First record of a breeding site of Ara militaris mexicanus (Ridgway, 1915), Military Macaw (Psittacidae), in the Alto Balsas Poblano Region, Mexico

. We report a new breeding site of Ara militaris mexicanus (Ridway, 1915), Military Macaw, in the state of Puebla in the Alto Balsas river basin, Mexico. The breeding site is in the Cañón Grande area along the Nexapa River. Observed temporality and breeding behaviors agree with the recorded breeding season of this species in Central Mexico. Notably, the region has no current designation as a protected natural area for conservation. We recommend collaboration with the local community to initiate efforts towards designate the Alto Balsas river basin as a protected natural area


INTRODUCTION
Ara militaris (Linnaeus, 1766), Military Macaw, has a wide and disjunct distribution, ranging from northern Mexico to South America.Its primary habitat includes ravines, canyons and slopes within tropical dry forests, including deciduous, semideciduous forest, and tropical oak forest (Rivera-Ortiz et al. 2013).According to the geographical distribution, three subspecies of the Ara militaris complex are recognized: A. m. militaris (Linnaeus, 1766), occurring in Venezuela, Colombia, Ecuador, and Peru;A. m. bolivianus (Reichenow, 1908), distributed in Bolivia and Argentina;and A. m. mexicanus (Ridway, 1915), found in Mexico (Juniper and Parr 1998;Dehasque 2016).Two primary lineages are identified within the complex: A. m. mexicanus and A. m. militaris-A.m. bolivianus.Historical isolation between the ancestral populations of these lineages is suggested by the discontinuous geographical distribution of this species (Rivera-Ortíz et al. 2023).Rivera-Ortíz et al. (2023) proposed elevating the subspecies A. m. mexicanus to a full species, as A. mexicanus, although conclusive evidence is lacking.Therefore, A. m. mexicanus should be retained as a subspecies of A. militaris.For these reasons, we attribute the breeding population in this note to the subspecies A. m. mexicanus.This subspecies has an approximate length of 67 cm and displays body plumage ranging from olive to bright green, with a matte green tone on the nape (Howell and Webb 1995;Macedo 2007).The flight feathers are blue on the upper part and olive-yellow on the interior part.The mantle is pale blue, with the upper tail covert feathers red and the lower cover feathers turquoise blue (Gaucin 2000).The forehead exhibits a crimson red tone, the auricular region varies from white to pink, featuring lines of small black and red feathers, and the beak is black (Juniper and Parr 1998;Dehasque 2016;Ortega 2024; Figure 1).Ara militaris undertakes altitudinal movements that range from 600 to 2200 m in elevation, and it has been suggested that these movements are influenced by variations in the abundance of food resources (Contreras-González et al. 2009) and the northern region of western Mexico by cooler temperatures in fall and winter (Nocedal et al. 2006).Its diet encompasses fruits, seeds, leaves, buds, and stems, making it a highly specialized species, as it consumes 10-23% of the available plant resources (Iñigo-Elías 1999;Contreras-González et al. 2009).This species exhibits gregarious nesting behavior, typically selecting cavities in vertical karstic walls or tree cavities with a diameter at breast height greater than 60 cm (Carreón 1997;Gómez 2004;Nocedal et al. 2006;Rivera-Ortíz et al. 2008, 2013).

Historically in western
Before 1999, the state of Puebla was not considered to be within the historical or potential distribution of A. militaris in Mexico, despite some unverified reports in the southern portion of Puebla (Arizmendi and Valiente-Banuet 1999). However, between 2008and 2010, Salazar et al. (2010) documented the first sightings of the species in the state, with an average of six individuals recorded in Cerro Colorado Tehuacán, Puebla (150 km from our study area).They also recorded macaws to the south of Puebla, bordering the state of Oaxaca, in the vicinity of Teotitlán de Flores Magón, about 180 km from our study area (Salazar et al. 2010) 2018) conducted a survey on A. militaris habitat conservation in the Alto Balsas basin in Guerrero, estimating an abundance of 100 individuals in the region (Naturam Sequi 2015, 2016).They also identified probable roosting and feeding areas near the locality of Santa María Cohetzala, Puebla, approximately 20 km from the Papalutla breeding colony (Figure 2).
In this study, we present, for the first time, evidence of the presence and establishment of a new breeding population of A. militaris in the state of Puebla within the Alto Balsas basin river.In addition, we include data spanning from 2016 to 2023, accompanied by detailed observations of their natural history.

METHODS
Our records were made in Santa María Cohetzala, located in the Alto Balsas basin in the southwestern portion of the state of Puebla, Mexico.The area is characterized by having rugged terrain with substantial elevated landforms and extensive ravines, intersected by the Nexapa and Atoyac rivers (Figure 2).The elevation varies from 700 to 1900 m a.s.l.The region has a warm subhumid climate, with an average annual temperature of 26 °C and an annual precipitation ranging of 600-800 mm (García 1998).The predominant vegetation type is tropical dry forest, including deciduous and semi-deciduous woodlands (Martínez-Gordillo et al. 1997) (Figure 3).
Five exploration and observation routes were surveyed, using existing trails in the tropical dry forest on Tecaballo Mountain and in the subdecidous tropical forest along the Nexapa River, each route with an average duration of 8 h.The total area explored was 5 km 2 , which included 3.5 km 2 in the deciduous forest and 1.5 km 2 in the subdeciduos forest.The routes were chosen based on previous studies by Naturam Sequi (2015Sequi ( , 2016) ) and local residents, who reported the presence of A. militaris in the area.The first route, in Peña del Sol, was surveyed on 12 December 2016 and had a length of 6 km.The second route, in Peña del Sol and Peña del Reloj in the Cañón Grande on Tecaballo Mountain, was surveyed on 12 February 2020 and had a route length of 8 km.The third route was surveyed on 6-7 March 2022 in Peña del Reloj and Cañón Grande and had a route length of 9 km.The fourth route, towards the upper part of the Cañón Grande in the Tecaballo Mountain, was surveyed on 5-7 June 2023, with the aim of locating viewpoints that would provide better visibility of the canyon walls and A. militaris individuals, had a route length of 12 km.Finally, the fifth route, in Peña del Sol, was surveyed on 21-23 May 2024 and had a route length of 10 km, and the search was specifically for cavities used as nests.In each route there were five observers and two local guides (Table 1).During the exploration trips, natural history data were recorded, such as vegetation type, geographical coordinates, and the activities of A. militaris individuals, including foraging and breeding behavior.Photographic evidence was captured.
Identification.The species was morphologically identified through observations, photographic documentation, vocalizations, direct sightings (Figure 1), and verification by specialists in Psittacidae who are co-authors (FARO, LDVR, and AMCG).Remarks.The first record of the macaws was from 12 December 2016, at 10:00 h by one of us (LDVR), who noted three pairs flying over the Nexapa River and perching on the emerging trees towards the northwest of the Peña del Sol site.It is possible that the birds were heading towards the Cañón Xicuhuetztlán in Papalutla, Guerrero.On 12 February 2020, at 14:00 h, 10 macaws were observed flying in circular movements over the Peña del Sol site along the course of the Nexapa River.They were heading towards Peña del Reloj, where they were seen entering the Cañón Grande (Table 1).Only their vocalizations could be heard, as the canyon was inaccessible to us, preventing further data recording.On 22 March 2022, at 12:00 h, four individuals of were observed flying over the course of the Nexapa River at the entrance of the Cañón Grande.They were feeding on the fruit of the Bonete Tree, Jacaratia mexicana A.DC. for approximately 30 min.After eating, two of these macaws flew and entered the Cañón Grande, while the other two macaws perched on a Parota Tree, Enterolobium cyclocarpum (Jacq.)Griseb.and engaged in courtship behavior for 5 min.Subsequently, they returned to the Bonete Tree, continued courtship for a couple of minutes, and were observed mating (Figure 1).At 13:00 h, they also entered the Cañón Grande.Later, at 16:00 h, a flock of 16 macaws was recorded, heading towards the Cañón Grande.On the same day, a walk through the canyon, six individuals were observed, and their vocalizations heard.On 7 March 2022, at 06:00 h, nine pairs of A. militaris were recorded as they exited the Cañón Grande.They were heading northwest, making circular movements over the Nexapa River, and then headed towards the Tecaballo Mountain (Table 1).Once again, the vocalizations of the macaws were heard from within the canyon.
On 6 June 2023, 16 macaws were recorded entering the Cañón Grande between 17:00 h and 19:30 h.They engaged in flights within the canyon, with some perching on the Cuajiote Colorado Tree, Bursera morelensis Ramírez and the Cacaloxuchil Tree, Plumeria rubra L. Others macaws entered the cavities in the walls of the Cañón Grande.On 7 June 2023, at 5:30 h, macaw vocalizations were heard within the canyon.At 6:15 h, a flock of 20 individuals emerged, spreading out over the course of the Nexapa River (Table 1).Cavities used to roost by the macaws were marked with guano on the walls of the Cañón Grande and identified, although their use for nesting were not identified.On 22 May 2024, two cavities occupied by A. militaris with reproductive activity were recorded.We observed that one individual, a male, arrived to feed another individual, the female, in the cavity.This occurred in both cavities throughout the day, indicating egg incubation (Figure 1).

DISCUSSION
Our results on the natural history and breeding biology of the Ana militaris mexicanus provide important information about the first record a breeding site of this species in the state of Puebla, Mexico.Breeding activities, including mating and courtship, strongly suggest that the Cañón Grande in the Peña del Reloj on Tecaballo Mountain represents a breeding site (Forshaw 2006).
In the semi-arid seasonal environments of central Mexico, the nesting season of A. m. mexicanus is known to start in March with courtship and mating, ending in late September with the first flight of fledglings (Rivera-Ortíz et al. 2008;Jiménez-Arcos et al. 2012;Vázquez-Reyes 2018).We recorded breeding activity in March, such as courtship and mating, as well as cavity use in June, which coincides with the patterns observed in the dry forests of central Mexico (Rivera-Ortíz et al. 2008;Jiménez-Arcos et al. 2012).The increase in the number of individuals in March likely corresponds to the onset of the breeding season (March-September) with the arrival of the first pairs.However, we note that our data are not consistent throughout the year, emphasizing the need for a permanent monitoring program of the population to document accurate patterns that may vary over time (Jiménez-Arcos et al. 2012).
Our observations in Santa María Cohetzala suggest that a metapopulation of A. m. mexicanus inhabits the Alto Balsas region, which is supported by field evidence from conservation projects (Naturam Sequi, 2015Sequi, , 2016) ) and ongoing monitoring data, which show fluctuating abundance between Papalutla and Santa María Cohetzala breeding sites and foraging areas in response to resource availability and even stochastic events.For instance, a census on 29 January 2024, recorded 46 individuals at the Xicuhuetztlán Canyon in Papalutla, while the Cañón Grande in Santa María Cohetzala had 25 individuals on 26 February 2024.Unfortunately, there was a forest fire near Cañón Grande on 12 March 2024 which disturbed nearby foraging areas.Subsequently, on 24 March 2024 at least 80 individuals were recorded in the roosting census at Xicuhuetztlán Canyon, likely due to macaws seeking refuge and foraging areas after the fire.This behavior aligns with the species' capacity for movement between nesting, resting, and foraging areas; this species can move up to 25 km per day, and it has an average home movement range of 87 km² (Juárez et al. 2012).
Our results may be found to be of importance in determining demographics and migratory movements between populations and for determining population connectivity through genetic and genomic analyses (landscape genetics) of the metapopulation (Ramírez-Albores 2016).Population genetics and phylogeography of A. m. mexicanus in the Sierra Madre Occidental and Sierra Madre del Sur show that these populations maintain genetic flow (Rivera-Ortíz et al. 2017;Rivera-Ortíz et al. 2023).The breeding site of A. m. mexicanus in Santa María Cohetzala is within the Important Bird Area (IBA) MX264 Papalutla-Tecaballo, as recognized by BirdLife International (2021).Furthermore, this IBA is acknowledged as within a Priority Region for Conservation in Guerrero, according to CONANP (2021).However, the distribution of this A. m. mexicanus population does not overlap with any natural protected area, despite its close proximity (37 km) to the Sierra de Huautla Biosphere Reserve, where A. m. mexicanus has been recorded at Rancho Viejo locality of Morelos (Urbina-Torres 2017).Regardless, these area designations are considered insufficient to reduce biodiversity loss in the region (Vázquez-Reyes 2018).This region is among those in which the least conservation efforts have been invested within Mexico.
Unfortunately, the breeding site of A. m. mexicanus in Santa María Cohetzala faced significant disturbances in 2016 and 2017 due to an uncontrolled wildfire, during which the State of Puebla deployed helicopters and helibuckets within the Cañón Grande (residents of Santa María Cohetzala pers.comm.).Despite such disturbance, the tropical dry forest in the area remains relatively unaffected, making it a potential area for A. m. mexicanus conservation.
Tecaballo Mountain, with its two breeding sites of A. m. mexicanus, one in Guerrero and one in Puebla, should be a focal point for conservation plans (Jiménez-Arcos et al. 2012).However, the conservation of the A. m. mexicanus population requires several important environmental policy actions and decisions.First, is necessary guarantee the protection and restoration of the forest habitats critical for foraging, roosting, and breeding of this species.However, this task is complicated by administrative complexities stemming from the macaw's distribution across Puebla and Guerrero states (Vázquez-Reyes 2018).Solid knowledge of the ecology of A. m. mexicanus in Santa María Cohetzala and on Tecaballo Mountain is of paramount importance.This will enable the formulation of effective conservation strategies in the region (De la Parra-Martínez et al. 2015).
Establishing biodiversity protection areas including two states has been successfully done in Mexico, as exemplified by the Tehuacán-Cuicatlán Biosphere Reserve which spans the states of Puebla and Oaxaca.This reserve has effectively allocated targeted funding for conservation focused on forest habitat of A. m. mexicanus.Indeed, A. m. mexicanus now acts as an umbrella species for biodiversity protection within the Tehuacán-Cuicatlán Biosphere Reserve (Vinculación Interdiciplinaria para el Desarrollo Ambiental y lo Social 2018).Again, coordinated efforts among Puebla, Guerrero, and the national environmental authorities are required.Fortunately, there important advances have already been made in documenting endemic biodiversity and threats to it, providing solid evidence to support an eventual designation of a federal protected natural area in the region (Vázquez-Reyes et al. 2018).
The protection of natural systems hinges on the establishment and development of environmental protection plans that encourage the active participation of residents in the conservation of vegetation cover and associated animal components (Vázquez-Reyes 2018).Reports like which are crucial for catalyzing the political will of authorities to allocate necessary budget and management efforts toward achieving the conservation of A. m. mexicanus and its habitat in the Alto Balsas region.
Mexico, A. militaris mexicanus extended continuously from Sonora to Chiapas, including the Balsas basin influence area in Michoacán, Guerrero, Puebla, and Oaxaca.Additionally, it is found in the eastern mountainous regions of Nuevo León, Tamaulipas, San Luis Potosí, and Querétaro (Iñigo-Elías 1999).However, its distribution and populations are currently in decline due to unsustainable capture for the illegal bird trade and habitat fragmentation resulting from land-use changes.Specifically, the habitat of this species in Mexico experiences an annual deforestation rate of 3.27-5.21%.Additionally, habitat loss has intensified in recent years of approximately 270,000 km² of potential habitat; by 2010, the loss had reached 150,000 km² and increased to 160,000 km² by 2016 (Rivera-Ortiz et al. 2013; Marín-Togo et al. 2012).These factors have led to the local extirpation of A. militaris over much of its range (Loza 1997; Rivera-Ortiz et al. 2008; Ríos-Muñoz and Navarro-Sigüenza 2009; Marín-Togo et al. 2012).Consequently, A. militaris is now listed under Appendix I of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES 2013), and as Vulnerable according to the International Union for Conservation of Nature (IUCN 2022), and in Mexico, it is considered as an endangered species (SEMARNAT 2019).
. It is noteworthy that within the Tehuacán-Cuicatlán Biosphere Reserve, located between Puebla and Oaxaca, approximately 190 km from our study area, exists a documented breeding population in Cañon del Sabino with an average of 80 individuals of A. militaris (Dirección de la Reserva de la Biosfera Tehuacán-Cuicatlán 2001; Rivera-Ortíz et al. 2008; Contreras-González et al. 2009).Additionally, 45 km from our study area in the Santa Cruz Achichiplilco in the municipality of Teotlalco, western Puebla, Hernández-Castán et al. (2012) reported an average of 36 macaws bordering the Sierra de Huautla Biosphere Reserve in the state of Morelos (Urbina-Torres et al. 2009) (Figure 2).Since 2006, records of A. militaris have been documented in the Alto Balsas basin, which includes parts of the state of Guerrero, more than 20 km to west of southeastern Puebla (Jiménez-Arcos et al. 2012; Vázquez-Reyes et al. 2018).Jiménez-Arcos et al. (2012) reported a breeding colony with an average abundance of 23 individuals in the Cañon Xicuhuetztlán of Cerro del Tecaballo, Papalutla in Guerrero.Vázquez-Reyes et al. (

Figure 1 .
Figure 1.Observation of Ara militaris mexicanus with breeding activity in the Cañón Grande in the Paraje del Reloj, Santa María Cohetzala Puebla.A, B. Courtship.C. Mating.D. Incubation of eggs.