First record of Camponotus textor Forel, 1899 (Hymenoptera, Formicidae) from Ecuador

. We present the first records of Camponotus textor Forel, 1899 (Hymenoptera, Formicidae) from Ecuador, representing the first records from west of the Andes. We explore the use of participatory science data from iNaturalist and scientific collections to understand the distribution of C. textor . While iNaturalist data alone cannot confirm the presence of C. textor due to the need for precise morphometric data, it can be a valuable tool for assessing the species’ potential distribution. This study increases Ecuador’s known ant species to 826 and Camponotus species to at least 61. Challenges in distinguishing C. textor and C. senex prevent us from determining C. textor ’s native status and highlight the need for further research on Camponotus in Ecuador.


INTRODUCTION
Ants are a group of eusocial insects accounting for more than 14,000 described extant species worldwide (Kass et al. 2022;Bolton 2024).More than 4,000 species and subspecies of ants are recorded in the Neotropics, and research efforts in the last decades have significantly advanced ant diversity characterisation in the region (Cardoso et al. 2010;Achury and Suarez 2017;Fernández et al. 2021;AntWeb 2024a).In Ecuador, recent studies have improved our understanding of the ant fauna, and currently 825 ant species and subspecies have been identified (Guénard et al. 2017;Pazmiño-Palomino and Troya 2022).However, further research is needed to enhance our understanding of regional and local biodiversity patterns and ecological processes of ant communities in this megadiverse country.
Camponotus Mayr, 1861 is the second largest ant genus, after Pheidole Westwood, 1839, with over 1,000 described species, further divided into 43 subgenera (Bolton 2024).Given the high morphological variability within Camponotus, a general taxonomic characterisation of this group poses challenges (Bolton 1973).Camponotus is distributed worldwide and comprises more than 450 species in the Neotropics (Fernández et al. 2021;AntWeb 2024a).Although the diversity of Camponotus in Ecuador has not been thoroughly revised, and the literature and specialised databases still have many inconsistencies, at least 60 species and five subspecies of Camponotus are known in mainland Ecuador (Salazar et al. 2015;Guénard et al. 2017;Pazmiño-Palomino and Troya 2022;AntWeb 2024b).
Traditional biodiversity data sources, such as museums and field surveys, are invaluable.However, these sources alone often struggle to keep pace with the vastness and complexity of ant diversity, making participative science an invaluable tool for researchers (Lucky et al. 2014).Thanks to the digital revolution, participative science now plays a crucial role in expanding our understanding of ant diversity by engaging a large number of people (Braschler 2009;Lucky et al. 2014;Sorvari 2022).This broad participation expands the geographical and temporal scope of biodiversity data collection efforts, providing data points that researchers may not otherwise have access, including the identification and monitoring of rare ants or the early detection of non-native species (Sheard et al. 2020;Báthori et al. 2022).
In this contribution, we present the first records of C. textor from Ecuador, based on specimens collected during fieldwork in the Pacific lowlands, and explore the potential of participative science in detecting this species.

METHODS
We conducted field expeditions to agricultural landscapes without remains of native forests nearby, and close to small human settlements in the Pacific lowlands of central Ecuador.Specimens were collected at two private farms: Hacienda La Chelita (HLC, −00.6170, −79.4689, 150 m elevation) and Hacienda Santa Teresa (HST, 00.5977, −079.3903,170 m elevation), in Los Ríos province, Ecuador.
Access to HLC was through the main street of the town of Los Ángeles, on the Troncal de la Costa highway connecting the cities of Santo Domingo de los Colorados and Quevedo.HST is 9 km northeast of HLC, 3 km after the town of Patricia Pilar on the Troncal de la Costa highway.Both farms have small houses surrounded by maize, cocoa trees, and African oil palm plantations.Since the early 1960s, intensive agricultural activities have eliminated all native vegetation, including secondary forests and shrublands.The closest area with native vegetation is approximately 30 km E (Samama Mumbes Wildlife Refuge).Both sites have average temperatures ranging from 22 to 24 °C and over 2,400 mm of annual precipitation, with a wet season between December and May and a dry season between June and November (Caicedo-Camposano et al. 2016).Three surveys were carried out: 9 October 2022 (HLC), 19 February 2023 (HLC), and 20 May 2023 (HST).One hour of sampling effort was dedicated to each survey.These surveys were conducted under the authorisation for specimens collection no.MAAE-ARSFC-2022-2203, issued by the Ministry of Environment, Water and Ecological Transition of Ecuador.
A total of 13 worker specimens were collected manually using entomological forceps and deposited in tubes with 70% ethanol.All collected specimens were dry-mounted and deposited at the Museo de Zoología, Universidad San Francisco de Quito, Quito, Ecuador (ZSFQ).Specimens were identified based on morphological and morphometric features following the guidelines of Fernández (2003) for genus identification, Longino (2006) and Mackay (2019) for subgenus identification, and Fox et al. (2017) for species identification.Specimens were examined using an Olympus SZX16 stereomicroscope and photographed with an attached Olympus DP73 digital camera.Eye length was measured between the anterior and posterior eye borders using cellSens Dimension v. 1.16.Measurements were made using photographs of all 13 workers in lateral view.
Occurrence data for C. textor from the Neotropics were obtained from iNaturalist, a participative science online portal by the California Academy of Science and National Geographic.Data search and extraction for iNaturalist were conducted on 20 September 2023 using the search terms "Camponotus textor" in the "Search" filter.We validated the taxonomic identification of each observation uploaded to iNaturalist and compiled geographic data, individually confirming all localities following the protocols described by Cisneros-Heredia and Peñaherrera-Romero (2020), and Cisneros-Heredia et al. (2023).iNaturalist data used for this study are available here: https://doi.org/10.5281/zenodo.10869127 To create the map showing the distribution of C. textor, we downloaded all occurrences available in the Global Biodiversity Information Facility (GBIF), excluding data from iNaturalist.Data search and extraction from GBIF were conducted on 26 July 2023 using the search terms "Camponotus textor" in the "scientific name" filter.These GBIF occurrences correspond to validated, vouchered records from the California Academy of Sciences, The Internacional Barcode of Life Consortium, the European Nucleotide Archive EMBL-EBI, and the Instituto de Ecología INECOL of Mexico.GBIF data used for map creation are available here: https:// doi.org/10.15468/dl.89w2ywAll collected specimens were workers (11 minor workers and two major workers).Ants were active during the morning and afternoon hours.At night, ants were inactive, and instead, Acromyrmex leaf cutter ants were seen in the same trails and locations used by C. textor workers during the day at HLC.No foraging behaviours were observed at HLC, while at HST, several workers were seen drinking water in a small plastic container near the nest.Other ant species found in sympatry with C. textor at both localities include Acromyrmex spp., Atta spp., Odontomachus spp., Tapinoma melanocephalum (Fabricius, 1793), Ectatomma spp., Crematogaster spp., and Paratrechina longicornis (Latreille, 1802).A silk nest of C. textor was found on top of a palm tree (Arecaceae: Bentickia sp.) at HST on 20 May 2023.The nest was about 60 cm long and approximately 4 m from the ground, built in a single palm leaf.A separate, smaller part of the nest was found in a different leaf from the same palm tree, but it seemed to be abandoned, as no ants were seen there.

Camponotus textor
Identification.We identified these specimens as Camponotus based on the following characteristics: posterior margin of clypeus apart from antennal alveoli by a distance equal to or greater than the diameter of alveoli and closure of metapleural gland (Fernández 2003).Specimens were identified as members of the subgenus Myrmobrachys Forel, 1912 because all workers exhibited a box-like propodeum with a broad, subrectangular dorsal face and the presence of gastral pubescence, a common feature in most species of Myrmobrachys (Longino 2006;Mackay 2019).One key characteristic differentiating C. textor from C. senex is the presence of appressed dense yellow pubescence on the gaster, giving C. textor a yellowish-golden colouration (Figure 1) (Longino 2006).Workers of C. textor generally have more reddish legs than C. senex workers (Figure 1) (Fox et al. 2017).However, leg colour in Camponotus spp.can be variable (Longino 2006), so eye length is the most useful characteristic for differentiating C. textor and C. senex (Fox et al. 2017).Eye length in C. textor workers ranges from 0.44-0.51mm, while in C. senex workers is 0.50-0.65 mm (Fox et al. 2017).All specimens examined for this study showed dense yellow pubescence on the gaster, reddish legs (Figure 1), and eye length 0.42-0.49mm (0.45 ± 0.02 mm, n = 13).Additionally, while C. senex is a species with high worker polymorphism, C. textor exhibits low worker polymorphism, as major workers are only slightly larger than minor workers (Longino 2006).Two workers collected at HST are most likely major workers, as they are slightly larger than the other collected workers.Also, the nest of C. textor is diagnostic, being a weaver ant building silk nests attached to leaves hanging from trees (Figure 2) (Longino 2006;Santos et al. 2016;Ramalho et al. 2016a).In contrast, C. senex is an opportunistic cavity nester, building nests in dead branches and twigs scattered on leaf litter (Longino 2006;Ramalho et al. 2016a).A total of 27 observations identified as C. textor were obtained from pictures in iNaturalist from the Neotropics, 10 of which were from Ecuador.These observations showed Camponotus (Myrmobrachys) ants with reddish legs and golden gaster.However, confirming the presence of C. textor based solely on photographs can be challenging.Eye length is the most reliable characteristic for distinguishing C. textor from C. senex, and it is not easy to measure in field-obtained photographs.We refrained from reporting the iNaturalist observations as confirmed records for C. textor.Nevertheless, iNaturalist observations suggest that the species occurrence in Ecuador is wider than can be inferred from our sampling in the Los Ríos province.Nine iNaturalist observations in Ecuador were from the Pacific lowlands, in the provinces of Los Ríos and El Oro, at elevations ranging from 10 to 490 m.One observation was from the inter-Andean valley in the Pichincha province at 2500 m.In addition to the Ecuadorian records, an observation of C. textor from Peru is also present in iNaturalist, from the Grapanazu sector, near Abra La Esperanza, Huancabamba district (iNaturalist 2022).

DISCUSSION
The records presented here are the first of Camponotus textor from Ecuador and the first west of the Andes, extending the geographic range of C. textor by approximately 1042 km from the nearest previously known locality (Platanilla, Panama; Figure 3) (AntWeb 2023b; GBIF.org 2023).Camponotus textor has been recorded in Central America (Honduras, Mexico, Costa Rica, and Panama;Longino 2006;Ramalho et al. 2017;Dáttilo et al. 2020) and South America (eastern Brazil, Bolivia, and French Guiana;Ramalho et al. 2017;AntWeb 2023a), making Panama the closest country to Ecuador in which C. textor is known (Figure 3).Given that C. textor was previously considered a subspecies of C. senex (Longino 2006;Ramalho et al. 2016a;Ramalho et al. 2016b), records of C. senex should be reviewed across its distribution to confirm its identification and avoid confusion with C. textor.Camponotus senex is considered a native species in several Central American countries and has also been reported in Venezuela, French Guiana, Guyana, Colombia, Peru, Ecuador, and Argentina (Escalante-Gutiérrez 1993;Fernández and Sendoya 2004;Longino 2006;Salazar et al. 2015;Franco et al. 2019).The paucity of information on the distribution of C. textor and the possible confusion with older records reported as C. senex in northern South America prevent us from determining if C. textor is native to Ecuador (and previously confused with C. senex) or corresponds to a non-native species recently introduced in the country.This situation is further exacerbated by the few records and scientific collections of ants made in the Pacific lowlands of Ecuador (Salazar et al. 2015).Species richness and composition of ant communities from western Ecuador are still poorly known (Salazar et al. 2015;Pazmiño-Palomino and Troya 2022;Salazar-Basurto et al. 2023).
Although participatory science data plays a significant role in providing geographic data for ant species (e.g., Salazar-Basurto et al. 2023), it is essential that identification characters can be clearly discernible in photographic records uploaded on participatory science platforms.Since C. textor and C. senex are cryptic species (Longino 2006;Ramalho et al. 2016a;Fox et al. 2017), key morphological characteristics used to diagnose C. textor are best observed in photographs taken under stereomicroscopic magnification (Fox et al. 2017), thus making their identification using field-taken photographs challenging.However, participatory science records could be valuable for identifying potential new locations where C. textor could be found.Based on the data on iNaturalist for the provinces of El Oro and Pichincha, we recommend a survey to uncover the species of Camponotus present here.
Even though Longino (2006) reported that C. textor inhabits mature forest canopy, all our records come from agricultural lands, suggesting that C. textor may adapt to disturbed landscapes.Santos et al. (2016) reported that C. textor can build its nests on 24 plant species belonging to 14 families, some of them being economically important plants, such as mango and citrus trees.Our record of a nest of C. textor on an introduced cultivated palm Bentinckia sp.adds another host plant for this weaver ant.
Taxonomic changes, like the separation of cryptic taxa previously considered as the same species, make it difficult to provide a precise number for species of Camponotus in Ecuador, as there are several inconsistencies across different lists and online databases.For example, Camponotus apicalis is not reported by Salazar et al. 2015, but is considered a valid species for Ecuador in Fernández and Sendoya (2004), Guénard et al. (2017) andAntWeb (2024b).This discrepancy is due to its taxonomic status, since C. apicalis is considered a senior synonym of Camponotus wheeleri (Fernández 2002;AntWeb 2023c).Fernández (2002) reported C. apicalis as a senior synonym of C. wheeleri, Fernández and Sendoya (2004) reported C. apicalis but not C. wheeleri, Salazar et al. (2015) reported C. wheeleri but not C. apicalis, and the databases GABI (Global Ant Biodiversity Informatics) (Guénard et al. 2017) and AntWeb (2024b) reported both C. apicalis and C. wheeleri for Ecuador.We suggest that further efforts should focus on a comprehensive revision of the genus Camponotus in Ecuador.
Our study highlights the importance of combining scientific collections and participatory science data to provide helpful information for identifying cryptic species.Although it is not possible to certainly identify observations from iNaturalist as C. textor, participative data can still be used as a first approach for assessing the potential distribution of this species in territories where they have not been previously reported (Báthori et al. 2022).While analyses based on morphological, ecological, behavioural, and molecular data are important to understand better the evolutionary relationships and taxonomy of Camponotus textor, participative data based on non-expert observations can aid in documenting this and other rare or poorly studied taxa.Further research on Camponotus ants in Ecuador should focus on their ecology.The ecological roles of C. textor need to be assessed to understand its impacts on agricultural ecosystems.Some carpenter ants have been reported as structural pests destroying wooden frameworks and electrical installations (e.g., Camponotus pennsylvanicus (De Geer, 1773) (Bueno and Campos-Farinha 1999); ecological facilitators of agricultural pests due to their aggressive behaviour safeguarding Homoptera (e.g., Camponotus rufipes (Fabricius, 1775) and Camponotus brasiliensis Mayr, 1862, Fowler 1991); or pest of honeybee colonies (e.g., Camponotus floridanus (Buckley, 1866), Akre and Hansen 1990).Carpenter ants may also be potential biocontrol agents in agricultural systems, with Camponotus senex demonstrating high efficiency in removing herbivores in mango trees (Santos et al. 2016) and reducing pests in shaded coffee (Philpott et al. 2004(Philpott et al. , 2005)).Further analysis and research could provide insight into the role of C. textor both as a potential pest and a biological pest control agent in agricultural systems.

Figure 1 .
Figure 1.Worker of Camponotus textor (ZSFQ-i11747) collected at Hacienda La Chelita, province of Los Ríos, Ecuador. A. Head in frontal view.B. Dorsal view.C. Lateral view.Red arrows show the characteristic appressed dense yellow pubescence of this species.Blue arrow shows the distinct reddish legs of C. textor.

Figure 2 .
Figure 2. Different views of the nest of Camponotus textor at Hacienda Santa Teresa, Los Ríos, Ecuador.

Figure 3 .
Figure 3. A. Map of Central and South America showing all known localities of Camponotus textor.B. Map of Ecuador showing the localities of C. textor in Ecuador.Yellow stars = New records of C. textor in Ecuador reported for this study; pink circles = iNaturalist observations reported as C. textor; green triangles = GBIF localities of C. textor.