Temporal Dynamics of Scout Release-Behavior of Termitophagous Ponerine ant, Megaponera Analis
J Insect Behav (2024) 37:233–242
https://doi.org/10.1007/s10905-024-09864-9
RESEARCH
Temporal Dynamics of Scout Release‑Behavior
of Termitophagous Ponerine ant, Megaponera Analis
Rifilwe V. Modiba · Christian W. W. Pirk
Abdullahi A. Yusuf
·
Received: 31 January 2024 / Revised: 26 September 2024 / Accepted: 8 October 2024 / Published online: 5 November 2024
© The Author(s) 2024
Abstract Megaponera analis is an obligate termitophagous species endemic to sub-Saharan Africa.
The species forages by sending out scouts to search
for termites. Once termites are located, the scouts
return to the nest and recruit nest mates. Scouts face
unpredictable environmental conditions, including
sudden flooding, temperature changes, high winds
and predation by natural enemies. We observed 519
scouting trips from 18 different colonies over three
years to determine how long it takes for the colony
to replace a scout that has likely died while on scouting duty or is lost due to environmental perturbations.
Study areas were at Maremani, Musina, Masebe and
D’nyala Nature Reserves in the Limpopo Province of
South Africa. We introduced definitions of scouting
turnover, scouting trip, and scouting session. Scouting turnover is the time it takes a successive scout
to leave the nest for a trip, whereas a scouting trip is
the journey a scout takes from the nest in search of
Supplementary Information The online version
contains supplementary material available at https://doi.
org/10.1007/s10905-024-09864-9.
R. V. Modiba (*) · C. W. W. Pirk · A. A. Yusuf
Social Insects Research Group, Department of Zoology
and Entomology, University of Pretoria, Pretoria 0028,
South Africa
e-mail: ;
R. V. Modiba
Department of Biological and Agricultural Sciences, Sol
Plaatje University, Kimberley 8300, South Africa
food. Since M. analis shows bimodal foraging behavior, the morning observations were recorded as the
first scouting session, while the subsequent session,
which starts in the afternoon, was recorded as the
second scouting session. The mean scouting turnover
for all 519 trips was 9.36 ± 0.64 (SD) minutes. When
turnover was divided into sessions, the first scouting
session lasted a mean of 7.77 ± 12.17 (SD) minutes,
while the second had 10.34 ± 18.32 (SD) minutes.
The minimum turnover was zero minutes, and the
maximum was 124 minutes.
Keywords Scouting dynamics · termite predation ·
foraging behavior · nestmate recruitment · bimodal
foraging
Introduction
In optimal foraging, communication is central to foraging success (Pyke and Starr 2021), and nest selection is equally critical for social insects as this affects
the colony’s fitness (Healey and Pratt 2008; Evison
et al. 2012). Generally, organisms approach foraging
in individual scouting or group searching (DechaumeMoncharmont et al. 2005). Foraging strategies utilized by different species of ants depend on the environment and habitat where they are found (Gibb and
Parr 2013), and these conditions have driven the foraging in some species to be bimodal.
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Benefits of group searching include sharing information such as food source availability (DechaumeMoncharmont et al. 2005); however, encountering
a predator or sudden change in environmental conditions can be detrimental to the colony, and this is
where individual scouting has an advantage. Scouts
play a pivotal role in locating and quantifying food
resources and making critical decisions that affect the
entire colony, such as hunting for new nest locations
(Franks et al. 2002).
Both communication and nest selection are mostly
the scouts’ responsibility, making scouts’ decisions
central to the colony’s survival. Biesmeijer and De
Vries (2001) have differentiated between a scout and
a recruit by defining a scout as an individual tasked
with searching for food independently, whereas the
recruit depends on the scout and nest mates for information on food sources. The scouting rate of ants is
significantly influenced by nest size, with polydomous
nests experiencing lower scouting rates than monodomous nests (Cao 2013). Scouts face unpredictable
natural phenomena like floods and gusty winds, and
how the colonies respond to these threats is unknown.
Eusocial insects are characterized by a high degree of
division of labor with specific tasks such as foraging,
caring for the young, and nest construction and maintenance allocated to individuals (Gordon 2002). Ants
are particularly sensitive to microclimatic changes,
which can influence nesting behavior (Karlik et al.
2016) and foraging (Hölldobler and Wilson 1990),
and scouts should communicate these changes to nest
mates for collective decision-making.
Scouting in Megaponera analis (syn. Megaponera
foetens and Pachycondyla analis) occurs bimodally,
with some scouting happening from the early morning
to about 11:00 am local time, while the next session
starts in the late afternoon and early evenings (Longhurst et al. 1979; Longhurst and Howse 1979; Yusuf
et al. 2013; Frank and Linsenmair 2017a, b; Frank et al.
2018a, b) and at night (Yusuf et al. 2013). Bimodal foraging is also practiced in non-ponerine ants, and the
driving factor is likely temperature, with the foraging
peaking at both sides of temperature extremes (Cerdá
et al. 2009). It is currently unknown why scouting is not
continuous, but this trait appears phylogenetically conserved as it is shared with Ophthalmopone berthoudi,
which is sister genus to M. analis (Schmidt 2013) and
also has bimodal foraging activity (Crewe and Peeters
1987). Literature, however, suggests that bimodality
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J Insect Behav (2024) 37:233–242
could be influenced by the unavailability of the termites
during hotter midday hours since termites are prone to
desiccation (Hewitt et al. 1972; Longhurst et al. 1979).
Scouting distance can reach 95 m, and searching
can last over an hour (Longhurst and Howse 1979;
Yusuf et al. 2014). Scouts use termite soil sheets as
kairomones to aid them in finding food sources quickly
(Longhurst and Howse 1978; Yusuf et al. 2014). When
the scout locates a food source, it quantifies it and
returns to the nest to recruit nestmates, laying a trail
pheromone (Longhurst et al. 1978). The trail lasts up
to 29 h (Hölldobler et al. 1994), thereby improving raid
position fidelity until termite prey are exhausted (Bayliss and Fielding 2002), and the scout always uses the
fastest way possible to the nest (Frank et al. 2018a).
When the scout arrives at the nest, it recruits between
200 and 800 nestmates back to the site, where it finds
termites (Frank et al. 2017; Yusuf et al. 2020). The
scout will lead a group of nest mates who walk in a column formation. The scout will be ahead of the column,
and behind it will be former scouts, while at the back
end of the column will be major ants, and these workers maintain position fidelity (Frank and Linsenmair
2017b). Although raids in M. analis have been studied
in detail, little is known about the dynamics of scouting and (...truncated)