Natural history on canvas: Brueghel knew about bird-eating noctule bats
Edited by Scott V. Edwards, Harvard University, Cambridge, MA; received December 22, 2025; accepted May 12, 2026
June 29, 2026
123 (27) e2536525123
- [Information & Authors](https://www.pnas.org/doi/10.1073/pnas.2536525123#core-collateral-info "Information & Authors")
- [Metrics & Citations](https://www.pnas.org/doi/10.1073/pnas.2536525123#core-collateral-metrics "Metrics & Citations")
- [View Options](https://www.pnas.org/doi/10.1073/pnas.2536525123#core-collateral-fulltext-options "View Options")
- [References](https://www.pnas.org/doi/10.1073/pnas.2536525123#core-collateral-references "References")
- [Figures](https://www.pnas.org/doi/10.1073/pnas.2536525123#core-collateral-figures "Figures")
- [Tables](https://www.pnas.org/doi/10.1073/pnas.2536525123#core-collateral-tables "Tables")
- [Media](https://www.pnas.org/doi/10.1073/pnas.2536525123#core-collateral-media "Media")
- [Share](https://www.pnas.org/doi/10.1073/pnas.2536525123#core-collateral-share "Share")
Abstract
A recent study has provided the first direct evidence that the greater noctule bat (_Nyctalus lasiopterus_) preys on and consume nocturnally migrating passerines in flight, using a combination of biologging, acoustic and movement monitoring, and molecular analyses. This result builds upon evidence accumulated over the past two decades. The research began with the first study reporting passerine feathers in the feces of this bat species. Subsequent fecal pellet analyses revealed peaks in the consumption of migratory avian prey, which coincided with migration periods. This body of evidence culminated in the recent publication demonstrating that greater noctules capture and consume migratory birds in flight at high altitudes. Here, we show that this interaction was depicted more than four centuries ago. In the canvas Air (1611), a painting by Jan Brueghel the Elder that contains detailed representations of more than 60 bird species, the artist also included three bat species. One of them, unmistakably identified as noctule bat, is represented in flight with a small passerine in its mouth. Although the behavior portrayed does not fully match contemporary descriptions of prey handling, the specificity of this predation scene suggests some familiarity of the artist with this behavior in the greater noctule. As the digitization of art collections accelerates and analytical tools continue to advance, the value of these sources to provide valuable data—previously difficult to extract and often overlooked—and complement modern research approaches will notably increase.
Sign up for PNAS alerts.
Get alerts for new articles, or get an alert when an article is cited.
Natural history knowledge is built on the accumulation of observations. The amount, precision, and spatial coverage of observations have notably increased historically, both due to the growing number of observers and to technological advances. This progress is exemplified by the understanding of bird migration in the field of movement ecology (1). Historical attempts to explain bird migration included imaginative approaches—such as Charles Morton’s proposal that birds traveled to the Moon (2)—and the interpretation of incidental observations, as illustrated by the Pfeilstorch, the White Stork (_Ciconia ciconia_) found in Germany with an African arrow lodged in its neck. Nowadays, the discipline is supported by tracking technologies, biologging, and global platforms such as Movebank (1, 3), allowing researchers to identify migration routes of populations and individuals. The ever-increasing potential of technological advances to reshape ecological research and generate natural history knowledge is undeniable. However, some ecological processes now clarified through modern scientific approaches may have been observed or hinted at in earlier centuries.
Stidsholt et al. (4) recently described how the largest bat species in Europe, the greater noctule bat (_Nyctalus lasiopterus_), preys on nocturnally migrating passerines, a finding built on evidence accumulated in the past two decades. Dondini & Vergari (5) had detected passerine feathers in noctule fecal samples and identified two common species based on plumage remains, the European robin (_Erithacus rubecula_) and the blue tit (_Cyanistes caeruleus_). Ibáñez et al. (6) provided the first robust indication that greater noctule regularly preys on migratory songbirds, showing that bird remains in bat pellets were more frequent during the periods of peak passerine migration. Furthermore, Ibáñez et al. (7) later employed DNA sequencing to prove that the greater noctule’s diet included over 30 distinct bird species, with almost all being night-flying sub-Saharan migrants. Stidsholt et al. (4) resolved the interaction by pairing ultralight 3-D biologging tags with acoustic, altitude, and movement monitoring and then implementing molecular confirmation of prey remains. Thus, demonstrating that greater noctule bats capture migrating passerines at high altitude required the integration of fine-scale movement data, echolocation analyses, and DNA/X-ray evidence.
Greater noctule predation on migrating passerines has been described in the scientific literature only recently, following years of research, relentless technological innovation, and international collaboration. Thus, it is striking that a 17th-century oil painting depicts a noctule bat carrying a passerine in flight. Although the image might not constitute a scientific proof of the artist’s direct observation of this behavior (it might be a stylistic convention or the author’s own imagination) it certainly depicts a scene consistent with the recent description of the phenomenon.
The painting Air (Fig. 1 _A_), completed in 1611 by the Flemish artist Jan Brueghel the Elder, is a work of remarkable interest to any naturalist. The piece evokes an allegory of the element Air, in which the Greek muse Urania, holding an armillary sphere, is surrounded by a swirling multitude of birds, while in the distance, the gods Apollo and Diana drive their chariots of the Sun and the Moon. What first captures the viewer’s attention is the astonishing diversity of avian species portrayed—some familiar European birds and others exotic species that seem extraordinary for a painting from 1611. The painting represents more than 60 species, including nearly 40 recognizable native European bird species, at least 14 exotic ones, and a few representatives of domestic taxa. In an allegory devoted to Air, Brueghel also included some Chiroptera, the only mammalian order capable of flight. At least four individuals can be clearly seen in the composition—one hidden on the left corner (Fig. 1 _B_), two just below Apollo’s chariot (Fig. 1 _C_), and the last one in the upper right corner (Fig. 1 _D_). The one depicted in Fig. 1 _B_ corresponds to the genus _Plecotus_, recognizable for its size and long ears (Fig. 1 _E_), but the species represented in Fig. 1 _C_ are not possible to identify at the genus level, although they seem to belong to the Vespertilionidae family (Fig. 1 _F_).
Fig. 1.
(_A_) The canvas Air painted by Jan Brueghel the Elder in 1611, an allegory of the air element. (_B_) Bat of the genus _Plecotus_. (_C_) Bats, probably belonging to the family Vespertilionidae. (_D_) Bat likely representing a greater noctule (_Nyctalus lasiopterus_) with a songbird caught on its mouth while flying. (_E_) _Plecotus austriacus_, coinciding with the genus represented on panel _B_. (_F_) _Pipistrellus kuhlii_, a vespertilionid bat that exemplifies the morphological type represented in panel _C_. (_G_) _Nyctalus lasiopterus_ in flight, as the individual represented on panel _D_. Photographs by Daniel Fernández (_E_) and Elena Tena (_F_ and _G_).
The sector of Brueghel’s painting enlarged in Fig. 1 _D_ represent a bat whose morphological traits align unequivocally with the genus _Nyctalus_ (noctule bats). Key diagnostic features include short, broad, rounded ears, long narrow wings, and long uniform brown to reddish-brown fur (8). Among European noctules, the greater noctule (_Nyctalus lasiopterus_) is the largest species—clearly larger than the common noctule (_Nyctalus noctula_)— and possesses a distinctively robust head profile ([8); Fig. 1 _G_]. Furthermore, based on its size and the depicted behavior—clutching a small passerine in its mouth while on the wing—the identification is highly consistent with a greater noctule, an identification further corroborated by several experts. The avian predation behavior portrayed by Brueghel, therefore, nicely aligns the one described by ref. 4. It is now known that once a greater noctule captures a passerine in flight, it removes the wings and probably the head too while still airborne, holding the prey with the feet together with the uropatagium and taking repeated bites and chewing. This process can take up to 20 min and involves normal echolocation calls while flying (4). The painting seems to show the initial stage of this behavior.
The greater noctule might have been difficult to observe in Brueghel’s native Belgium by the 17th-century. However, prior to painting Air, Brueghel spent time in Italy, where the species does occur and where he could have encountered it directly. It is anyway improbable that he had actually observed events of bat predation passerines, especially given that such events occur at night. Even so, occasional diurnal activity has been documented in a closely related species, the common noctule, during autumn migration in Central Europe (9), raising the possibility that Brueghel could have witnessed similar behavior in daylight. Anecdotal records of bird consumption also exist for common noctule (10). Thus, although the proportions in the painting align more closely with a greater noctule—particularly relative to the size of the bird—the depicted bat could conceivably represent a common noctule as well. The passerine prey cannot be reliably identified beyond a general assignment due to the limited detail and partial view in the painting. Ibáñez et al. (7) identified up to 31 bird species from prey remains, most belonging to the Sylviidae and Muscicapidae families, which are consistent with the passerine depicted. However, the limited visibility and wide range of possibilities preclude assignment to a specific family. On the other hand, much like Dondini & Vergari (5) did almost 400 y later, Brueghel may have seen or been told about the presence of passerine feathers associated to noctule bats, and then represented this behavior in the painting with a degree of artistic license. We acknowledge interpretations of historical artworks must be cautious, particularly in the case of allegorical paintings, which often incorporated symbolic elements. For example, bats and birds could carry contrasting meanings such as darkness and light. However, the fact that a noctule bat, and no other bat species, is represented in the scene suggests an observational inspiration rather than purely symbolic convention. Several versions of Air were produced—by Brueghel or apprentices—and only the version analyzed here includes the scene of noctule predation. Yet the mere depiction of this now well-known predation phenomenon centuries before the advent of modern biologging or ultrasound detectors constitutes a remarkable fact.
Our finding highlights how unconventional sources (i.e., those not directly related to the Natural Sciences) can yield valuable insights for ecological knowledge (e.g., refs. 11–13). Since the term historical ecology was first introduced in the twentieth century, the use of historical data to investigate ecological processes has steadily expanded (14)—yet unconventional archives, including art and literature, remain largely underexplored. This is now beginning to change. In a recent Perspective, Navarro et al. (15) highlighted how extracting information from these unconventional sources can reveal long-term biodiversity data that are otherwise inaccessible.
As with movement ecology, technological progress is rapidly turning the recovery of historical data into a practical reality for ecologists. Thus, the ongoing digitization of museum collections and art galleries is opening new possibilities for ecologists to access and analyze these data more efficiently. Perhaps, as in the case of the canvas Air from Brueghel the Elder, many hidden observations are awaiting in pieces of art to be spotted by ecologists.
Materials and Methods
The artwork analyzed here was identified through a systematic review of artworks aimed at documenting bird and mammal species depicted in art. The painting corresponds to the original version of _Air_ by Jan Brueghel the Elder (1611), currently housed at the Musée des Beaux-Arts de Lyon (France), although other versions were produced later by the artist.
The bat species depicted—with particular attention to the bat depicted capturing a passerine in flight—were identified by experts in bat biology with extensive experience on the genus _Nyctalus_, particularly _Nyctalus lasiopterus_.
Data, Materials, and Software Availability
All study data are included in the main text.
Acknowledgments
Funding was provided by Training, Recruitment, Incorporation, and Mobility Program for R&D&I Human Resources of the Andalusian Research, Development, and Innovation Strategy (EIDIA), Horizon 2027 (BOJA reference no. 209, 31/10/2023) and Grant «Excelencia Severo Ochoa» CEX2024-001498-S funded by Ministerio de Ciencia, Innovación y Universidades/Agencia Estatal de Investigación/10.13039/501100011033. We thank Javier Juste and Carlos Ibáñez for expert identification of the bat species, particularly greater noctule bat (_Nyctalus lasiopterus_), together with the two coauthors of this paper S.S.-N. and E.T. We thank Carlos Gutiérrez-Expósito for his helps with bird species identification. We also thank Javier Juste for confirming Italy as a country with the presence of the greater noctule bat during Jan Brueghel the Elder’s stay, where the artist was able to observe the event. We additionally thank Carlos Ibáñez for reading the final version of the manuscript and providing valuable suggestions. Two anonymous reviewers greatly helped to improve the manuscript.
Author contributions
P.R.-V. and M.C. designed research; P.R.-V., S.S.-N., E.T., and M.C. performed research; P.R.-V., S.S.-N., E.T., and M.C. analyzed data; and P.R.-V. and M.C. wrote the paper.
Competing interests
The authors declare no competing interest.
References
1
2
H. S. Gladstone, An early work on bird migration. _Br. Birds_**22**, 220–224 (1928).
3