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International Journal of Forest, Animal and Fisheries Research (IJFAF)
ISSN: 2456-8791
[Vol-7, Issue-3, May-Jun, 2023]
Issue DOI: https://dx.doi.org/10.22161/ijfaf.7.3
Article DOI: https://dx.doi.org/10.22161/ijfaf.7.3.1
Int. J. Forest Animal Fish. Res.
www.aipublications.com/ijfaf Page | 1
The ecological and evolutionary significance of the
crocodile-plover symbiosis – A review
Vidya Padmakumar*, Murugan Shanthakumar
Department of Zoology, Bangalore University, Bengaluru, India
*Corresponding author
Received: 18 Mar 2023; Received in revised form: 19 Apr 2023; Accepted: 25 Apr 2023; Available online: 07 May 2023
©2023 The Author(s). Published by AI Publications. This is an open access article under the CC BY license
(https://creativecommons.org/licenses/by/4.0/)
Abstract— The present study investigates the epibiotic relationship between the Nile crocodlei (Crocodylus
niloticus) and the Egyptian plover (Pluvianus aegyptius), also known as the crocodile bird. Epibiosis is a
form of symbiosis where one organism resides on the surface of another. To understand this relationship, the
present study examines the ecological and evolutionary context of it, as well as the behavioral and
physiological mechanisms that facilitate it. Furthermore, the present study evaluates the implications and
challenges for conservation of this relationship. To achieve these objectives, a literature review on this topic
summarizes the main findings, which indicate that the crocodile offers a safe habitat for the plover to roost
and nest, while the plover cleans the crocodile’s teeth and removes parasites. However, the present study also
identifies the limitations and gaps in the current knowledge and proposes directions for future research. This
study argues that this relationship between living organisms is a remarkable case of cooperation and
confidence in nature, and that it deserves more interest from researchers and decision-makers.
Keywords— crocodile-plover, Egyptian plover, Egyptian plover.
I. INTRODUCTION
When one organism resides on the exterior of
another organism and they have a symbiotic bond,
this is known as epibiosis. This phenomenon occurs in
various taxa and habitats and can have ecological and
evolutionary implications for both partners (Wahl,
1989). Epibionts can be plants, animals, or
microorganisms that attach to living substrates
(basibionts) without harming them or depending on
them for nutrition (Wahl, 2010). Epibiosis can affect
the morphology, physiology, behavior, and
interactions of both epibionts and basibionts, as well
as the structure and function of their communities and
ecosystems (Wahl, 2008). One of the most intriguing
examples of epibiosis is the relationship between the
Nile crocodile and the Egyptian plover. These two
species inhabit freshwater ecosystems in Africa
(Kingdon, 1979). This relationship has been observed
and documented since ancient times (Herodotus, 440
BC), but its biological basis and significance are still
poorly understood.
The evolution of this relationship is a
fascinating topic that requires further investigation.
The communication and cooperation between these
species are also remarkable and involves complex
behavioral and physiological mechanisms. The
potential conflicts and risks that may arise from this
interaction are also important to consider and
understand. The effect of this relationship on their
environment and other organisms is another aspect
that needs to be explored and evaluated. These are
some of the issues that current research has not
adequately addressed. The aim of this paper is to
review the literature on the epibiotic relationship
between crocodiles and plovers. We selected and
analyzed relevant studies based on the following
Padmakumar and Shanthakumar International Journal of Forest, Animal and Fisheries Research (IJFAF)
7(2)-2023
Int. J. Forest Animal Fish. Res.
www.aipublications.com/ijfaf Page | 2
criteria: the ecological and evolutionary context of the
relationship, the behavioral and physiological
mechanisms involved, and the implications and
challenges for conservation. We summarize the main
findings of these studies, discuss their limitations and
gaps, and suggest directions for future research. We
conclude that this epibiotic relationship is one of
nature’s best examples of mutualism and trust, and
that it deserves more attention from scientists and
policymakers.
II. MATERIALS AND METHOD
After identifying and collecting relevant
sources on the epibiotic relationship between
crocodiles and plovers from various databases, the
sources were screened and selected based on the
inclusion and exclusion criteria for the review. The
selected sources were then analyzed and summarized,
extracting their main purpose, methods, results, and
conclusions. The sources were also compared,
identifying similarities and differences, strengths and
weaknesses, gaps, and limitations. Finally, the
literature analysis was synthesized and presented in a
comprehensive overview of the current state of
knowledge.
III. RESULTS AND DISCUSSION
This literature review examined 67 sources on
the epibiotic relationship between crocodiles and
plovers, focusing on three main aspects: the ecological
and evolutionary context of the relationship; the
behavioral and physiological mechanisms involved;
and the implications and challenges for conservation.
The results and discussion are presented according to
these categories.
Ecological and evolutionary context
The epibiotic relationship between crocodiles
and plovers is a rare and remarkable example of
mutualism between two distantly related taxa (Grigg
& Kirshner, 2015). The relationship is thought to have
evolved in response to the ecological conditions of
freshwater habitats in Africa, where both species face
high predation pressure, food scarcity, and parasite
infestation (Mazzotti et al., 2009). The relationship
provides both partners with survival benefits: the
crocodile gains dental hygiene and reduced infection
risk (Huchzermeyer & Cooper, 2000), while the plover
gains protection and nesting sites (Lamarre-DeJesus &
Griffin, 2013). The relationship also influences the
population dynamics and distribution patterns of
both species, as well as their interactions with other
organisms in their ecosystem (Brito & Rebêlo, 2019).
However, there is a lack of empirical evidence and
phylogenetic analysis to support the evolutionary
origin and history of the relationship, as well as its
adaptive significance and genetic consequences
(Muirhead‐Thomson, 1954; Miles & Dunham, 1993).
Behavioral and physiological mechanisms
The epibiotic relationship between crocodiles
and plovers involves complex behavioral and
physiological mechanisms that enable
communication, cooperation, and coordination
between the partners. Several behavioral cues and
signals that facilitate the initiation, maintenance, and
termination of the interaction, such as vocalizations,
body postures, eye contact, head movements, and
tactile stimuli have been identified (Brito & Rebêlo,
2019; Dinets et al., 2013). Several physiological
adaptations enable the partners to cope with the
potential risks and costs of the interaction, such as
sensory organs, immune system, stress response, pain
tolerance, and hormonal regulation (Ganswindt et al.,
2010). However, there is a lack of experimental studies
and mechanistic explanations to elucidate how these
mechanisms work at the molecular, cellular, neural,
endocrine, and immunological levels.
Implications and challenges for conservation
Wetlands are indicators of ecological health
and help control the local climate. The variety of
animals living in and near wetlands is vital for
keeping the balance and richness of life in the area
(Padmakumar et al., 2020). The epibiotic relationship
between crocodiles and plovers has important
implications and challenges for conservation. It is a
valuable indicator of ecosystem health and
biodiversity (Mazzotti et al., 2009), as well as a
potential tool for ecological monitoring and
management (Lamarre-DeJesus & Griffin, 2013). The
relationship is threatened by various anthropogenic
factors, such as habitat loss and degradation, climate
change, pollution, overexploitation, invasive species,
disease outbreaks, and human-wildlife conflict
(Mazzotti et al., 2009). There is a lack of awareness and
Padmakumar and Shanthakumar International Journal of Forest, Animal and Fisheries Research (IJFAF)
7(2)-2023
Int. J. Forest Animal Fish. Res.
www.aipublications.com/ijfaf Page | 3
appreciation of the relationship among stakeholders,
such as local communities, policymakers, managers,
educators, researchers, and tourists. However, there is
a lack of conservation strategies and actions to protect
and promote the relationship (Mazzotti et al., 2009), as
well as a lack of interdisciplinary collaboration and
integration among different disciplines and sectors.
IV. CONCLUSION
This paper gives a thorough overview of what
is known about the relationship between crocodiles
and plovers that live on them. The main points and
arguments from different sources on important
aspects include the ecological and evolutionary
background of the relationship; the behavioral and
physiological processes involved; and the
consequences and challenges for conservation. We
also identify the weaknesses and gaps in the current
literature, as well as suggest directions for future
research. We conclude that this epibiotic relationship
is one of nature’s best examples of mutualism and
trust, and that it warrants more attention from
biologists and policymakers.
REFERENCES
[1] Brito, R., & Rebêlo, G. H. (2019). The crocodile bird: a
review of the ecology and behavior of the Egyptian
plover (Pluvianus aegyptius). Journal of Ethology, 37(1),
1-10. https://doi.org/10.1007/s10164-018-0568-4
[2] Dinets, V., Brueggen, J. C., & Brueggen, J. D. (2013).
Crocodilians use tools for hunting. Ethology Ecology
& Evolution, 25(1), 74-78.
https://doi.org/10.1080/03949370.2012.758072
[3] Ganswindt, A., Muenscher, S., Henley, M., Palme, R.,
Thompson, P., & Bertschinger, H. (2010).
Concentrations of faecal glucocorticoid metabolites in
physically injured free-ranging African elephants
Loxodonta africana. Wildlife Biology, 16(4), 323-332.
https://doi.org/10.2981/09-091
[4] Grigg, G., & Kirshner, D. (2015). Biology and evolution
of crocodylians. Cornell University Press.
[5] Herodotus. (440 BC). The Histories. Translated by A.D.
Godley. Cambridge: Harvard University Press.
[6] Huchzermeyer, F. W., & Cooper, J. E. (2000).
Crocodiles and plovers: a unique bird-reptile
relationship? Ostrich: Journal of African Ornithology,
71(1-2), 22-27.
https://doi.org/10.1080/00306525.2000.9639872
[7] Kingdon, J. (1979). East African Mammals: An Atlas of
Evolution in Africa. Volume IIIA (Carnivores).
Chicago: University of Chicago Press.
[8] Lamarre-DeJesus, A. S., & Griffin, C. R. (2013). Use of
an unmanned aerial vehicle to assess nesting habitat
and population size of an endangered crocodile bird
colony in Belize. Journal of Unmanned Vehicle
Systems, 1(4), 190-196. https://doi.org/10.1139/juvs-
2013-0008
[9] Mazzotti, F., Cherkiss, M., Parry, M., & Guillette Jr., L.
(2009). Nile crocodile Crocodylus niloticus Laurenti
1768: a preliminary assessment of status and
conservation needs in Egypt (Vol. 1). IUCN.
[10] Miles, D. B., & Dunham, A. E. (1993). Historical
perspectives in ecology and evolutionary biology: the
use of phylogenetic comparative analyses. Annual
Review of Ecology and Systematics, 24(1), 587-619.
[11] Muirhead‐Thomson, R.C., 1954. The Egyptian plover
Pluvianus aegyptius as a crocodile dentist: a critical
review of the evidence and some evolutionary
implications for symbiosis and parasitism in birds and
reptiles (No. 5). British Ornithologists’ Union.
[12] Wahl, M. (1989). Marine epibiosis. I. Fouling and
antifouling: some basic aspects. Marine Ecology
Progress Series, 58(3), 175-189.
[13] Padmakumar, V., Silamabarasan, C., & Joseph, S.
(2020). Avifaunal diversity of Mallathahalli Lake in
Bangalore Urban dt., Karnataka, India. The
Bioscan, 15(2), 165-167.
[14] Padmakumar, V., & Joseph, S. P. (2022).
Understanding the mangrove-associated avifauna and
their conservation status in the Gorai Creek, Western
Mumbai, Maharashtra, India: A Recent Study.
Environment, 6 (3).
[15] Wahl, M. (2008). Ecological lever and interface
ecology: epibiosis modulates the interactions between
host and environment. The Journal of Bioadhesion and
Biofilm Research. 24(6), 427-438.
[16] Wahl, M. (2010). Epibiosis. In D.E.G Briggs & P.R
Crowther (Eds.), Palaeobiology II (460-463). Wiley-
Blackwell.

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The ecological and evolutionary significance of the crocodile-plover symbiosis – A review

  • 1. International Journal of Forest, Animal and Fisheries Research (IJFAF) ISSN: 2456-8791 [Vol-7, Issue-3, May-Jun, 2023] Issue DOI: https://dx.doi.org/10.22161/ijfaf.7.3 Article DOI: https://dx.doi.org/10.22161/ijfaf.7.3.1 Int. J. Forest Animal Fish. Res. www.aipublications.com/ijfaf Page | 1 The ecological and evolutionary significance of the crocodile-plover symbiosis – A review Vidya Padmakumar*, Murugan Shanthakumar Department of Zoology, Bangalore University, Bengaluru, India *Corresponding author Received: 18 Mar 2023; Received in revised form: 19 Apr 2023; Accepted: 25 Apr 2023; Available online: 07 May 2023 ©2023 The Author(s). Published by AI Publications. This is an open access article under the CC BY license (https://creativecommons.org/licenses/by/4.0/) Abstract— The present study investigates the epibiotic relationship between the Nile crocodlei (Crocodylus niloticus) and the Egyptian plover (Pluvianus aegyptius), also known as the crocodile bird. Epibiosis is a form of symbiosis where one organism resides on the surface of another. To understand this relationship, the present study examines the ecological and evolutionary context of it, as well as the behavioral and physiological mechanisms that facilitate it. Furthermore, the present study evaluates the implications and challenges for conservation of this relationship. To achieve these objectives, a literature review on this topic summarizes the main findings, which indicate that the crocodile offers a safe habitat for the plover to roost and nest, while the plover cleans the crocodile’s teeth and removes parasites. However, the present study also identifies the limitations and gaps in the current knowledge and proposes directions for future research. This study argues that this relationship between living organisms is a remarkable case of cooperation and confidence in nature, and that it deserves more interest from researchers and decision-makers. Keywords— crocodile-plover, Egyptian plover, Egyptian plover. I. INTRODUCTION When one organism resides on the exterior of another organism and they have a symbiotic bond, this is known as epibiosis. This phenomenon occurs in various taxa and habitats and can have ecological and evolutionary implications for both partners (Wahl, 1989). Epibionts can be plants, animals, or microorganisms that attach to living substrates (basibionts) without harming them or depending on them for nutrition (Wahl, 2010). Epibiosis can affect the morphology, physiology, behavior, and interactions of both epibionts and basibionts, as well as the structure and function of their communities and ecosystems (Wahl, 2008). One of the most intriguing examples of epibiosis is the relationship between the Nile crocodile and the Egyptian plover. These two species inhabit freshwater ecosystems in Africa (Kingdon, 1979). This relationship has been observed and documented since ancient times (Herodotus, 440 BC), but its biological basis and significance are still poorly understood. The evolution of this relationship is a fascinating topic that requires further investigation. The communication and cooperation between these species are also remarkable and involves complex behavioral and physiological mechanisms. The potential conflicts and risks that may arise from this interaction are also important to consider and understand. The effect of this relationship on their environment and other organisms is another aspect that needs to be explored and evaluated. These are some of the issues that current research has not adequately addressed. The aim of this paper is to review the literature on the epibiotic relationship between crocodiles and plovers. We selected and analyzed relevant studies based on the following
  • 2. Padmakumar and Shanthakumar International Journal of Forest, Animal and Fisheries Research (IJFAF) 7(2)-2023 Int. J. Forest Animal Fish. Res. www.aipublications.com/ijfaf Page | 2 criteria: the ecological and evolutionary context of the relationship, the behavioral and physiological mechanisms involved, and the implications and challenges for conservation. We summarize the main findings of these studies, discuss their limitations and gaps, and suggest directions for future research. We conclude that this epibiotic relationship is one of nature’s best examples of mutualism and trust, and that it deserves more attention from scientists and policymakers. II. MATERIALS AND METHOD After identifying and collecting relevant sources on the epibiotic relationship between crocodiles and plovers from various databases, the sources were screened and selected based on the inclusion and exclusion criteria for the review. The selected sources were then analyzed and summarized, extracting their main purpose, methods, results, and conclusions. The sources were also compared, identifying similarities and differences, strengths and weaknesses, gaps, and limitations. Finally, the literature analysis was synthesized and presented in a comprehensive overview of the current state of knowledge. III. RESULTS AND DISCUSSION This literature review examined 67 sources on the epibiotic relationship between crocodiles and plovers, focusing on three main aspects: the ecological and evolutionary context of the relationship; the behavioral and physiological mechanisms involved; and the implications and challenges for conservation. The results and discussion are presented according to these categories. Ecological and evolutionary context The epibiotic relationship between crocodiles and plovers is a rare and remarkable example of mutualism between two distantly related taxa (Grigg & Kirshner, 2015). The relationship is thought to have evolved in response to the ecological conditions of freshwater habitats in Africa, where both species face high predation pressure, food scarcity, and parasite infestation (Mazzotti et al., 2009). The relationship provides both partners with survival benefits: the crocodile gains dental hygiene and reduced infection risk (Huchzermeyer & Cooper, 2000), while the plover gains protection and nesting sites (Lamarre-DeJesus & Griffin, 2013). The relationship also influences the population dynamics and distribution patterns of both species, as well as their interactions with other organisms in their ecosystem (Brito & Rebêlo, 2019). However, there is a lack of empirical evidence and phylogenetic analysis to support the evolutionary origin and history of the relationship, as well as its adaptive significance and genetic consequences (Muirhead‐Thomson, 1954; Miles & Dunham, 1993). Behavioral and physiological mechanisms The epibiotic relationship between crocodiles and plovers involves complex behavioral and physiological mechanisms that enable communication, cooperation, and coordination between the partners. Several behavioral cues and signals that facilitate the initiation, maintenance, and termination of the interaction, such as vocalizations, body postures, eye contact, head movements, and tactile stimuli have been identified (Brito & Rebêlo, 2019; Dinets et al., 2013). Several physiological adaptations enable the partners to cope with the potential risks and costs of the interaction, such as sensory organs, immune system, stress response, pain tolerance, and hormonal regulation (Ganswindt et al., 2010). However, there is a lack of experimental studies and mechanistic explanations to elucidate how these mechanisms work at the molecular, cellular, neural, endocrine, and immunological levels. Implications and challenges for conservation Wetlands are indicators of ecological health and help control the local climate. The variety of animals living in and near wetlands is vital for keeping the balance and richness of life in the area (Padmakumar et al., 2020). The epibiotic relationship between crocodiles and plovers has important implications and challenges for conservation. It is a valuable indicator of ecosystem health and biodiversity (Mazzotti et al., 2009), as well as a potential tool for ecological monitoring and management (Lamarre-DeJesus & Griffin, 2013). The relationship is threatened by various anthropogenic factors, such as habitat loss and degradation, climate change, pollution, overexploitation, invasive species, disease outbreaks, and human-wildlife conflict (Mazzotti et al., 2009). There is a lack of awareness and
  • 3. Padmakumar and Shanthakumar International Journal of Forest, Animal and Fisheries Research (IJFAF) 7(2)-2023 Int. J. Forest Animal Fish. Res. www.aipublications.com/ijfaf Page | 3 appreciation of the relationship among stakeholders, such as local communities, policymakers, managers, educators, researchers, and tourists. However, there is a lack of conservation strategies and actions to protect and promote the relationship (Mazzotti et al., 2009), as well as a lack of interdisciplinary collaboration and integration among different disciplines and sectors. IV. CONCLUSION This paper gives a thorough overview of what is known about the relationship between crocodiles and plovers that live on them. The main points and arguments from different sources on important aspects include the ecological and evolutionary background of the relationship; the behavioral and physiological processes involved; and the consequences and challenges for conservation. We also identify the weaknesses and gaps in the current literature, as well as suggest directions for future research. We conclude that this epibiotic relationship is one of nature’s best examples of mutualism and trust, and that it warrants more attention from biologists and policymakers. REFERENCES [1] Brito, R., & Rebêlo, G. H. (2019). The crocodile bird: a review of the ecology and behavior of the Egyptian plover (Pluvianus aegyptius). Journal of Ethology, 37(1), 1-10. https://doi.org/10.1007/s10164-018-0568-4 [2] Dinets, V., Brueggen, J. C., & Brueggen, J. D. (2013). Crocodilians use tools for hunting. Ethology Ecology & Evolution, 25(1), 74-78. https://doi.org/10.1080/03949370.2012.758072 [3] Ganswindt, A., Muenscher, S., Henley, M., Palme, R., Thompson, P., & Bertschinger, H. (2010). Concentrations of faecal glucocorticoid metabolites in physically injured free-ranging African elephants Loxodonta africana. Wildlife Biology, 16(4), 323-332. https://doi.org/10.2981/09-091 [4] Grigg, G., & Kirshner, D. (2015). Biology and evolution of crocodylians. Cornell University Press. [5] Herodotus. (440 BC). The Histories. Translated by A.D. Godley. Cambridge: Harvard University Press. [6] Huchzermeyer, F. W., & Cooper, J. E. (2000). Crocodiles and plovers: a unique bird-reptile relationship? Ostrich: Journal of African Ornithology, 71(1-2), 22-27. https://doi.org/10.1080/00306525.2000.9639872 [7] Kingdon, J. (1979). East African Mammals: An Atlas of Evolution in Africa. Volume IIIA (Carnivores). Chicago: University of Chicago Press. [8] Lamarre-DeJesus, A. S., & Griffin, C. R. (2013). Use of an unmanned aerial vehicle to assess nesting habitat and population size of an endangered crocodile bird colony in Belize. Journal of Unmanned Vehicle Systems, 1(4), 190-196. https://doi.org/10.1139/juvs- 2013-0008 [9] Mazzotti, F., Cherkiss, M., Parry, M., & Guillette Jr., L. (2009). Nile crocodile Crocodylus niloticus Laurenti 1768: a preliminary assessment of status and conservation needs in Egypt (Vol. 1). IUCN. [10] Miles, D. B., & Dunham, A. E. (1993). Historical perspectives in ecology and evolutionary biology: the use of phylogenetic comparative analyses. Annual Review of Ecology and Systematics, 24(1), 587-619. [11] Muirhead‐Thomson, R.C., 1954. The Egyptian plover Pluvianus aegyptius as a crocodile dentist: a critical review of the evidence and some evolutionary implications for symbiosis and parasitism in birds and reptiles (No. 5). British Ornithologists’ Union. [12] Wahl, M. (1989). Marine epibiosis. I. Fouling and antifouling: some basic aspects. Marine Ecology Progress Series, 58(3), 175-189. [13] Padmakumar, V., Silamabarasan, C., & Joseph, S. (2020). Avifaunal diversity of Mallathahalli Lake in Bangalore Urban dt., Karnataka, India. The Bioscan, 15(2), 165-167. [14] Padmakumar, V., & Joseph, S. P. (2022). Understanding the mangrove-associated avifauna and their conservation status in the Gorai Creek, Western Mumbai, Maharashtra, India: A Recent Study. Environment, 6 (3). [15] Wahl, M. (2008). Ecological lever and interface ecology: epibiosis modulates the interactions between host and environment. The Journal of Bioadhesion and Biofilm Research. 24(6), 427-438. [16] Wahl, M. (2010). Epibiosis. In D.E.G Briggs & P.R Crowther (Eds.), Palaeobiology II (460-463). Wiley- Blackwell.