B49AED
• Data Availability Statement
  Datasets generated or analyzed in this study are available from the corresponding author on reasonable request.
• Funding
  This study was supported by Universidade do Extremo Sul Catarinense, Fundação de Amparo à Pesquisa e Inovação de Santa Catarina (FAPESC 2022TR1975); and Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES Financial Code 001).
• Ethical Statement
  Not applicable
• AI Statement
  No artificial intelligence tools were used in the preparation of this manuscript.
• How to cite this article
  Mottin V, Moraes AGF, Bogoni JA, Castilho PV, Carvalho F, Graipel ME, Ribeiro M, Miranda JMD, Zocche JJ (2026) Circadian rhythm overlap between felids (Carnivora: Felidae) and their prey in a Subtropical Atlantic Forest protected area. Zoologia 43: e25020. https://doi.org/10.1590/S1984-4689.v43.e25020
• Published by
  Sociedade Brasileira de Zoologia at Scientific Electronic Library Online - https://www.scielo.br/zool

Edited by

• Editorial responsibility
  Jorge Salazar-Bravo

Data availability

Datasets generated or analyzed in this study are available from the corresponding author on reasonable request.

Publication Dates

• Publication in this collection
  12 June 2026
• Date of issue
  2026

History

• Received
  13 Mar 2025
• Accepted
  17 Nov 2025