Received: 14 July 2022 | Revised: 10 December 2022 | Accepted: 22 December 2022

DOI: 10.1111/aje.13107

S H O R T C O M M U N I C AT I O N

Biofluorescence and predator avoidance in the giraffe

Kin Morandi1 | Anna K. Lindholm1 | Derek E. Lee2,3 | Monica L. Bond1,2

1
Department of Evolutionary Biology
and Environmental Studies, University of Abstract
Zurich, Zurich, Switzerland
Biofluorescence of mammalian pelage may serve to hide prey from predators sensi-
2
Wild Nature Institute, Concord, New
Hampshire, USA
tive to ultraviolet radiation, among other potential functions. To date biofluorescence
3
Department of Biology, Pennsylvania has been documented in nocturnal-crepuscular and fossorial mammals that are active
State University, University Park, under low-light conditions. Giraffes are primarily diurnal, but biofluorescent pelage
Pennsylvania, USA
might offer camouflage from their nocturnal felid predators. Using a full-spectrum
Correspondence camera we qualitatively analyzed UV reflectance and absorption in giraffe pelage
Kin Morandi, Department of Evolutionary
Biology and Environmental Studies, from a museum specimen. We found no trace of UV biofluorescence in the giraffe
University of Zurich, Zurich, Switzerland. pelage, suggesting that this trait may not be ecologically or biologically relevant in
Email: kin.morandi@me.com
giraffes. The function of biofluorescence in mammals remains elusive, but our study
Funding information contributes to the growing body of data about biofluorescence, or its lack thereof, in
Claraz Schenkung
diurnal versus nocturnal-crepuscular or fossorial mammal species.

Résumé
La biofluorescence du pelage des mammifères peut servir à cacher les proies des
prédateurs sensibles aux rayons ultraviolets, entre autres fonctions potentielles.
Jusqu'à présent, la biofluorescence a été documentée chez les mammifères nocturnes-
crépusculaires et fossoriels qui sont actifs dans des conditions de faible luminosité. Bien
que les girafes soient principalement diurnes, leur pelage biofluorescent pourrait leur
permettre de se camoufler de leurs prédateurs félins nocturnes. À l'aide d'une caméra
à spectre complet, nous avons analysé qualitativement la réflectance et l'absorption
des UV dans le pelage des girafes provenant d'un spécimen de musée. Aucune trace de
biofluorescence UV n'a été trouvée dans le pelage de la girafe, ce qui suggère que cette
caractéristique n'est pas pertinente d'un point de vue écologique ou biologique chez les
girafes. Le rôle de la biofluorescence chez les mammifères reste insaisissable, mais notre
étude contribue au nombre croissant de données sur la biofluorescence, ou son absence,
chez les espèces de mammifères diurnes ou nocturnes, crépusculaires ou fossiles.

Many organisms use cues from the ultraviolet (UV) wavelength of
light for foraging or intra-­and inter-­specific signalling (Douglas &
Jeffery, 2014; Macel et al., 2020; Olson et al., 2021). Biofluorescence
occurs when short wavelengths in the UV spectrum are absorbed
by organisms and longer spectral wavelengths are projected out-
wards, causing them to glow in a variety of colours when exposed
to UV light (Olson et al., 2021). Kohler et al. (2019) and Anich
et al. (2021) proposed that absorption of UV light and fluorescence
of longer wavelengths may ‘hide’ individuals from UV-­sensitive
predators and help them blend into a UV-­saturated environment.
Although less UV light is available during the night, most of the bio-
fluorescent mammals identified thus far are nocturnal-­crepuscular
(Douglas & Jeffery, 2014). Biofluorescence of the pelage under
UV light has been documented in marsupial opossums (Didelphids;
Meisner, 1983; Pine et al., 1985), the monotreme duck-­billed platy-
pus (Ornithorhynchus anatinus; Anich et al., 2021), springhares
(Pedetidae; Olson et al., 2021), pocket gophers (Geomys spp.; Pynne
et al., 2021) and flying squirrels (Glaucomys spp.; Kohler et al., 2019).

Afr J Ecol. 2023;00:1–4. wileyonlinelibrary.com/journal/aje© 2023 John Wiley & Sons Ltd. | 1
2 |
To the contrary, diurnal squirrels (Sciurus or Tamiasciurus spp.) do
not exhibit biofluorescence (Kohler et al., 2019). Thus, the common
theme among biofluorescent mammals discovered to-­date is that
they are active in low-­light conditions, such as during dawn, dusk,
and night-­time or underground.
The giraffe (Giraffa camelopardalis) is thought to have a high vi-
sual acuity, with relatively large eyes and excellent vision (Ishengoma
et al., 2017), the capacity to solve spatial details (Veilleux &
Kirk, 2014), and the ability to perceive movement to a distance of 2 km
(Dagg & Foster, 1976). Giraffes are active mostly in the day (Burger
et al., 2020; Scheijen et al., 2021; van der Jeugd & Prins, 2000), with
group members sharing in vigilance at night while taking turns sleep-
ing for short periods (Burger et al., 2020). Two of the giraffe's main
predators include two members of the family Felidae, African lions
(Panthera leo) and leopards (Panthera pardus) (Dagg & Foster, 1976).
Mammals with eye lenses that transmit UV light to the retina—­called
‘UV sensitivity’—­are able to see in a wider spectrum of wavelengths.
Among carnivores, the lenses of domestic cats (Felis catus) appear
highly perceptive to UV light (Douglas & Jeffery, 2014) suggesting
that other felids whose lenses have not yet been tested, including
lions and leopards, may be similarly UV sensitive. One function of
the complex and variable spots of the giraffe pelage is thought to be
camouflage, to break up their silhouettes from background trees and
bushes (Lee et al., 2018; Mitchell & Skinner, 2003), but giraffes might
remain vulnerable if their felid predators are able to perceive UV
reflectance. Therefore, biofluorescence of the giraffe's pelage may
serve an adaptive role as additional camouflage against predation
from UV-­sensitive predators through visual confusion, as hypothe-
sized by Kohler et al. (2019) and Anich et al. (2021). Here, we test
whether the pelage of a diurnal mammal species, the giraffe, exhibits
biofluorescence that has been documented to-­date in mammals that
are active under low-­light conditions.
We quantitatively analysed UV reflectance and absorption
in giraffe pelage based on the experiment carried out by Anich
et al. (2021) on the platypus coat from a museum specimen. We used
a full-­spectrum camera to photograph a pelage sample of a female
individual Masai giraffe (G. camelopardalis tippelskirchi subspecies or
G. tippelskirchi species) that was born in captivity in Vienna in 1961
and died in 1983 in Rapperswil, Circus Knie, and is housed in the
Zoological Museum collections of the University of Zurich. Normal
cameras re-­create pictures in the human-­visible spectrum (about
420–­680 nm), and empirical colour analyses require calibrated pho-
tographs from a full-­spectrum camera (Stevens et al., 2007). We
used a Full-­Spectrum Converted IR/UV Canon EOS RP Mirrorless
digital camera from the Forensics kit (Kolari Vision, 2017) with a
Canon EF 50 mm f/1.8 II lens. Filters were required in order to take
pictures in the desired spectrum: UV photography required a UV-­
pass filter (Kolari UV Bandpass Lens Filter) and a visible-­spectrum-­
pass filter (Kolari 52 mm Colour-­Correcting UV/IR-­Cut Lens Filter).
For image normalisation, we used standards (Stevens et al., 2007),
including diffuse grey standards and colour charts (X-­Rite Colour
Checker Passport). The photographs were taken in a dark room, so
MORANDI et al.
we artificially directed UV light to the giraffe pelage sample using a
Kolari Vision KV-­FL1 Multispectral IR/UV Flash, which also includes
UV and IR light and an UV-­only flashlight (Picatinny Rail-­mounted
UV Focus Assist Flashlight).
We placed the giraffe pelage on a table with the right thoracic
part facing upwards. The camera, with the multispectral flash and
the EF 50 mm lens mounted, was fixed framing a plain area of the
sample. The colour standard was placed on the pelage so that it was
included in all photographs. Different photographs were taken in
different lighting conditions (the room was always darkened before
taking each image) and with different filters mounted on the lens:
(1) under visible light with a visible-­spectrum-­pass filter, (2) under
UV light with a visible-­spectrum-­pass filter (to check UV reflec-
tance, namely biofluorescence), (3) under UV light with a UV-­pass
filter (to check UV absorption), (4) under visible light (daylight lamp/
multispectral flash) with UV-­pass filter. The photographs were then
qualitatively analysed to assess the “response” of the giraffe's pelage
against different types of lights.
From the giraffe pelage empirical imaging analysis, we detected
a colour difference between the raw and the normalised picture
(Figure 1a,b) provided by the camera's human-­biased colour resolu-
tion. No traces of UV biofluorescence were detected on the giraffe
pelage (Figure 1d). This was demonstrated by photographing the gi-
raffe sample under only UV lights with a visible-­light pass filter. A
slight UV reflectance can be noticed in some white areas around the
spots and on the mane hairs, but since it was not spread all over the
sample, we concluded that this was disturbance caused by dust and
other particles.
By photographing the pelage with a multispectral flash (which
again produces at the same time visible, UV and IR light spectra) and
controlling for VIS and IR lights using a UV bandpass filter which
enables only UV light to be captured in the photo, no signal was re-
corded. The resulting picture was almost black, also after normalisa-
tion, indicating that the giraffe pelage has no response under normal
light conditions against UV spectra portion.
Macel et al. (2020) noted that it is widely accepted in the scien-
tific community that biofluorescence plays a role in communication,
predation and camouflage in several taxa. However, data are sparse
on biofluorescence in mammals, particularly for those most active in
brighter light conditions such as diurnal, non-­fossorial species. We
had hypothesized that biofluorescence might serve to camouflage
giraffes, which are mostly active during the day, from their nocturnal
felid predators. However, we did not find any trace of UV biofluo-
rescence in the giraffe pelage. UV biofluorescence of the giraffe's
coat may therefore not be ecologically relevant. UV reflection traits
may indeed be adaptive in some way for nocturnal-­crepuscular or
fossorial mammals (Anich et al., 2021; Kohler et al., 2019; Olson
et al., 2021; Pynne et al., 2021), or, alternatively, there may be no
ecological or biological role to biofluorescence for any mammal
species. Our study contributes to the growing body of data about
biofluorescence, or its lack thereof, in diurnal versus nocturnal-­
crepuscular and fossorial mammal species.
MORANDI et al. | 3

F I G U R E 1 Giraffe pelage images (a) (b)

under different light environments. (a)
illustrates the giraffe pelage sample
photographed under normal light
(produced by multispectral flash) with a
visible light pass filter, which was then
normalised in ImageJ against the grey
standards (b). (c) shows the same giraffe
sample photographed under only UV light
with a visible light pass filter. Again, the
picture was then normalised against the
grey standards using ImageJ (d). No UV (c) (d)
reflectance was detected.

AU T H O R C O N T R I B U T I O N S Douglas, R. H., & Jeffery, G. (2014). The spectral transmission of ocular

All the authors have accepted responsibility for the entire content of
this submitted manuscript and approved submission.
AC K N O​W L E​D G E​M E N T S
We thank Martina Schenkel of the Zoologisches Museum of the
University of Zurich for enabling the imaging of the giraffe pelage. We
also thank the subject editor F. Bercovitch, reviewer T. Caro, and one
anonymous reviewer for their helpful comments and suggestions.
F U N D I N G I N FO R M AT I O N
Claraz Schenkung provided funding to purchase the full-­spectrum
camera and associated equipment.
C O N FL I C T O F I N T E R E S T
The authors declare no conflict of interest.
DATA AVA I L A B I L I T Y S TAT E M E N T
Data sharing is not applicable to this article as no new data were cre-
ated or analyzed in this study.
ORCID
Derek E. Lee https://orcid.org/0000-0002-1042-9543
Monica L. Bond https://orcid.org/0000-0001-8500-6564
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