The complex courtship display of the reddish hermit, Phaethornis ruber pygmaeus (Aves: Trochilidae), in the Atlantic Forest of northeastern BrazilABSTRACT
The courtship displays of hummingbirds are remarkable in their acrobatic flights and fluttering sounds, which are important examples of sexual selection. The courtship display of the reddish hermit, Phaethornis ruber (Linnaeus, 1758), was pointed out as one of the most complex among the Trochilidae, but it has been only partially described in the literature. Here, we describe the complete courtship display and the copulation behavior of this species. We provide the first video recording of this behavior, and using the slow-motion approach, we reveal unprecedented details of the courtship display of a hermit hummingbird. Each male’s performance session comprised four main displays: arc, pendulum, sliding horizontal, and pre-copulatory approach. Remarkable variations occurred in some of these displays, followed by characteristic sounds. While lekking behaviors have been thoroughly described for many hermit hummingbird species, courtship displays are cryptic behaviors that are difficult to capture, and there is still much to be learned about this subject. Because of the complexity of the observed behaviors, our study provides critical insights into the role of complex aerial displays in mate choice and the evolution of sexual selection in hummingbirds.
KEYWORDS:
Courtship behavior; hummingbird; mating behavior; pre-copulatory display; sexual selection
Hummingbirds (Aves, Trochilidae) have long called the attention of ornithologists due to the evolution of various outstanding characteristics, including their iridescent structural and pigmentary colors (Gruson et al. 2021), miniaturization, high speed and maneuverability (Clark 2009, Nuñez-Rosas et al. 2018), nocturnal torpor (Shankar et al. 2020), and highly specialized diet (Leimberger et al. 2022). Furthermore, they play important functional roles in communities, as more than 7000 plant species depend on hummingbirds for pollination, a type of mutualistic interaction that can potentially involve coevolutionary responses (Leimberger et al. 2022). When performing hovering flights, hummingbirds present one of the highest metabolic rates documented for vertebrates (Suarez 1992, Chen and Welch 2014), and the energy expenditure may be even higher during the reproductive displays, rich in vocal and fluttering sounds and acrobatic flights (Clark et al. 2013, Wilcox and Clark 2022). For this reason, their breeding displays are of ecological, behavioral and physiological interest (Cowell 2000, Clark et al. 2013, Wilcox and Clark 2022), yet the complete array of behavioral repertories is still unknown for many species (Ruschi 1986, Sick 1997, Hinkelmann et al. 2020).
The Trochilidae are also good examples of Rensch’s sexual size dimorphism rule, which predicts that within a clade, females are larger in the smaller species and males are larger in the larger species (Rensch 1960, Cowell 2000). Potential hypotheses to explain this pattern involve foraging niche divergences between sexes in habitats where food is scarce; eggs’ size influencing female body size for fecundity, and the sexual selection hypothesis. The latter predicts that female selection for complex courtship displays could favor smaller males with increased flight agility (Rensch 1960, Cowell 2000, Wilcox and Clark 2022).
The hermit hummingbirds comprise a well-supported monophyletic clade within the Trochilidae-Eutoxeres, Ramphodon, Glaucis, Threnetes, and Phaethornis (McGuire et al. 2007, 2014). Diagnostic traits include the predominance of pigmentary colors, mainly brown, grayish, and reddish, and in most species, males display at leks to attract mates (Hinkelmann and Schuchmann 1997). During the breeding season, assemblages of males can display at the lekking sites almost all day, exhibiting characteristic sounds and body movements (Schuchmann 1987). This is likely the reason why the lekking behaviors have been described in detail for many of the representatives of this hummingbird clade (Skutch 1964, Wiley 1971, Snow 1973, 1974, 1977, Stiles and Wolf 1979, Harger and Lyon 1980, Schuchmann 1987, 1995, Ramjohn et al. 2003, Felton et al. 2008). However, preliminary observations on at least two species of hermit hummingbirds, the reddish hermit, Phaethornis ruber (Linnaeus, 1758), and the grey-chinned hermit, Phaethornis griseogularis Gould, 1851, revealed that in addition to the conspicuous lekking displays, these birds also perform highly complex pre-copulatory courtship behaviors (Mobbs 1971, Schuchmann 1987). Because courtship displays are cryptic and likely less frequent than lekking behavior, they have remained poorly known for most hermit hummingbird species, even though this behavior can be an excellent and remarkable example of sexual selection.
The reddish hermit, Phaethornis ruber pygmaeus (Spix, 1824), is one of the smallest hummingbirds (8-9 cm; 1.8-3 g), widely distributed in South America, occurring in Ecuador, Bolivia, Colombia, Guyana, eastern Peru, southward to most of Brazil. They inhabit the understory of a variety of forested habitats, such as cloud and lowland forests, as well as forest borders and some types of savanna (Sick 1997, Hinkelmann et al. 2020). Males have a band of dark feathers on the breast and are slightly smaller than females (Cowell 2000, Hinkelmann et al. 2020), while the females have paler underparts. The courtship behavior of the reddish hermit was described for a pair maintained in captivity (Mobbs 1971) and for a pair observed in nature in the state of Mato Grosso, Brazil (Sick 1997), but only parts of the display sequence were captured. Here, we provide the first record of the complete courtship display and copulation behavior of the reddish hermit, with slow-motion videos permitting the description of details never observed for a hermit hummingbird.
We performed this study in an approximately 1,036 ha Atlantic Forest fragment in Alagoas, northeastern Brazil (10°00’S, 36°16’W). This area is a Private Natural Heritage Reserve (RPPN Mutum de Alagoas), which belongs to an ethanol and sugar industry (Usina Coruripe) surrounded by sugar cane plantations. The vegetation is classified as an open ombrophilous Atlantic Forest, and the climate is classified as AS’ according to Köppen: tropical with a well-defined dry and warm season from October through January and a long rainy season during the autumn and winter (Alvares et al. 2013). The average annual rainfall is 1600-1700 mm, and the average minimum and maximum temperatures range from 21-22 °C to 30-31 °C (Roda and Santos 2005, Barros et al. 2012). The observations occurred about 5 m from a trail, far from the forest border, in an area in late regeneration stage, with the predominance of emergent trees and no evidence of degradation.
We captured the courtship display while performing ad libitum searches for bird nests, and we recorded the movements using a semi-professional camera Sony Cyber-shot DSC-HX1 (Sony Corp., Japan). Due to the high speed of some movements, some of the display details were not observable by human eyes. For this reason, we also generated slow-motion videos of each display separately. Although most of the displays we observed did not perfectly match those described for other hummingbird species, whenever possible, we classified the displays based on the nomenclatures of similar behaviors already published in the literature-e.g., “pendulum” (Myers et al. 2022), “arc” (Schuchmann 1987, Felton et al. 2008), and “pre-copulatory approach” displays (Schuchmann 1987). For comparative purposes, we also searched for potential photographic or video records of pre-copulatory displays of hermit hummingbirds in the citizen science databases eBird (https://ebird.org), Macaulay Library (https://www.macaulaylibrary.org), and Wikiaves (https://www.wikiaves.com.br).
On 15 October 2022, at 3:30 pm, the buzz sound (drone sound) of a male reddish hermit’s wings caught our attention. At this moment, we could observe an individual perched on a slender horizontal twig, about 30 cm above ground, and a male performing an “arc display”, hovering about 30 cm above this individual. We immediately started the video recording, and we could observe two sequential and likely complete male courtship display sessions that together lasted almost 8 min (Suppl. Video 1). Although we could not observe the underparts of the perched individual, we assumed that it was a female because copulation occurred (see below). In the first display session, the female was more active, while in the second session, the male left the displaying area at the moment in which the copulation was about to occur, apparently because the female had not signalized interest. Each session comprised four main aerial displays that we classified as arc, pendulum, sliding horizontal line, and pre-copulatory approach. The sequences of the displays were the same in the two sessions, and below we describe these displays and their variations in the order they were performed.
Arc display
The male hovered approximately 30 cm above the female, forming an arc of approximately 140°, resembling a lateral jump in the air. Its head and tail were positioned upwards, the chest was projected forward, and the back was arched. During this display, the yellow base of the mandible, the whitish chin, and the white thigh feathers were exposed to the female. Notably, the feathers of the black chest patch were erected, likely due to the wind generated by rapid wing movements (Suppl. Video 1). Three distinct variations were observed. During most of the time, the male was constantly facing the female (Fig. 1A, Suppl. Video 2). At certain moments, however, the drone sound, the erection of the chest feathers, and the body curvature were intensified, while the time to perform each arc did not change. During these highly accelerated arcs, when reaching the extremity of each movement, the male turned its body outward (to the right or left; Fig. 1B, Suppl. Video 3). Another notable variation occurred at the end of the arc display when the male flew upwards, shook its body laterally, and wide opened the bill to show the yellow interior of the mandible to the female. This was followed by one or two pendulum dives before assuming the regular arc movement again, which was accompanied by a “té-tek, té-tek” sound (Fig. 1C, Suppl. Video 4).
Arc display observed in the pre-copulatory behavior of the reddish hermit, Phaethornis ruber pygmaeus, and its variations: (A) Display performed facing the female; (B) Accelerated display performed pointing the body towards outside at the end of the arc; (C) Shaking ascendant flight followed by diving.
Pendulum display
In this movement, the male performed sequential U-shaped flights within a very limited space, only slightly larger than its own body. With shallow dives, the male stretched its whole body, with the head and beak pointing forward. The display alternated between a dive that involved a forward and a backward flight, and another one that involved body contortion to sharply change the body direction (Fig. 2, Suppl. Video 5). During these flights, the male constantly exhibited the yellow basis of the mandible, the whitish chin, and the white thigh feathers, while constantly protruding its tongue. This movement was accompanied by a distinctive “treeek; treeek; treeek” sound, likely produced by the wing feathers.
Pendulum display observed in the pre-copulatory behavior of the reddish hermit, characterized by male’s sequential U-shaped flights.
Sliding horizontal line
The male moved back and forth across a horizontal line approximately 60 cm long, keeping its body perpendicular to the female. This movement was so fast that it was barely visible to human eyes. Slow-motion video evidenced that during these flights, the male changed its body direction before the end of the line, creating the impression that, near the end of the line, his body was sliding backward. The premature direction change ensured that the yellow beak remained visible to the female for most of the movement. This display was accompanied by another distinctive, metallic “tóoii, tóoii, tóoii, tóoii” sound (Fig. 3, Suppl. Video 6).
Sliding horizontal line display, a component of the pre-copulatory behavior of the reddish hermit, in which the male moves back and forth across a horizontal line about 60 cm long, with its body perpendicular to the female.
Pre-copulatory approach
The male approached the female frontally, shaking its body laterally, while keeping its tail closed. Its head and beak positions gave the body a concave shape (Fig. 4, Suppl. Video 7). Although the lateral movements were constant, it approached the female slowly. After that, it kept performing this movement while stopping right in front of her for 5-6 seconds, before the copulation occurred. During the display sessions, the female consistently moved its head to follow the male’s movements, held her wings lowered and moved her tail up and down. The female’s tail movements became broader and more frequent near the end of the first male display session, likely signalizing its interest in copulating. Copulation itself lasted approximately 18 seconds, with the male mounting the female from her left side while continuing to protrude its tongue (Suppl. Video 8).
Pre-copulatory approach display, in which the male reddish hermit approaches the female frontally, shaking its body laterally, assuming a concave body shape.
Despite being a widespread and common species in Brazil, few records were found about the courtship display of the reddish hermit in citizen science platforms. Of the 3,635 photos of this species available at WikiAves, only six show parts of the display (WA 282.800 and 282.801 for P. ruber ruber, and WA 587.935, 986.281, 2.125.124, 2.794.845 for P. r. pygmaeus), and two videos from Macaulay Library showed brief parts of the arc display.
Our observations of the arc and of the sliding horizontal displays matched the descriptions provided by Sick (1997). However, fewer details were provided, especially for the sliding horizontal movement, likely because it was observed only based on human eyes. Mobbs (1971) also provided detailed descriptions of the arc and the pre-copulatory approach displays for a pair of reddish hermits brought from Brazil and held in captivity in England. Yet, the other phases of the courtship display and the copulation were not mentioned by the above authors, suggesting that they have observed only incomplete performances (Sick 1997). A painting provided by Ruschi (1986) depicts an individual with a protruding tongue hovering very close to a perching individual, showing a body silhouette that seemed to be associated with the pre-copulatory approach display, but detailed information was not provided.
The most detailed record of a hermit hummingbird’s courtship behavior was described for the grey-chinned hermit from Colombia (Schuchmann 1987). This species’ aerial displays were divided into three phases: i) horizontal arc, in which the male’s tail and head positions were similar to that we observed for the reddish hermit; however, variations in speed, in body directions, and the alternations of the arc movement with brief pendulous dives we observed were not reported for the grey-chinned hermit; ii) a rapid back and forth movement, that seemed analogous to the sliding horizontal display we mentioned, but for which few details were described. Based on the provided drawing, this display seemed to follow a curve rather than a straight line, as in the reddish hermit, but it is to be confirmed; and iii) the pre-copulatory approach, where a drawing evidenced the similar concave body silhouette we observed for the Reddish Hermit, but further details were not provided. The pendulous display was not reported. It remains unclear whether the lower number of details provided for the courtship display of the grey-chinned hermit was because its display is less complex than that of the reddish hermit or if the published description was simplified. Our data revealed the importance of slow-motion videos for detailed behavioral descriptions, meaning that further video recordings are needed to confirm the pre-copulatory steps of other Hermits. Because these events seem not to occur frequently, citizen science records could play an important role in generating this type of data.
The arc display we observed seemed homologous to the rotation and arc movements recorded for a male reddish hermit hovering in front of another male in a lowland forest in Bolivia (Felton et al. 2008), but there were key differences. In the “rotation display”, the tail of the displaying male was rapidly raised and lowered, and it rhythmically opened and closed its bill to show the yellow gape. This movement was sometimes alternated with an approximately 100° “arc” display. Because both the perching and the displaying individuals were males, these behaviors were interpreted as antagonistic interactions between territorial individuals (Felton et al. 2008). Although simpler aerial displays were observed for males hovering in front of other males, apparently in territorial defense (Snow 1973, Felton et al. 2008), detailed descriptions of the reddish hermit’s singing assemblages (leks) indicate that these gatherings did not involve aerial movements (Davis 1934, Felton et al. 2008). This suggests that the courtship display is way more complex than the lekking behavior in this species.
Because we have observed two complete display sessions from the beginning, one culminating in copulation, we are confident that we have captured all the steps of the courtship display. In many hummingbird genera, courtship displays involve long diving and shuttle flights across the sky, covering dozens of meters (Clark 2009). The displays of the reddish hermit, however, occurred very close to the female, within a minimal space. This may be an adaptation to the forest understory environment, where visibility reaches only a few meters. Although analyzing the diversity and evolution of hummingbird courtship displays is beyond the scope of this manuscript, to our knowledge, this is the first record of the complete courtship display of a hermit hummingbird. While lekking behaviors have been thoroughly described for many hummingbird species, courtship displays are secretive and punctual behaviors that are difficult to capture, and much has still to be learned about this subject. Differences in the courtship display may be important to provide additional support for the taxonomic hypothesis, as P. ruber and P. pygmaeus have been considered sufficiently distinct to merit specific status (Piacentini, pers. comm.). Because our observations occurred within a well-preserved Atlantic Forest understory area, we do not have evidence about how habitat disturbance could affect this behavior, which is of conservation concern and should be investigated in future works.
Here, we present the first complete documentation of a hermit hummingbird’s courtship display, revealing an intricate and highly energetic performance. Given the high energetic cost of these displays, our findings provided further evidence for the importance of energetic performance in the sexual selection of hummingbirds. Although hypotheses based on pre-copulatory behaviors to account for sexual size dimorphism have not been tested for the hermits (see Wilcox and Clark 2022), this complex display associated with the fact that males have smaller body size, provide support for the sexual selection hypothesis that female selection for complex courtship behaviors could favor smaller males with increased flight agility (Cowell 2000, Wilcox and Clark 2022). As Mobbs (1971) noted after observing courtship behaviors of various hummingbird species in captivity: “the display of the male Reddish Hermit really has to be seen to be believed”.
ACKNOWLEDGMENTS
We are grateful to the Usina Coruripe for authorizing field work in the private area. Fernando Pinto, Sônia Roda, Marcela Daher, Carlos Monteiro, and Luzenilton Brito provided logistical support in the field. We also thank Flor Maria Guedes Las-Casas, Wallace Rodrigues Telino Junior, Guilherme Santos Toledo de Lima, Manoel Martins Dias Filho, and three anonymous referees for their important suggestions on previous versions of the manuscript.
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ADDITIONAL NOTES
- ZooBank register
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Data Availability Statement
All data generated and/or analyzed are included in this article and its Supplementary Materials.
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Funding Statement
This research was supported by the ARCA project from the Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP 2017/23548-2). LWLA was supported by a PhD fellowship from the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq 142308/2019-6); MCC received a post-doc fellowship from FAPESP (2020/13489-1). Productivity Research Fellowships were granted by CNPq to MRF (308702/2019-0 and 304213/2022-5) and LFS (308337/2019-0).
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Ethic Statement
Field work was authorized by SISBio/MMA (Proc# 66157-6). The methods were reviewed and approved by the Ethics Committee on Animal Use of the Universidade Federal de São Carlos (Proc# 1405291118).
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AI Statement
No artificial intelligence tools were used in the preparation of this manuscript.
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How to cite this article
Lobo-Araújo LW, Costa MC, Santos EL, Silveira LF, Francisco MR (2026) The complex courtship display of the reddish hermit, Phaethornis ruber pygmaeus (Aves: Trochilidae), in the Atlantic Forest of northeastern Brazil. Zoologia 43: e24082. https://doi.org/10.1590/S1984-4689.v43.e24082
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Published by
Sociedade Brasileira de Zoologia at Scientific Electronic Library Online - https://www.scielo.br/zool
Supplementary Material
Supplementary Video 1.
Supplementary Video 2.
Supplementary Video 3.
Supplementary Video 4.
Supplementary Video 5.
Supplementary Video 6.
Supplementary Video 7.
Supplementary Video 8.
Authors: LW Lobo-Araújo, MC Costa, EL Santos, LF Silveira and MR Francisco.
Data type: Species behavior.
Copyright notice: This dataset is made available under the Open Database License - ODBbL (https://opendatacommons.org/licenses/odbl/1.0/). The ODbL is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
All data generated and/or analyzed are included in this article and its Supplementary Materials.
