Ground Nesting Behavior of Aotus griseimembra: Rare Terrestrial Evidence in A Strictly Arboreal Species

Abstract

We report a novel observation of ground nesting behavior in a couple of Aotus griseimembra within a successional inter-Andean Forest patch of Colombia. This behavior, previously unrecorded for strictly arboreal primates of the Genus Aotus, challenges conventional understanding. The monkeys exhibited typical species actions but sought refuge on the ground, possibly influenced by habitat alterations. Their visits to the ground sleeping site were monitored and confirmed the vulnerability to predators, competitors in the forest patch. These findings call the attention for further research into the response strategies of neotropical primates to environmental stressors and habitat disturbance.

Full text

NOTES
Natural History
Mammalogy Notes 2020, 6(2), mn0113; https://doi.org/10.47603/mano.v10n2.433 1/7
Mammalogy Notes ISSN 2382-3704
https://doi.org/10.47603/mano.v10n2.433
Ground Nesting Behavior of Aotus griseimembra: Rare
Terrestrial Evidence in A Strictly Arboreal Species
Julián Arango-Lozano1* Karime Angarita-Corzo2, Jose Julio-Guzmán3, Camilo
Ernesto Angarita-Yanes4, Sebastián O. Montilla5
1 Maestría en Ciencias Biológicas, Universidad de Caldas, Calle 65 #26-10, 170001, Manizales, Colombia.
2 Escuela de Medicina Veterinaria. Facultad de Ciencias Agrarias. Universidad de Antioquia, Carrera 75 # 65-87, 050034,
Medellín, Colombia.
3 Grupo de investigación en biodiversidad del caribe colombiano, Universidad del Atlántico, UA, 081001, Carrera 30 # 8-49,
Puerto Colombia, Colombia
4 Biólogo, Investigador independiente
5 Laboratorio de Ecología de Bosques Tropicales y Primatología; Universidad de los Andes, Bogotá D.C., Colombia
* Correspondencia: arangolozanoj1@gmail.com
Resumen
Reportamos una observación novedosa del comportamiento de anidación en el suelo en
una pareja de Aotus griseimembra, dentro de un parche sucesional de bosque interandino
en Colombia. Este comportamiento, previamente no registrado para primates
estrictamente arbóreos del género Aotus, desafía la comprensión convencional. Los monos
exhibieron acciones típicas de la especie, pero buscaron refugio en el suelo, posiblemente
influenciados por alteraciones del hábitat. Sus visitas al sitio de descanso en el suelo
fueron monitoreadas y confirmaron la vulnerabilidad a depredadores y competidores en
el parche de bosque. Estos hallazgos destacan la necesidad de nuevas investigaciones
sobre las estrategias de respuesta de los primates neotropicales a los factores estresantes
ambientales y la perturbación del hábitat
Palabras clave: Fragmentación del bosque, Monos nocturnos del Caribe, Primates neotropicales.
Abstract
We report a novel observation of ground nesting behavior in a couple of Aotus
griseimembra within a successional inter-Andean Forest patch of Colombia. This behavior,
previously unrecorded for strictly arboreal primates of the Genus Aotus, challenges
conventional understanding. The monkeys exhibited typical species actions but sought
refuge on the ground, possibly influenced by habitat alterations. Their visits to the ground
sleeping site were monitored and confirmed the vulnerability to predators, competitors in
the forest patch. These findings call the attention for further research into the response
strategies of neotropical primates to environmental stressors and habitat disturbance.
Keywords: Caribbean Owl monkey, Forest fragmentation, Neotropical primates.

Mammalogy Notes 2020, 6(2), mn0113; https://doi.org/10.47603/mano.v10n2.433 2/7
Neotropical primates are one of the most threatened mammal groups due to their ecology
restricted to forest habitats (Alfaro et al., 2015; Benchimol & Peres 2014; Túnez et al., 2021).
These populations in central and South America are declining as their ideal habitats are
reduced by the expansion of agricultural frontiers, deforestation, pollution, and climate
change (de Almeida et al., 2017; Estrada et al., 2017; Carvalho et al., 2019). In response to
these environmental stressors, Neotropical primates may exhibit behavioural changes in
response to these new challenges (Jung et al., 2015; Morelos-Juárez et al. 2015; Corrêa et al.,
2018). These changes include alterations in foraging strategies, social structures, and
ranging patterns to cope with habitat fragmentation and resource scarcity (Schwitzer et al.,
2011; Estrada et al., 2012; de Almeida et al., 2017; Ramsay et al., 2023). Additionally, shifts in
reproductive behaviors and increased tolerance to human presence show an adaptive
response to habitat disturbance (Tokuda et al. 2018; Mancini et al. 2023). While vulnerability
increases over time due to habitat loss, some resilient species as Sapajus nigritus and
Alouatta guariba clamitans underscore their capacity to persist in human-altered
landscapes (Corrêa et al., 2018; Tokuda et al. 2018).
Terrestrial behavior, recognized as those activities performed by primates on the ground
rather than in trees, has been poorly studied in the Neotropics because most monkey 
species are highly associated with the forest canopy (Campbell et al., 2005; Eppley et al.,
2016; Souza-Alvez et al., 2019; Monteza-Moreno et al., 2020; Eppley et al., 2022). However,
some patterns have been detailed, such as: (i) use of stones and sticks on the ground to
open shelled fruits and extract insects in Cebus spp. (Waga et al., 2006), use of waterholes
during dry season (Freese 1978). (ii) Foraging behavior at terrestrial salt licks and
waterholes in Ateles spp. and Alouatta spp. (Campbell 2005; Ferrari 2008; Link et al., 2010,
2011). (iii) Ground vocalizations near highly degraded forests by Aotus spp. (Shanee &
Shanee 2011). And finally, and more generally, (iv) road crossings by various groups of
neotropical primates, that may result in roadkills (Praill et al., 2023). However, no nesting
or sleeping sites on the ground have been recorded before for strictly arboreal primates
of the genus Aotus. Here we report an abnormal sleeping site behavior on the ground by
Aotus griseimembra in the inter-Andean basin of Colombia.
During field trips from April 2 to May 2, 2024, we identified an adult pair of Aotus
griseimembra in the Santa Lucia rural area, vereda La Rochela, Simacota, Santander, within
the inter-Andean basin of the Mid-Magdalena River Basin, Colombia (latitude: 6°44'7.43" N,
longitude: 73°52'14.34" W). These observations occurred in a successional forest patch
surrounded by mixed grasslands and two nearby roadways at an elevation of 100 m.
The first observation occurred on April 4, 2024, at 09:59 h, when we detected the male at
the forest boundaries on the ground, possibly altered by human presence. On April 8, 2024,
at 12:47 h and 16:10 h, we observed the pair heading towards a sleeping site based on a
tree of the species Xilopia aromatica (Annonaceae). The animals were resting and covering
themselves with a mixed structure of sticks, leaves, and leaf litter on the ground, a type of
structure already documented in trees but different from the usual holes or cavities in
trees (Aquino & Encarnación 1986; Case 2013; Helenbrook et al., 2019). Throughout our
observations, we identified the male climbing in and out of the trees and returning to the
sleeping sites (possibly by the researcher’s presence); however, the female remained in
the refuge, a possible territorial display like other neotropical primates (Rutberg 1983;

Mammalogy Notes 2020, 6(2), mn0113; https://doi.org/10.47603/mano.v10n2.433 3/7
Janson, 1986; Spence-Aizenberg et al. 2023; Figure 1; video in Supplementary material,
Arango-Lozano 2024).
FIGURE 1. Detection site of: (a, b, c) the A. griseimembra ground sleeping site at the base of a Xilopia
aromatica tree, and the night monkey couple in the sleeping site. Arrows indicate the sleeping site.
(d and e) records via camera trap of the presence of Leopardus pardalis and Cebus versicolor,
respectively. Video is available in Mendeley data repositorium (Arango-Lozano 2024,
https://data.mendeley.com/datasets/j6tcm29gnw/1).
We conducted a total of nine separate excursions, excluding the initial encounter on April
4th when we set up camera traps to monitor the activity of the individuals at the sleeping
site. These excursions took place on April 5th, 8th, 9th, 14th, 17th, 21st, 22nd, and 23rd, and
May 2nd, between 17:00 and 24:00 hours. It is noteworthy that between April 9th and April
21st, there were no sightings of the pair, either at the sleeping site or on the camera trap
footage. However, on April 22nd at 19:30 hours, the pair was observed in the gallery forest
association, moving through the canopy of trees, but not near the ground sleeping site.
This observation was repeated on April 23rd at 19:15 hours.
Aotus griseimembra is a night monkey species highly associated to forest cover,
categorized as Vulnerable (VU) due to illegal trade, transformation, and degradation of
habitat (Defler, 2003; Mantilla-Meluk & Ortega 2011; Henao-Díaz et al.,2020; Montilla et al.,
2021). This night monkey inhabits from conserved to highly degraded forest patches in the
Caribbean and inter Andean valleys of Colombia and the western slopes of Northern
Venezuela always restricted below 1000 m in elevation (Shanee et al., 2023). Several authors
have called the attention about the A. griseimembra tolerance to inhabiting degraded
patches in Colombia, using the surrounding available resources for feeding and sleeping
site in the canopy trees (Garcés-Restrepo et al., 2016; Montilla et al., 2021, Shanee et al.,
2023). For A. griseimembra, however, this is the first evidence using of roosts other than
tree holes. It is also the first observation of ground-level sleeping sites for night monkeys.

Mammalogy Notes 2020, 6(2), mn0113; https://doi.org/10.47603/mano.v10n2.433 4/7
We are not sure what led this night monkey couple to use a terrestrial sleeping site.
However, with camera traps records and recognized traces we identified the presence of a
pair of Leopardus pardalis and a troop of Cebus versicolor (Figure 1d and 3e). The former
is a potential predator (Miranda et al., 2005; Bianchi & Mendes 2007; de Oliveira et al., 2009),
and the latter is a potential resource competitor (Link et al., 2010; Marsh et al., 2016). This
could indicate that resources such as fruit or leaves for primates are not scarce in the
mixed matrix of grasslands and remnant successional forest. Additionally, the persistence
of this nest may be due to the behavior of the Aotus genus, which is known to maintain
multiple sleeping sites within their home range (Bustamante-Manrique et al., 2021; Montilla
et al., 2021). It is possible that the ground sleeping site served as a previously established
refuge for the night monkey pair, which they continued to visit after the structure collapsed
from the tree due to unknown circumstances.
The proximity to the ground heightens the vulnerability of these primates to predators
such as wild cats, both during the day and night. Additionally, it increases the likelihood of
encounters with domestic cats and dogs, thereby exposing them to zoonotic diseases like
Toxoplasma gondii, as observed in other primates, including those of the Aotus genus
(Gyimesi et al., 2006). Despite this unexpected behavior, the couple exhibited typical
characteristics of their species, displaying heightened responsiveness to stimuli, and
seeking refuge in canopy cover when humans were present (Shanee et al., 2023). However,
due to the rarity of this occurrence, we are motivated to maintain non-invasive surveillance
of the couple using camera traps to gather more comprehensive data. Currently, we are
seeking permission from the owners of private properties surrounding the forest patch
where the encounter took place to monitor the couple's movements and ascertain whether
they are attempting to access other nearby forest remnants.
ACKNOWLEDGMENTS
The authors thank Proyecto Troncal Magdalena 1 for the support of field logistics.
REFERENCES
Arango-Lozano J. 2024, “GROUND NESTING BEHAVIOR OF Aotus griseimembra: RARE TERRESTRIAL
EVIDENCE IN A STRICTLY ARBOREAL SPECIES”, Mendeley Data, V1, doi: 10.17632/j6tcm29gnw.1
available at: https://data.mendeley.com/datasets/j6tcm29gnw/1
Alfaro JWL, Cortés-Ortiz L, Di Fiore A, Boubli JP. 2015. Comparative biogeography of Neotropical
primates. Molecular Phylogenetics and Evolution 82:518–529.
https://doi.org/10.1016/j.ympev.2014.09.027
Aquino R, Encarnación F. 1986. Characteristics and use of sleeping sites in Aotus (Cebidae: Primates)
in the Amazon lowlands of Peru. American Journal of Primatology 11(4):319–331.
https://doi.org/10.1002/ajp.1350110403
Benchimol M, Peres CA. 2014. Predicting primate local extinctions within “real‐world” forest
fragments: a pan‐neotropical analysis. American Journal of Primatology 76(3):289–302.
https://doi.org/10.1002/ajp.22233
Bianchi RDC, Mendes SL. 2007. Ocelot (Leopardus pardalis) predation on primates in Caratinga
Biological Station, southeast Brazil. American Journal of Primatology: Official Journal of the
American Society of Primatologists 69(10):1173–1178. https://doi.org/10.1002/ajp.20415

Mammalogy Notes 2020, 6(2), mn0113; https://doi.org/10.47603/mano.v10n2.433 5/7
Bustamante-Manrique S, Botero-Henao N, Castaño JH, Link A. 2021. Activity budget, home range and
diet of the Colombian night monkey (Aotus lemurinus) in peri-urban forest fragments.
Primates 62(3):529–536. https://doi.org/10.1007/s10329-021-00895-w
Campbell CJ, Aureli F, Chapman CA, Ramos-Fernández G, Matthews K, Russo SE, Vick L. 2005.
Terrestrial behavior of Ateles spp. International Journal of Primatology 26:1039–1051.
https://doi.org/10.1007/s10764-005-6457-1
Case L. 2013. Understanding Behavior and Nest Box Usage in Three Species of Owl Monkeys: Azara's
Owl Monkey (Aotus Azarai), Spix's Owl Monkey (A. Vociferans) and Nancy's Owl Monkey (A.
Nancymaae). (Master's Thesis). Retrieved from:
https://digital.library.txst.edu/server/api/core/bitstreams/d3202538-e42d-4778-8bd1-
0914ef624dba/content
Carvalho JS, Graham B, Rebelo H, Bocksberger G, Meyer CF, Wich S, Kühl HS. 2019. A global risk
assessment of primates under climate and land use/cover scenarios. Global Change Biology
25(9):3163–3178. https://doi.org/10.1111/gcb.14671
Corrêa FM, Chaves ÓM, Printes RC, Romanowski HP. 2018. Surviving in the urban–rural interface:
Feeding and ranging behavior of brown howlers (Alouatta guariba clamitans) in an urban
fragment in southern Brazil. American Journal of Primatology 80(6):e22865.
https://doi.org/10.1002/ajp.22865
de Almeida-Rocha JM, Peres CA, Oliveira LC. 2017. Primate responses to anthropogenic habitat
disturbance: a pantropical meta-analysis. Biological Conservation 215:30–38.
https://doi.org/10.1016/j.biocon.2017.08.018
Defler TR. 2003. Primates de Colombia Conservación Internacional, serie de guías tropicales de
campo 4. Conservación Internacional, Bogotá DC, 543pp.
de Oliveira Calleia F, Rohe F, Gordo M. 2009. Hunting strategy of the margay (Leopardus wiedii) to
attract the wild pied tamarin (Saguinus bicolor). Neotropical Primates 16(1):32–34.
https://doi.org/10.1896/044.016.0107
Eppley TM, Donati G, Ganzhorn JU. 2016. Determinants of terrestrial feeding in an arboreal primate:
The case of the southern bamboo lemur (Hapalemur meridionalis). American Journal of
Physical Anthropology 161(2):328–342. https://doi.org/10.1002/ajpa.23034
Eppley TM, Hoeks S, Chapman CA, Ganzhorn JU, Hall K, Owen MA, Santini L. 2022. Factors influencing
terrestriality in primates of the Americas and Madagascar. Proceedings of the National
Academy of Sciences 119(42):e2121105119. https://doi.org/10.1073/pnas.2121105119
Estrada A, Raboy BE, Oliveira LC. 2012. Agroecosystems and primate conservation in the tropics: a
review. American journal of primatology 74(8):696–711. https://doi.org/10.1002/ajp.22033
Estrada A, Garber PA, Rylands AB, Roos C, Fernandez-Duque E, Di Fiore A, Li B. 2017. Impending
extinction crisis of the world’s primates: Why primates matter. Science advances
3(1):e1600946. doi: 10.1126/sciadv.1600946
Ferrari SF, Veiga LM, Urbani B. 2008. Geophagy in New World monkeys (Platyrrhini): ecological and
geographic patterns. Folia Primatologica 79(5):402–415. https://doi.org/10.1159/000141901
Freese CH. 1978. The behaviour of white faced capuchins (Cebus capucinus) at a dry-season
waterhole. Primates 19:275–286. https://doi.org/10.1007/BF02382797
Garcés-Restrepo MF, Quintero Ángel A, Cuéllar N, Giraldo A. 2016. Mammal Diversity in an Area with
Relicts of Dry Forest in the Mid-Magdalena Valley (Caldas, Colombia). Revista de Ciencias
20(2). https://core.ac.uk/download/pdf/267165753.pdf

Mammalogy Notes 2020, 6(2), mn0113; https://doi.org/10.47603/mano.v10n2.433 6/7
Gyimesi ZS, Lappin MR, Dubey JP. 2006. Application of assays for the diagnosis of toxoplasmosis in
a colony of woolly monkeys (Lagothrix lagotricha). Journal of Zoo and Wildlife Medicine
37(3):276–280. https://doi.org/10.1638/05-018.1
Henao-Díaz F, Stevenson P, Carretero-Pinzón X, Castillo-Ayala C, Pacheco J, Defer T, García Villalba J,
Caro D, Link A, Moreno M, Palacios E, Duque N, Soto-Calderón I, Soto L, Velásquez-Tibata J,
Olaya-Rodríguez M, Noguera-Urbano E, Valencia L. 2020. Atlas de la biodiversidad de
Colombia. Primates. Mejores modelos con el apoyo de expertos. Atlas de la Biodiversidad
de Colombia. Instituto de Investigación de Recursos Biológicos Alexander von Humboldt,
Bogotá D. C., Colombia. 51 pp.
Helenbrook WD, Wilkinson ML, Suarez JA. 2019. Habitat use, fruit consumption, and population
density of the black-headed night monkey, Aotus nigriceps, in southeastern Peru. Acta
Amazonica 50(1):37–43. https://doi.org/10.1590/1809-4392201900172
Janson CH. 1986. The mating system as a determinant of social evolution in capuchinmonkeys
(Cebus). In: Else J, Lee PC (eds) Proceedings of the Xth International Congress of Primatology.
Cambridge University Press, Cambridge, pp 169–179.
Jung L, Mourthe I, Grelle CE, Strier KB, Boubli JP. 2015. Effects of local habitat variation on the
behavioral ecology of two sympatric groups of brown howler monkey (Alouatta clamitans).
PLoS One 10(7):e0129789. https://doi.org/10.1371/journal.pone.0129789
Link A, De Luna AG, Alfonso F, Giraldo-Beltran P, Ramirez F. 2010. Initial effects of fragmentation on
the density of three neotropical primate species in two lowland forests of Colombia.
Endangered Species Research 13(1):41–50. https://doi.org/10.3354/esr00312
Link A, De Luna A G, Arango R, Diaz MC. 2011. Geophagy in brown spider monkeys (Ateles hybridus)
in a lowland tropical rainforest in Colombia. Folia primatologica 82(1):25–32.
https://doi.org/10.1159/000326056
Marsh C, Link A, King-Bailey G, Donati G. 2016. Effects of fragment and vegetation structure on the
population abundance of Ateles hybridus, Alouatta seniculus and Cebus albifrons in
Magdalena Valley, Colombia. Folia Primatologica 87(1):17–30.
https://doi.org/10.1159/000443929
Mancini G, Benítez-López A, Di Marco M, Pacifici M, Rondinini C, Santini L. 2023. Synergistic effects of
habitat fragmentation and hunting on the extinction risk of neotropical primates.
Biodiversity and Conservation 32(8):2655–2669. https://doi.org/10.1007/s10531-023-02623-w
Mantilla-Meluk H, Jiménez-Ortega AM. 2011. Revisando el estatus taxonómico y la partición ecológica
de los monos nocturnos del género Aotus en el noroccidente de Colombia, con notas sobre
Aotus zonalis Goldman, 1914. Revista Biodiversidad Neotropical 1(1 Ene-Jun):28–37.
https://doi.org/10.18636/bioneotropical.v1i1.26
Miranda J, Bernardi IP, Abreu KC, Passos FC. 2005. Predation on Alouatta guariba clamitans Cabrera
(Primates, Atelidae) by Leopardus pardalis (Linnaeus)(Carnivora, Felidae). Revista Brasileira
de Zoologia 22:793–795. https://doi.org/10.1590/S0101-81752005000300043
Monteza-Moreno CM, Crofoot MC, Grote MN, Jansen PA. 2020. Increased terrestriality in a Neotropical
primate living on islands with reduced predation risk. Journal of human evolution 143: 02768.
https://doi.org/10.1016/j.jhevol.2020.102768
Montilla SO, Mopán-Chilito AM, Murcia LNS, Triana JDM, Ruiz OMC, Montoya-Cepeda J, Link A. 2021.
Activity patterns, diet and home range of night monkeys (Aotus griseimembra and Aotus
lemurinus) in tropical lowland and mountain forests of central Colombia. International
Journal of Primatology 42:130–153. https://doi.org/10.1007/s10764-020-00192-1

Mammalogy Notes 2020, 6(2), mn0113; https://doi.org/10.47603/mano.v10n2.433 7/7
Morelos-Juarez C, Tapia A, Conde G, Peck M. 2015. Diet of the critically endangered spider monkey
(Ateles fusciceps fusciceps) in the Ecuadorian Chocó: Conflict between primates and loggers
over fruiting tree species. PeerJ Preprints 3: e1574V1.
https://doi.org/10.7287/peerj.preprints.1574v1
Praill LC, Eppley TM, Shanee S, Cunneyworth PM, Abra FD, Allgas N, Svensson MS. 2023. Road
Infrastructure and Primate Conservation: Introducing the Global Primate Roadkill Database.
Animals 13(10):1692. https://doi.org/10.3390/ani13101692
Ramsay MS, Mercado Malabet F, Klass K, Ahmed T, Muzaffar S. 2023. Consequences of Habitat Loss
and Fragmentation for Primate Behavioral Ecology. In Primates in anthropogenic
landscapes: Exploring primate Behavioural flexibility across human contexts (pp. 9–28).
Cham: Springer International Publishing.
Rutberg AT. 1983. The evolution of monogamy in primates. Journal of Theoretical Biology 104(1):93–
112. https://doi.org/10.1016/0022-5193(83)90403-4
Shanee S, Shanee N. 2011. Activity budget and behavioural patterns of free-ranging yellow-tailed
woolly monkeys Oreonax flavicauda (Mammalia: Primates), at La Esperanza, northeastern
Peru. Contributions to Zoology 80(4):269–277. https://doi.org/10.1163/18759866-08004004
Shanee S. 2023. Threats and conservation of owl monkeys (Aotus spp.) in the Andes. In Owl Monkeys:
Biology, Adaptive Radiation, and Behavioral Ecology of the Only Nocturnal Primate in the
Americas (pp. 649-671). Cham: Springer International Publishing.
https://doi.org/10.1007/978-3-031-13555-2_22
Schwitzer C, Glatt L, Nekaris KAI, Ganzhorn JU. 2011. Responses of animals to habitat alteration: an
overview focussing on primates. Endangered Species Research 14(1):31–38.
Souza-Alves JP, Mourthe I, Hilário RR, Bicca-Marques JC, Rehg J, Gestich CC, … & Barnett AA. 2019.
Terrestrial behavior in titi monkeys (Callicebus, Cheracebus, and Plecturocebus): Potential
correlates, patterns, and differences between genera. International Journal of Primatology
40:553-572.
Spence-Aizenberg A, García de la Chica A, Evans S, Fernandez-Duque E. 2023. Communication in owl
monkeys. In Owl Monkeys: Biology, Adaptive Radiation, and Behavioral Ecology of the Only
Nocturnal Primate in the Americas (pp. 497–533). Cham: Springer International Publishing.
https://doi.org/10.1007/978-3-031-13555-2_17
Tokuda M, Martins MM, Izar P. 2018. Socio-genetic correlates of unbiased sex dispersal in a
population of black capuchin monkeys (Sapajus nigritus). Acta Ethologica 21,1–11 (2018).
https://doi.org/10.1007/s10211-017-0277-0
Túnez JI, Nardelli M, Ibañez EA, Peralta DM, Byrne MS. 2021. A review of the conservation status of
Neotropical mammals. Molecular Ecology and Conservation Genetics of Neotropical
Mammals 11–33. https://doi.org/10.1007/978-3-030-65606-5_2
Waga IC, AK Dacier, PS Pinha, MCH Tavares 2006. Spontaneous tool use by wild capuchin monkeys
(Cebus libidinosus) in the Cerrado. Folia Primatológica 77:337–344.
https://doi.org/10.1159/000093698
Editor: Camilo A. Calderón-Acevedo
Received: 2024-05-16
Reviewed: 2024-05-20
Accepted: 2024-08-10
Published: 2024-10-22