2024_Camponotus_textor
2024_Camponotus_textor
2024_Camponotus_textor
biodiversity data
1 Universidad San Francisco de Quito USFQ, Colegio de Ciencias Biológicas y Ambientales, Instituto de Biodiversidad Tropical IBIOTROP, Laboratorio de Zoología
Terrestre, Museo de Zoología, Quito 170901, Ecuador
2 Università di Bologna, Department of Biological, Geological, and Environmental Sciences, Bologna, Italy
3 Instituto Nacional de Biodiversidad INABIO, Quito, Ecuador
Corresponding authors: Ignacio J. Moreno-Buitrón (ignaciomorenob99@gmail.com); Diego F. Cisneros-Heredia (diego.cisnerosheredia@gmail.com)
Abstract. 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.
Key words. Camponotini, distribution, Formicinae, insect, Pacific lowlands, new records, weaver ant
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 Neo‑
tropics, 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).
Camponotus textor Forel, 1899 is a weaver ant that inhabits across the Neotropics, with populations
Academic editor: Filippo Di Giovanni
reported in Mexico, Honduras, Costa Rica, Panama, French Guiana, Brazil, and Bolivia (Longino 2006;
Received: 10 November 2023 Ramalho et al. 2017; Dáttilo et al. 2020; AntWeb 2023a). Camponotus textor was previously considered
Accepted: 5 April 2024 a subspecies of the cavity-dwelling C. senex (Smith, 1858), until Longino (2006) validated it as a separate
Published: 7 May 2024 species. Its recognition as a distinctive species is currently supported by morphological, behavioural,
ecological, biochemical (cuticular hydrocarbons), and molecular traits (Longino 2006; Ramalho et al. 2016a;
Fox et al. 2017).
Copyright © The authors. This is an open‑access Traditional biodiversity data sources, such as museums and field surveys, are invaluable. However,
article distributed under terms of the Creative
Commons Attribution License (Attribution 4.0 these sources alone often struggle to keep pace with the vastness and complexity of ant diversity, making
International – CC BY 4.0) participative science an invaluable tool for researchers (Lucky et al. 2014). Thanks to the digital revolution,
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Moreno-Buitrón et al. · Camponotus textor in Ecuador 578
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 collect‑
ed 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-Cam‑
posano 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 identifi‑
cation, Longino (2006) and Mackay (2019) for subgenus identification, and Fox et al. (2017) for species iden‑
tification. 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 Cisne‑
ros-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 Acad‑
emy 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.89w2yw
RESULTS
Camponotus textor Forel, 1899
New records. ECUADOR – Los Ríos • Los Ángeles, Hacienda La Chelita; −00.6170, −079.4689; 150 m alt.;
09.X.2022; J. Montalvo & I. J. Moreno-Buitrón; hand collected; 1 ♀, ZSFQ-i11759, dry preserved • same
locality; 19.II.2023; J. Montalvo & I. J. Moreno-Buitrón; hand collected; 3 ♀, ZSFQ-i11747, dry preserved; 3
♀, ZSFQ-i11748, dry preserved; 3 ♀, ZSFQ-i11749, dry preserved • Patricia Pilar, Hacienda Santa Teresa;
−00.5977, −079.3903; 170 m alt.; 20.V.2023; J. Montalvo & I. J. Moreno-Buitrón; hand collected; 3 ♀,
ZSFQ-i11755, dry preserved.
All 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 forag‑
ing 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.
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 0.44–0.51 mm, 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 0.42–0.49 mm (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; San‑
tos 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
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.
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 report‑
ed by Salazar et al. 2015, but is considered a valid species for Ecuador in Fernández and Sendoya (2004),
Guénard et al. (2017) and AntWeb (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 data‑
bases 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 assess‑
ing 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 eco‑
logical 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 instal‑
lations (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., Campono-
tus 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, 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.
ACKNOWLEDGEMENTS
We are grateful to the owners of Hacienda La Chelita and Hacienda Santa Teresa for their hospitality and
for granting us complete access to their properties for carrying out this research project, especially Jaime
Andrade, Emma Andrade, Hilda Andrade, Walter Andrade, Nancy Andrade, María Fernanda Andrade, and
Manuel León. We thank the staff of the Laboratory of Terrestrial Zoology, Institute of Tropical Biodiversity
IBIOTROP, Universidad San Francisco de Quito USFQ (Ecuador) for their assistance during our use of its high
magnification stereomicroscopy equipment and museum facilities, and the support offered by Margarita
López-García, Emilia Peñaherrera, and Jorge Montalvo at the Museum of Zoology of Universidad San Fran‑
cisco de Quito. Nelson Miranda, professor at USFQ and curator of the Herbarium of Economic Botany of
Ecuador, helped us identify the host palm where the nest of C. textor was found. We thank Emilia Peñaher‑
rera and Naia Andrade for their help with the map.
ADDITIONAL INFORMATION
Conflict of interest
The authors declare that no competing interests exist.
Ethical statement
This study was authorised by the Ministry of Environment, Water and Ecological Transition of Ecuador (no.
MAAE-ARSFC-2022-2203)
Funding
This study was financially supported by Universidad San Francisco de Quito USFQ, through research,
outreach and operative funds assigned to the Institute of Tropical Biodiversity IBIOTROP, and a publication
grant awarded to Diego F. Cisneros-Heredia by the office of the Dean of Research.
Author contributions
Conceptualization: IJMB, SER, DFCH. Data curation: IJMB. Formal analysis: IJMB, DFCH. Funding acquisi‑
tion: DFCH. Investigation: IJMB, DFCH. Methodology: IJMB, DFCH. Resources: IJMB, DFCH. Supervision:
SER, DFCH. Visualization: IJMB. Project administration: IJMB. Validation: IJMB, SER, DFCH. Writing – original
draft: IJMB, SER, IBM, DFCH. Writing – review and editing: IJMB, SER, IBM, DFCH.
Author ORCID iDs
Ignacio J. Moreno-Buitrón https://orcid.org/0009-0003-1067-7760
Selene Escobar-Ramírez https://orcid.org/0000-0002-1603-4710
Isabel Becdach-Mesia https://orcid.org/0009-0004-4626-501X
Diego F. Cisneros-Heredia https://orcid.org/0000-0002-6132-2738
Data availability
All data supporting this study’s findings are available in the main text and the following depositories: https://
doi.org/10.5281/zenodo.10869127 and https://doi.org/10.15468/dl.89w2yw.
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