Journal of Threatened
Taxa | www.threatenedtaxa.org | 26 April 2026 | 18(4): 28607–28614
ISSN 0974-7907 (Online) | ISSN 0974-7893 (Print)
https://doi.org/10.11609/jott.9740.18.4.28607-28614
#9740 | Received 07 March 2025 | Final received 08 March 2026| Finally
accepted 16 March 2026
Altered nocturnal vocal activity
patterns in Tropical Kingbird Tyrannus melancholicus
(Passeriformes: Tyrannidae) at a site with artificial
lighting
David Ramírez-Adame 1 , Claudia Cristina Valenzuela-Inzunza
2 , Rosa Gabriela Beltrán-López
3 ,
Eduardo Michael Acosta-Morán
4 & José Antonio Guerrero 5
¹,2 Maestría en Biología
Integrativa de la Biodiversidad
y la Conservación, Centro de Investigación
en Biodiversidad y Conservación, Universidad Autónoma
del Estado de Morelos, Av. Universidad 1001, Chamilpa,
Cuernavaca, Morelos, CP 62209, México.
3 Laboratorio de Ictiología,
Centro de Investigaciones Biológicas,
Universidad Autónoma del Estado de Morelos, Av.
Universidad 1001, Chamilpa, Cuernavaca, Morelos, CP 62209, México.
4 Grupo de Investigación
de Mamíferos Marinos A. C.,
Puerto Vallarta, Jalisco, México.
5 Laboratorio de Monitoreo
y Conservación de Fauna, Facultad
de Ciencias Biológicas,
Universidad Autónoma del Estado de Morelos, Av.
Universidad 1001, Chamilpa, Cuernavaca, Morelos, CP
62209, México.
1 adame.ram95@gmail.com, 2 clau.crisva@gmail.com,
3 rosa.beltran@uaem.mx, 4 michael.moran.2112@gmail.com,
5 aguerrero@uaem.mx (corresponding author)
Editor: Aditya Srinivasulu, Zoo Outreach Organisation,
Hyderabad, Telangana, India. Date of publication: 26 April 2026 (online & print)
Citation: Ramírez-Adame, D., C.C.
Valenzuela-Inzunza, R.G. Beltrán-López,
E.M. Acosta-Morán & J.A. Guerrero (2026).
Altered nocturnal vocal activity patterns in Tropical Kingbird Tyrannus melancholicus (Passeriformes: Tyrannidae)
at a site with artificial lighting. Journal of Threatened
Taxa 18(4): 28607–28614. https://doi.org/10.11609/jott.9740.18.4.28607-28614
Copyright: © Ramírez-Adame et al. 2026. Creative Commons Attribution 4.0
International License. JoTT allows unrestricted use,
reproduction, and distribution of this article in any medium by providing
adequate credit to the author(s) and the source of publication.
Funding: The study does not receive any funding.
Competing interests: The authors declare no competing interests.
Author details: David Ramírez Adame is master of science from the Center for Research in Biodiversity and
Conservation (CIByC) at the Autonomous University of
the State of Morelos, with an interest in avian ecology and conservation. Claudia Cristina Valenzuela Inzunza
is master of science from the Center for Research in Biodiversity and
Conservation (CIByC) at the Universidad Autónoma del Estado de Morelos. Interested in evolutionary
biology, phylogeography, and the genetic variation
and conservation of birds. Rosa Gabriela Beltrán López, PhD in biological sciences, currently
she’s professor and researcher in Universidad Autónoma
del Estado de Morelos, her research is focused in molecular systematics, phylogeography and biogeography for conservation of
vertebrates and freshwater taxa. Eduardo
Michael Acosta Moran is a biologist active member of marine mammals research group (GRIMMA) with interest of Humpback
Whale ecology, contributing to humpback whale photograph catalog of Bahía de
Banderas, México. Currently working as a specialist nature tour guide. Dr. José Antonio Guerrero is a professor
and researcher in the Faculty of Biological Sciences at the Autonomous
University of the State of Morelos specializing in genetics and ecology of
wildlife, especially in mammals. He is a leading expert in the ecology and
conservation of Volcano Rabbit.
Author contribution: DRA and CCVI designed the study, conducted the fieldwork, performed the
recordings and analyses, and wrote the manuscript. RGBL parti-cipated
in the fieldwork, contributed to the recordings, and reviewed the manuscript.
EMAM assisted in the fieldwork and recordings. JAGE wrote and reviewed the
manuscript and the statistical analyses.
Acknowledgements: We thank the Comisión Nacional de Áreas Naturales Protegidas (CONANP) for facilitating access to the Islas Marías Biosphere Reserve, as well as Pablo Zamorano, director of the Biosphere Reserve, and park rangers Verónica Escobar, Sinead Gómez, Alba Solis, and Juan Moya, for their support in the management of permits and for their accompaniment during the field trips. We also thank the Navy for their logistical support. David Ramírez-Adame and Cristina Valenzuela-Inzunza thank the Secretaría de Ciencia, Humanidades, Tecnología e Innovación (SECIHTI) for the scholarships granted for the completion of master’s studies. We appreciate Lynna Kiere’s English review and comments that improved the clarity of the article.
Abstract: Artificial lighting at night
(ALAN) modifies the vocal activity patterns of numerous birds. In this study,
we evaluated the impact of ALAN on the vocal activity patterns of the Tropical
Kingbird Tyrannus melancholicus on María Madre
Island, located in the state of Nayarit, Mexico. Vocalizations were recorded
using autonomous Song Meter Micro recorders at one site with ALAN and another
without. We analyzed sound spectrograms to quantify vocalizations and performed
behavioural observations. Our results showed
differences in vocal activity between the ALAN site and the non-illuminated
site. The ALAN site had higher total vocal activity, producing 3,947
vocalizations with at least one vocalization in every recording, compared to
228 distributed across 40% of the recordings at the non-ALAN site. Tropical
Kingbirds at the ALAN site also had a substantially longer and earlier-shifted
period of pre-dawn vocalization activity, from 0200 h to 0500 h, compared to a
very brief period around 0500 h at the site without ALAN, as well as
vocalizations throughout the night compared to a brief peak around twilight
with no nocturnal vocalizations. Finally, we observed that individuals at the
ALAN site continued to forage into the night, displaying increased agonistic
interactions such as chases, wing-fluttering, and physical confrontations. Our
results suggest that artificial light significantly alters the activity pattern
of the Tropical Kingbird, extending its vocal and foraging activity. Our study
could contribute to understanding the effects of light pollution on
insectivorous birds in island environments and shows the importance of reducing
the impacts of artificial light in natural habitats.
Keywords: ALAN, behaviour,
insectivorous birds, islands, light pollution, nocturnal foraging,
vocalization.
Spanish Resumen
(español): La iluminación
artificial nocturna (ALAN, por
sus siglas en inglés) modifica
los patrones de actividad de numerosas especies de aves, alterando su comportamiento
y actividad vocal. En este estudio, evaluamos
el impacto de la ALAN en los patrones de actividad vocal del Tirano Pirirí (Tyrannus melancholicus)
en la Isla María Madre, ubicada
en el estado de Nayarit,
México. Las vocalizaciones de esta
especie se registraron en un sitio con ALAN y otro sin
ALAN, utilizando grabadoras
autónomas Song Meter Micro. Analizamos
los espectrogramas para cuantificar las vocalizaciones y realizamos observaciones de comportamiento. Nuestros resultados mostraron diferencias claras en la actividad vocal en el sitio con ALAN en comparación con el sitio no iluminado.
El sitio con ALAN presentó una
mayor actividad vocal total, con 3,947 vocalizaciones con al menos una en cada
grabación, en contraste con 228 vocalizaciones distribuidas en el 40% de las grabaciones en el sitio sin ALAN.
Los Tiranos Pirirís en el sitio con ALAN tuvieron un período de vocalización previo al amanecer sustancialmente más prolongado y adelantado, de 0200
h a 0500 h, en comparación
con un período muy breve alrededor de las 0500 h en el
sitio sin ALAN, así como vocalizaciones a lo largo de toda
la noche, en contraste con un breve pico al anochecer sin vocalizaciones nocturnas. Finalmente, observamos que los individuos en el sitio con ALAN continuaron forrajeando durante la noche, mostrando también un aumento en las interacciones agonísticas, como persecuciones, aleteos y confrontaciones físicas. Nuestros resultados sugieren que la luz
artificial altera significativamente
el patrón de actividad del Tirano Pirirí, extendiendo su actividad vocal y de forrajeo.
Este estudio podría contribuir a comprender los efectos de la contaminación lumínica en aves insectívoras
en ambientes insulares y muestra la importancia de reducir los impactos de la luz artificial
en los hábitats
naturales.
Introduction
Increasing urbanization in economically
developing regions has often resulted in substantial increases in artificial
lighting at night (ALAN) (Cinzano 2003; Hölker et al.
2010). ALAN adversely affects ecosystems by modifying natural light regimes, a
phenomenon called “ecological light pollution” (Longcore
& Rich 2004). This affects terrestrial and marine habitats globally through
both direct illumination and indirect reflection of sky glow (Spoelstra & Visser 2013).
However, the effects and consequences of light pollution on biodiversity, ecosystems,
and ecological and evolutionary processes remain poorly understood (Dominoni 2015).
Most organisms are subject to
circadian cycles that are regulated by natural light cycles (Arendt 1998;
Bell-Pedersen et al. 2005). Light pollution alters these cycles in numerous
species by interfering with the natural light regimes that regulate their
biological rhythms (Cassone 2014). Beyond disrupting
sleep and activity patterns, ALAN can also have detrimental effects on behaviour, reproduction, and even survival (Raap et al. 2015). Desynchronization of the circadian clock
due to artificial light can have consequences at the physiological and
biochemical levels, altering processes such as hormone secretion and immune
responses, which potentially compromises organisms’ ability to adapt to their
environment and perform activities that are essential for their survival
(Gaston et al. 2013).
In addition to the direct
negative effects of artificial night-time lighting, ALAN can alter the foraging
activity of insectivorous by attracting arthropods, thereby providing a
consistent food source for insectivorous species. Opportunistic nocturnal
feeding has been observed at locations where ALAN leads to nighttime arthropod
activity, primarily in some species of amphibians (Perry et al. 2008), reptiles
(Garber 1978; Amadi et al. 2021; Baxter-Gilbert et
al. 2021), and birds (Santos et al. 2010; Ortiz 2013).
In avian species, one of the most
frequently reported effects of light pollution is the alteration of vocal
activity patterns and feeding behaviour, which in
some instances has been found to impact reproduction and survival (Longcore & Rich 2004; Dominoni
et al. 2013; Dominoni 2015). While birds commonly
exhibit marked vocal peaks at dawn and dusk (Gil & Llusia
2020), ALAN can substantially modify these vocalization rhythms. In species
such as Eurasian Blue Tit Cyanistes caeruleus, European Robin Erithacus
rubecula, Eurasian Blackbird Turdus
merula, Great Tit Parus
major, and American Robins Turdus migratorius, an advancement in the onset of morning
song and an extension of vocal activity after dusk have been observed in
illuminated environments (Miller 2006; Kempenaers et
al. 2010; Da Silva et al. 2014). However, most of these studies have focused
solely on identifying the first and last vocal events of the day, without
examining song patterns across the full 24-hour cycle, limiting our
understanding of continuous vocal activity (Dominoni
2015). Moreover, ALAN has been shown to affect not only vocal rhythms but also
foraging behaviours. Although some species are
typically diurnal, the Northern Mockingbirds Mimus
polyglottos have been observed utilizing the
visibility provided by ALAN to continue to feed their offspring after dusk (Stracey et al. 2014), and the Eurasian Blackbird has been
observed using artificial lighting to forage at night in urban areas (Russ et
al. 2015). It has been documented that in anthropogenically altered locations,
some visual-hunting waders demonstrate increased activity in illuminated areas,
increasing their ingestion rate by 83% (Santos et al. 2010). Illuminated areas
may also attract avian species to degraded habitats near urban areas where they
may be subject to fitness costs such as increased predation by domestic &
feral dogs and cats, lower quality nesting habitats, and noise & chemical
pollution, among others.
The Tropical Kingbird Tyrannus
melancholicus (Image 1), a passerine bird of the
family Tyranidae, is widely distributed, ranging from
the southwestern United States to southern Argentina (Stouffer et al. 2024). In
Mexico, this species is resident from southern Sonora, including the Islas Marías on the west coast and from central Tamaulipas on the
east coast extending southward along both coasts, including the Yucatán
Peninsula, and inland throughout the central region of the country (Stager
1957; Howell & Webb 1995). Both sexes vocalize throughout the day, mostly
while perched but also occasionally during flight. A distinctive dawn song,
characterized by a series of brief notes and delicate ascending trills, is
typically performed from shortly before dawn until sunrise and does not resume
for the remainder of the day (Skutch 1965; Smith
1966). Additionally, these birds emit social calls that are perceived as
high-pitched, thin warbles that exhibit contextual variations (Skutch 1965). The Tropical Kingbird feeds primarily by
aerial hunting of flying insects, with frequent returns to the same perch after
capturing prey (Fitzpatrick 1980). While foraging habits are predominantly
diurnal, there are anecdotal reports of Tropical Kingbirds hunting insects at
night in artificially illuminated areas in Brazil (Sick & Teixeira 1981;
Stouffer et al. 2024). Nevertheless, the influence of ALAN on the activity
patterns of this species remains unclear, and the specific effects on vocal
activity in less anthropized insular populations, such as those inhabiting
isolated regions like Islas Marías, have yet to be
elucidated.
Given that artificial
illumination continues to expand into remote regions, it is crucial to
understand its effects on avian populations. To address this, we compared the
vocal activity of the Tropical Kingbird at locations with and without
artificial nocturnal lighting in an island population that is not exposed to
other aspects of urbanization. We hypothesized that ALAN induces an extension
of vocal activity during nighttime hours, thereby altering the natural activity
patterns of this species.
Methods
During July and August 2024, we
conducted field research on María Madre Island (21.634° N, 106.541° W; Image 2)
with the main objective of studying the Elf Owl Micrathene
whitneyi. María Madre is the largest island in
the Islas Marías archipelago, situated within the
Islas Marías Biosphere Reserve in Nayarit, Mexico.
The climate is arid, with moderate precipitation occurring from June to October
and an average annual temperature of 24.9°C. In the Islas Marías,
during the sampling period, sunrise occurred at approximately 0500 h and sunset
around 1900 h. The island exhibits four distinct vegetation types: medium
sub-deciduous forest, low deciduous forest, crassicaulous
scrub with low thorny forest, and coastal dune associations (Sánchez-Mejorada 1984; CONANP 2021). Furthermore, the island has an
urbanized area, consisting of paved roads, houses for tourists and island
personnel, as well as some areas designated for naval bases.
During our field trips on the
island, we noted that individuals of Tropical Kingbird inhabiting the urbanized
area exhibited nocturnal activity. Consequently, we designed an observational
study to be conducted at two sites with contrasting lighting conditions. The
first site was situated in an area with consistent ALAN, where multiple lamps
are activated at dusk and remain on until dawn, providing approximately 10
hours of continuous artificial illumination each night. These lamps are
positioned at 30 m intervals. The second recording location was established in
the territory of a Tropical Kingbird, approximately 410 m from the nearest lamp
in an area without ALAN (Image 2). We implemented the same recording protocol
at both sites using autonomous Song Meter Micro recorders model SM-MIC
(Wildlife Acoustics, Inc., Maynard, USA) with built-in omnidirectional
microphones. Recorders were mounted at 2 m height on tree branches within
Tropical Kingbird territories, carefully positioned to avoid disturbance. The
recording protocol consisted of 10 min recording periods followed by 10 min
intervals without recordings, continuing until a complete 24-hour cycle was
achieved at both sites. The recordings were made continuously, with one day of
separation between both recording sets and a similar lunar phase (last quarter)
at both sites. All recordings were made in WAV format at 44.1 kHz/16-bit
resolution.
The vocal activity pattern of
Tropical Kingbird was characterized by quantifying the number of vocalizations
from 72 recordings per site, corresponding to a complete 24-hour recording
cycle. Data were extracted through systematic visual inspection of spectrograms
configured with a Hann window, 256 points, 90% overlap, and a discrete Fourier
transform size of 512 samples, using Raven Pro 1.6 software (Yang 2024).
Vocalizations were identified and counted by manually selecting all clear
acoustic events across the 24-hour cycle. Each detection was verified aurally
and visually by two independent observers to minimize false positives.
Vocalizations were manually identified and categorized as either dawn songs or
social calls (Figure 1; Audio
1; Audio
2). Dawn songs are apparently
produced only by males and defined as long, ascending trills composed of
stereotyped sequences of multiple syllables, typically emitted during pre-dawn
hours, and characterized by slightly longer and more grating notes. Social calls
are high-pitched, thin, and emitted by both sexes, consisting of short,
repeated notes (Howell & Webb 1995; Stouffer et al. 2024). Circular charts
were constructed utilizing the “ggplot2” package (Wickham 2016) of R Software R
4.4.2 (R Core Team 2024) to visualize the frequency distribution of dawn songs
and social calls over a 24-hour cycle. Subsequently, acoustic activity patterns
were statistically compared using the Mardia-Watson-Wheeler
circular nonparametric test (Zar 1998) using Oriana
4.02 software (Kovach 2013).
In addition to the recordings, to
document the nocturnal behaviour of Tropical
Kingbirds, direct observations were conducted during six nights at sites with
ALAN, simultaneously, were performed these six same nights walks at sites without
ALAN, between 1800 h and 0200 h, with a single walk conducted per site.
Results
At the site with ALAN, 100% of
recordings contained at least one Tropical Kingbird vocalization, totaling
3,947 vocalizations, of which 1,040 were dawn songs, and 2,907 were social
calls. In contrast, at the site without ALAN, 40% of recordings contained
vocalizations, for a total of 228 vocalizations, of which 13 were dawn songs
and 215 were social calls. During the sampling period, we observed that the
territories were generally occupied by a single individual; however, at the
site with ALAN, more agonistic interactions between individuals were observed,
including chases, wing-fluttering, and physical confrontations.
At the site with ALAN, dawn songs
were recorded from 0200–0500 h, exhibiting a peak around 0300 h (Figure 2A). In
contrast, at the site without ALAN, dawn song activity was more temporally
concentrated, occurring primarily near dawn at approximately 0500 h (Figure
2B). Statistical analysis revealed significant differences in dawn song
emission patterns between the site with ALAN compared to the site without ALAN
(W = 26.145, P < 0.001).
Social call patterns exhibited
notable differences between the sites with and without ALAN. At the
artificially illuminated location, social calls occurred throughout the 24-hour
cycle, with heightened activity during nighttime hours and a peak after sunset
(1900 h; Figure 2C). In contrast, in the site lacking artificial light, social
calls were less frequent, limited to daytime hours, and primarily concentrated
around twilight periods, with peak activity observed before sunset at 1800 h
(Figure 2D). Statistical analysis revealed significant differences between the
two sites (W = 56.456, P < 0.001).
Our field observations at ALAN
site revealed that Tropical Kingbirds foraged at night, frequently pursuing
insects drawn to the light sources (Video 1). These feeding behaviours were recorded from
dusk until 0200 h, covering the entire period during which field surveys were
conducted. Additionally, these birds displayed agonistic behaviours
towards conspecifics. In contrast, during six nighttime surveys at locations
without ALAN, we did not observe feeding behaviours
or detect any vocalizations of Tropical Kingbird after nightfall.
Discussion
The results of this study show
clear differences in vocalization patterns, both in dawn songs and in social
calls, of the Tropical Kingbird between a site with ALAN and a site without
ALAN. At the site with ALAN, an earlier onset of dawn singing
and a longer duration of vocal activity were observed, in contrast to the site
without ALAN, where vocal activity was more concentrated at dawn. This difference
was statistically significant. Such shifts in the timing of morning song in the
presence of ALAN have been reported in other bird species exposed to similar
conditions. For instance, previous studies have documented that artificial
light can advance the onset of vocal activity and extend its duration in urban
birds, which can in turn affect key processes such as reproductive success and
disrupt circadian rhythms (Kempenaers et al. 2010; Da
Silva et al. 2015).
Regarding social calls, we
observed significantly higher activity at the site with ALAN, with calls
recorded throughout the 24-hour cycle, including a peak in vocal activity after
sunset and a high concentration of calls during the night. In contrast, at the
site without ALAN, social calls were considerably less frequent and were only
recorded during daylight hours, peaking before sunset. This difference in the
temporal pattern of social calls was statistically significant, reinforcing
evidence that artificial night lighting substantially alters vocal
communication in this species on María Madre Island. While our findings
partially align with previous studies that have documented increased nocturnal
vocalizations in birds exposed to artificial light (Da Silva et al. 2014; Dominoni et al. 2015; Ursino et
al. 2022), most of those studies report this phenomenon only within narrow time
windows, shortly before or after sunset. In contrast, the present study
documents extended nocturnal vocal activity lasting throughout the night.
Furthermore, we observed nocturnal
foraging behaviour at the site with ALAN, including
active pursuit of insects attracted to the light and increased agonistic
interactions between individuals, such as chasing, fluttering, and physical
confrontations. These behaviours were not observed
during nighttime hours at the site without ALAN, where individuals remained
vocally and behaviourally inactive after sunset. This
type of behaviour is consistent with observations in
other bird species, in which opportunistic nocturnal foraging induced by ALAN
has been documented, facilitated by the attraction of insects to light sources
(Santos et al. 2010; Stracey et al. 2014; Russ et al.
2015). These studies indicate that ALAN can extend birds’ foraging activity
beyond the daytime period, altering natural foraging patterns.
Although the Tropical Kingbird is
widely distributed across the American continent and is common even in urban
areas with high levels of artificial light (Stouffer et al. 2024), records of
nocturnal vocal activity in this species are scarce in the literature. To date,
there is only a single isolated report of nocturnal vocal activity from a
continental population, where individuals were observed feeding on insects and
emitting social calls from 2300 h to 2400 h (Sick & Teixeira 1981).
Therefore, the more pronounced and extended response to ALAN observed in this
study could potentially be intensified by the ecological conditions of an insular
environment.
Previous work has shown that
island species often exhibit greater behavioural plasticity
and niche expansion (Scott et al. 2003; Losos & Ricklefs 2009; Dufour et al. 2024). These traits have been
associated with the evolution of relatively larger brains in island
populations, which facilitates the adoption of new foraging strategies and
space use (Sol et al. 2005; Cnotka et al. 2008; Sayol et al. 2018). In this context, the high levels of
nocturnal activity recorded in our study could be interpreted as an adaptive
strategy by which birds exploit the abundance of insects attracted by artificial
light, a resource that would otherwise be unavailable under natural conditions.
A potential limitation of the
study is the limited sampling effort, the lack of quantification of the number
of individuals per site, along with the absence of data on the reproductive
status of the birds during the brief sampling period, which could influence
vocal activity and foraging patterns of the individuals. Furthermore, our
results establish a foundation for developing conservation strategies aimed at
mitigating the impact of artificial illumination on species susceptible to
human-induced environmental changes.
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