Content uploaded by Shahmshad Ahmed Khan
Author content
All content in this area was uploaded by Shahmshad Ahmed Khan on Dec 29, 2023
Content may be subject to copyright.
Shahmshad Ahmed Khan Science Reviews - Biology, 2023, 2(4), 1-6
1
Review of the Role of Bees as Ecosystem Engineers in
Nature
Shahmshad Ahmed Khan1, PhD candidate
1University of Sargodha, Pakistan
Corresponding author: shamshadahmeduca@gmail.com
https://doi.org/10.57098/SciRevs.Biology.2.4.1
Received November 08, 2023. Revised November 18, 2023. Published online December 28, 2023.
Abstract: Bees are a highly important group, providing a multitude of services to ecosystems, most notably
through pollination. While much of the research on bees has traditionally focused on their role in pollinating a
variety of wild and cultivated crops, this review paper expands the discourse by examining their functions as
ecosystem engineers. Traditional definitions of ecosystem engineers exclude pollinators due to their non-
physical modifications of habitats for other species. However, contemporary studies challenge this perspective,
recognizing pollinators as integral ecosystem engineers who offer a range of direct and indirect ecological
services. For instance, bees’ pollination activities lead to the formation of dry fruits that subsequently serve as
shelters for various organisms. In addition, ground-nesting bees modify soil composition through mechanisms
such as aeration and bioturbation. This review aims to elucidate the transformative impact bees have as
ecosystem engineers, thereby enhancing our understanding of their ecological importance.
Keywords: Ecosystem engineers, Ecosystem function, Habitat modification, Soil Bioturbation, Ground nesting
soil
Introduction
Ecosystem engineers are organisms that modulate
the availability of resources for other species, either
directly or indirectly, by altering the physical state
of biotic and abiotic materials within their ecosys-
tem (Jones, Lawton, & Shachak, 1994). These engi-
neers, through modifications of abiotic conditions,
instigate changes in the biological responses of res-
ident species, thereby influencing ecosystem func-
tioning. Ecosystem functioning encompasses both
physiochemical and biological processes within the
ecosystem, which are closely associated to human
well-being. This connection is evident in services
such as carbon sequestration, productivity, and nu-
trient cycling (Byers, 2022).
Overview of Bees as Ecosystem Engineers
Mills, Soulé, and Doak (1993) identified five catego-
ries of keystone species, among which includes key-
stone habitat modifiers or ecosystem engineers
(Lawton & Jones, 1995). These engineers are further
classified as either autogenic or allogenic (Jones et
al., 1994). Autogenic engineers modify environ-
ments with their physical structures, such as living
or decaying tissues. Conversely, allogenic engineers
transform environments by manipulating biotic or
abiotic components through various activities
(Jones et al., 1994; Jones, Lawton, & Shachak, 1997).
However, the complexity inherent in species rich-
ness makes classifying certain organisms challeng-
ing. For instance, pollinators like bees were not tra-
ditionally considered ecosystem engineers by Jones
et al. (1994) because they primarily modulate re-
sources supply for seed and fruit consumers. None-
theless, recent perspectives, such as those put forth
by Wilby (2002), suggest expanding the term ‘eco-
system engineering’ to include organisms whose
actions create or modify habitats, even if their influ-
ence does not align with traditional definitions of
engineering (Casas-Crivillé & Valera, 2005). Under
this broader definition, pollinators like bees can in-
deed be considered ecosystem engineers due to
their direct and indirect impacts on ecosystems
(Cardoso, Rezende, Caetano, & Oliveira, 2023;
Casas-Crivillé & Valera, 2005).
Shahmshad Ahmed Khan Science Reviews - Biology, 2023, 2(4), 1-6
2
Bees form a monophyletic group of insects with
over 20,000 described species across seven families,
inhabiting a diverse array of latitudes and terres-
trial ecosystems (Patel, Pauli, Biggs, Barbour, & Bo-
ruff, 2021). Globally, both wild bees and managed
honeybees are economically significant as pollina-
tors (Kremen & Chaplin-Kramer, 2007; Kremen,
James, & Pitts-Singer, 2008). Their efficient pollen
transport, reliance on floral resources, and social be-
haviors, from semi-social to eusocial classifications,
enable them to be prolific pollinators for a wide
spectrum of plant species (Klein, Boreux, Fornoff,
Mupepele, & Pufal, 2018; Ollerton, 2017). Bees are
responsible for pollinating more than 90% of the
world’s flowering crops (Klein et al., 2007). In addi-
tion to increasing crop yields (Aizen et al., 2019;
Aizen & Harder, 2009), bees enhance the nutritional
value and quality of fruits and vegetables (Ahmad
et al., 2021; Khan et al., 2022; Klatt et al., 2014), indi-
rectly mitigating food waste associated with aes-
thetic and quality imperfections (Gunders & Bloom,
2017). The crucial role of wild and managed bees in
pollinating wild plants in forest ecosystems is also
well-documented (Senapathi et al., 2015).
In this review, we aim to understand the role of bees
as ecosystem engineers and explore their ecological
functions across various ecosystems. While tradi-
tionally pollinators, specifically bees, were not
acknowledged as ecosystem engineers due to their
focus on resource provisioning (Jones et al., 1994),
recent studies indicate that pollinator bees might
fulfill the role of ecosystem engineers providing di-
verse environmental services. Bees also serve as bi-
oindicators for contaminants resulting from in-
creasing anthropogenic activities. The honeybee
(Apis mellifera), in particular, is recognized as a val-
uable bioindicator due to its close association with
the ecosystems it inhabits (Sadeghi, Mozafari, Bah-
mani, & Shokri, 2012). Bees encounter various con-
taminants during foraging, in flight, and through
nectar and pollen consumption (Ruschioni et al.,
2013). Analysis of the honeybees can reveal the
presence of elements such as Cu, Cr, Zn, Mn, and Fe
concentrations, which correlate with seasonal
changes and agricultural practices (Skorbiłowicz,
Skorbiłowicz, & Cieśluk, 2018).
Bees' Role in Pollination and Biodiversity
Pollination is important for the stability of natural
ecosystems and the production of various vegeta-
bles and crops, forming a critical link between agri-
culture and the cycle of life. So the process of
pollination significantly contributes to economic
improvement (Gill et al., 2016; Hristov, Neov,
Shumkova, & Palova, 2020). The term ‘pollination’
is the transfer of pollen from the male anthers to the
female stigma, either within the same flower or be-
tween different flowers on same or on different
plants.
Over 75% of food crops are partially or completely
dependent on animal pollinators, with pollinating
bees playing a crucial role in the production of
fruits, vegetables, coffee, cocoa, and almonds (Klein
et al., 2007). Pollinators such as bees are also key in
supporting biodiversity, as evidenced by the posi-
tive correlation between plant and pollinator diver-
sity (Ollerton, 2017). The abundance and species
richness of pollinators in an area are considered as
an indicator of the overall health of that ecosystem.
Certain crops, including cherries, blueberries, and
apples, solely rely on bee pollinators for up to 90 %
of their yield, indicating the interdependence be-
tween bee pollinators and plant diversity (Biesmei-
jer et al., 2006).
Pollination as Ecosystem Engineering
For pollinators to function as ecosystem engineers,
certain conditions must be met. Firstly, the plant
species must require biotic factors for pollination.
Secondly, the resultant fruits should be of a dry type
that is not consumed by frugivores and predators,
allowing them to remain in the environment for an
extended period. Thirdly, post-seed dispersal, these
fruits should create habitats for other organisms
(Cardoso et al., 2023). In this scenario, a plant acts
as an autogenic engineer, modifying the environ-
ment through its physical structures. Conversely,
the pollinator serves as an allogenic engineer, effect-
ing change through various activities (Figure 1)
(Cardoso et al., 2023).
Habitat Construction and Maintenance
Bees from the Andrenidae, Fideliidae, Halicitidae,
Melittidae, Oxaeidae, and Stenotritidae families
build their nests underground across wide range of
habitats. These habitats include weathered sand-
stone, prehistoric walls, dense clay, and sandy
dunes of deserts and beaches (Custer, 1928; Roubik
& Roubik, 1992; Stephen, 1965). Most species prefer
soils that are well-drained, ranging from horizontal
to vertical orientations for nesting (Linsley, 1958).
However, species such as Epicharis, Dasypoda, and
Nomia, are well-known for building nests in areas
prone to submersion. In populated areas, solitary
Science Reviews - Biology, 2023, 2(4), 1-6 Shahmshad Ahmed Khan
3
bees select nesting sites based on specific surface
characteristics. For instance, Nomia melanderi, pre-
fers soils with less than 8% clay content and mini-
mal surface water seepage. This species digs deeper
nests in hotter conditions to regulate temperature
(Stephen, 1960). Similarly, species like Colletes, An-
drena, and halictine bees often nest in soil with thin-
ner organic layers compared to adjacent areas (Os-
good Jr, 1972).
Nutrient Cycling and Soil Fertility
Bees contribute to nutrient cycling and soil fertility,
in addition to their well-known role in pollinating
diverse crops. In many terrestrial ecosystems, nitro-
gen is vital and often limiting factor for plant
growth and productivity. Research has shown that
nitrogen supplementation can substantially in-
crease both growth and productivity in plants (Vi-
tousek & Howarth, 1991). Honeybees feces, rich in
organic nitrogen, decomposes in the soil, releasing
inorganic nitrogen that plants can readily assimilate
(Mishra, Afik, Cabrera, Delaplane, & Mowrer,
2013). The wild bee, Osmia bicornis has been identi-
fied as a significant participant in the nutrient cycle.
Biogeochemical niche analysis has underscored its
role in this process. Pollen, the principal diet of bees,
is a potent contributor to nutrient cycling due to its
dense nutritional profile, which provides essential
nutrients. The death of O. bicornis individuals also
contributes to the flow of nutrients within ecosys-
tems (Filipiak, 2023).
Figure 1: Schematic diagram of bees as ecosystem engineers.
Conservation Implications and Strategies
Bees, as important pollinators for both wild and cul-
tivated crops, serve as ecosystem engineers in ter-
restrial environments (Klein et al., 2007). Despite
their ecological significance, bees are declining, pri-
marily due to anthropogenic activities (Biesmeijer et
al., 2006; Fitzpatrick et al., 2007). Habitat loss
emerges as the major factor in this decline, with
habitat fragmentation exacerbating the situation by
causing genetic isolation and inbreeding (Zayed,
2009). The potential decline of bee populations
poses a substantial threat to ecosystem services,
leading to concerns about disastrous impacts on the
ecosystem functioning. This has prompted action
from researchers, educators, and policymakers
(Brown & Paxton, 2009; Dicks et al., 2013; Potts et
al., 2016). Effective conservation strategies should
prioritize mitigate habitat loss, promoting pollina-
tor-friendly agricultural practices, and enhancing
public awareness about the bees. The presence of
natural patches of flowering plants can positively
contribute to the species richness and abundance of
Shahmshad Ahmed Khan Science Reviews - Biology, 2023, 2(4), 1-6
4
bees by providing a continuous source of pollen,
nectar, and shelter (Ruiz-Toledo, Vandame, Penilla-
Navarro, Gómez, & Sánchez, 2020). A study con-
ducted in Poland found that six ornamental flower-
ing plants in the city of Lublin, each with different
corolla lengths, attracted various species of pollina-
tors (Jachuła, Denisow, & Strzałkowska-Abramek,
2019). Flowering plants in city gardens and parks
act as a refuges for most pollinators, catering to their
relatively small requirements such as habitat range,
life cycle, and nesting behavior (Hall et al., 2017).
Additional measures should focus on assessing the
effects of invasive bee species on native bees and
understanding the potential impacts of climatic
changes on bee diversity and abundance.
Conclusion
Bees are indispensable not only as pollinators of
crops but also as ecosystem engineers, providing a
range of multifaceted services. Beyond pollination,
bees contribute to the ecosystem by creating habi-
tats, such as hiding places for various organisms in
abandoned fruits after seed dispersal. Additionally,
they influence soil composition and aeration
through their burrowing activities, which enhances
plant root development. However, the full scope of
bees’ role as ecosystem engineers remains underex-
plored. This gap presents a critical area for future
research to further elucidate their contributions to
environmental processes and biodiversity.
References
1. Ahmad, S., Khalofah, A., Khan, S. A., Khan, K. A., Jilani, M. J., Hussain, T., . . . Ahmad, Z. (2021).
Effects of native pollinator communities on the physiological and chemical parameters of loquat tree
(Eriobotrya japonica) under open field condition. Saudi Journal of Biological Sciences, 28(6), 3235-3241.
2. Aizen, M. A., Aguiar, S., Biesmeijer, J. C., Garibaldi, L. A., Inouye, D. W., Jung, C., . . . Ngo, H.
(2019). Global agricultural productivity is threatened by increasing pollinator dependence without a
parallel increase in crop diversification. Global change biology, 25(10), 3516-3527.
3. Aizen, M. A., & Harder, L. D. (2009). The global stock of domesticated honey bees is growing slower
than agricultural demand for pollination. Current biology, 19(11), 915-918.
4. Biesmeijer, J. C., Roberts, S. P., Reemer, M., Ohlemuller, R., Edwards, M., Peeters, T., . . . Thomas,
C. (2006). Parallel declines in pollinators and insect-pollinated plants in Britain and the Netherlands.
Science, 313(5785), 351-354.
5. Brown, M. J., & Paxton, R. J. (2009). The conservation of bees: a global perspective. Apidologie, 40(3),
410-416.
6. Byers, J. E. (2022). Using ecosystem engineers to enhance multiple ecosystem processes.
Functional Ecology, 1-15.
7. Cardoso, J. C., Rezende, U. C., Caetano, A. P., & Oliveira, P. E. (2023). Pollinators and plants as
ecosystem engineers: post‐dispersal fruits provide new habitats for other organisms. Oikos, e09819.
8. Casas-Crivillé, A., & Valera, F. (2005). The European bee-eater (Merops apiaster) as an ecosystem
engineer in arid environments. Journal of Arid Environments, 60(2), 227-238.
9. Custer, C. P. (1928). The bee that works in stone; Perdita opuntiae Cockerell. Psyche: A Journal of
Entomology, 35, 67-84.
10. Dicks, L. V., Abrahams, A., Atkinson, J., Biesmeijer, J., Bourn, N., Brown, C., . . . Cresswell, J. E.
(2013). Identifying key knowledge needs for evidence‐based conservation of wild insect pollinators:
collaborative cross‐sectoral exercise. Insect Conservation and Diversity, 6(3), 435-446.
11. Filipiak, M. (2023). Wild bee is an important contributor to the nutrient cycle, as shown by
biogeochemical niche analysis of Osmia bicornis. Available at SSRN 4591616.
12. Fitzpatrick, Ú., Murray, T. E., Paxton, R. J., Breen, J., Cotton, D., Santorum, V., & Brown, M. J.
(2007). Rarity and decline in bumblebees–a test of causes and correlates in the Irish fauna. Biological
conservation, 136(2), 185-194.
Science Reviews - Biology, 2023, 2(4), 1-6 Shahmshad Ahmed Khan
5
13. Gill, R. J., Baldock, K. C., Brown, M. J., Cresswell, J. E., Dicks, L. V., Fountain, M. T., . . . Holland,
J. M. (2016). Protecting an ecosystem service: approaches to understanding and mitigating threats to
wild insect pollinators. In Advances in ecological research (Vol. 54, pp. 135-206): Elsevier.
14. Gunders, D., & Bloom, J. (2017). Wasted: How America is losing up to 40 percent of its food
from farm to fork to landfill.
15. Hall, D. M., Camilo, G. R., Tonietto, R. K., Ollerton, J., Ahrné, K., Arduser, M., . . . Frankie, G.
(2017). The city as a refuge for insect pollinators. Conservation Biology, 31(1), 24-29.
16. Hristov, P., Neov, B., Shumkova, R., & Palova, N. (2020). Significance of apoidea as main
pollinators. ecological and economic impact and implications for human nutrition. Diversity, 12(7),
280.
17. Jachuła, J., Denisow, B., & Strzałkowska-Abramek, M. (2019). Floral reward and insect visitors in
six ornamental Lonicera species–plants suitable for urban bee-friendly gardens. Urban Forestry &
Urban Greening, 44, 126390.
18. Jones, C. G., Lawton, J. H., & Shachak, M. (1994). Organisms as ecosystem engineers. Oikos, 373-
386.
19. Jones, C. G., Lawton, J. H., & Shachak, M. (1997). Positive and negative effects of organisms as
physical ecosystem engineers. Ecology, 78(7), 1946-1957.
20. Khan, S. A., Tanveer, M., Ahmad, S., Mars, M., Naeem, M., Naveed, Z., . . . Goulson, D. (2022).
Declining abundance of pollinating insects drives falls in loquat (Eriobotrya japonica) fruit yields in
the Pothwar region of Pakistan. Agriculture, Ecosystems & Environment, 339, 108138.
21. Klatt, B. K., Holzschuh, A., Westphal, C., Clough, Y., Smit, I., Pawelzik, E., & Tscharntke, T. (2014).
Bee pollination improves crop quality, shelf life and commercial value. Proceedings of the royal society
B: biological sciences, 281(1775), 20132440.
22. Klein, A.-M., Boreux, V., Fornoff, F., Mupepele, A.-C., & Pufal, G. (2018). Relevance of wild and
managed bees for human well-being. Current Opinion in Insect Science, 26, 82-88.
23. Klein, A.-M., Vaissière, B. E., Cane, J. H., Steffan-Dewenter, I., Cunningham, S. A., Kremen, C., &
Tscharntke, T. (2007). Importance of pollinators in changing landscapes for world crops. Proceedings
of the royal society B: biological sciences, 274(1608), 303-313.
24. Kremen, C., & Chaplin-Kramer, R. (2007). Insects as providers of ecosystem services: crop pollination
and pest control. Paper presented at the Insect conservation biology: proceedings of the royal
entomological society’s 23rd symposium.
25. Kremen, C., James, R., & Pitts-Singer, T. (2008). Crop pollination services from wild bees. Bee
pollination in agricultural ecosystems, 10-26.
26. Lawton, J. H., & Jones, C. G. (1995). Linking species and ecosystems: organisms as ecosystem
engineers. In Linking species & ecosystems (pp. 141-150): Springer.
27. Linsley, E. (1958). The ecology of solitary bees. Hilgardia, 27(19), 543-599.
28. Mills, L. S., Soulé, M. E., & Doak, D. F. (1993). The keystone-species concept in ecology and
conservation. BioScience, 43(4), 219-224.
29. Mishra, A., Afik, O., Cabrera, M. L., Delaplane, K. S., & Mowrer, J. E. (2013). Inorganic nitrogen
derived from foraging honey bees could have adaptive benefits for the plants they visit. PloS one, 8(7),
e70591.
30. Ollerton, J. (2017). Pollinator diversity: distribution, ecological function, and conservation.
Annual review of ecology, evolution, and systematics, 48, 353-376.
Shahmshad Ahmed Khan Science Reviews - Biology, 2023, 2(4), 1-6
6
31. Osgood Jr, E. (1972). Soil characteristics of nesting sites of native bees associated with the low-
bush blueberry in Maine. Maine Agrie. Exp. Stn. Tech. Bull, 59, 1-8.
32. Patel, V., Pauli, N., Biggs, E., Barbour, L., & Boruff, B. (2021). Why bees are critical for achieving
sustainable development. Ambio, 50, 49-59.
33. Potts, S. G., Imperatriz-Fonseca, V., Ngo, H. T., Aizen, M. A., Biesmeijer, J. C., Breeze, T. D., . . .
Settele, J. (2016). Safeguarding pollinators and their values to human well-being. Nature, 540(7632),
220-229.
34. Roubik, D. W., & Roubik, D. W. (1992). Ecology and natural history of tropical bees: Cambridge
University Press.
35. Ruiz-Toledo, J., Vandame, R., Penilla-Navarro, P., Gómez, J., & Sánchez, D. (2020). Seasonal
abundance and diversity of native bees in a patchy agricultural landscape in Southern Mexico.
Agriculture, Ecosystems & Environment, 292, 106807.
36. Ruschioni, S., Riolo, P., Minuz, R. L., Stefano, M., Cannella, M., Porrini, C., & Isidoro, N. (2013).
Biomonitoring with honeybees of heavy metals and pesticides in nature reserves of the Marche
Region (Italy). Biological trace element research, 154, 226-233.
37. Sadeghi, A., Mozafari, A.-A., Bahmani, R., & Shokri, K. (2012). Use of honeybees as bio-indicators
of environmental pollution in the Kurdistan province of Iran. Journal of Apicultural Science, 56(2), 83-
88.
38. Senapathi, D., Biesmeijer, J. C., Breeze, T. D., Kleijn, D., Potts, S. G., & Carvalheiro, L. G. (2015).
Pollinator conservation—the difference between managing for pollination services and preserving
pollinator diversity. Current Opinion in Insect Science, 12, 93-101.
39. Skorbiłowicz, E., Skorbiłowicz, M., & Cieśluk, I. (2018). Bees as bioindicators of environmental
pollution with metals in an urban area. Journal of Ecological Engineering, 19(3), 229-234.
40. Stephen, W. (1960). Artificial bee beds for the propagation of the Alkali bee, Nomia melander.
Journal of Economic Entomology, 53(6), 1025-1030.
41. Stephen, W. (1965). Effects of soil moisture on survival of prepupae of the alkali bee. Journal of
Economic Entomology, 58(3), 472-474.
42. Vitousek, P. M., & Howarth, R. W. (1991). Nitrogen limitation on land and in the sea: how can it
occur? Biogeochemistry, 13, 87-115.
43. Wilby, A. (2002). Ecosystem engineering: a trivialized concept? Trends in Ecology & Evolution, 17(7),
307.
44. Zayed, A. (2009). Bee genetics and conservation. Apidologie, 40(3), 237-262.