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Adaptive Modification of Appendages for Grooming (Cleaning, Antifouling) and Reproduction in the Crustacea
Abstract
Appendages used primarily for feeding and locomotion have become secondarily modified for grooming and reproductive purposes in many crustaceans. Crustaceans use specialized brushes and combs composed of setae with a complex microstructure for scraping surfaces clean. Decapod crustaceans have the best-described cleaning behavior, with gill cleaning by a variety of mechanisms necessitated by enclosure of gills in a branchial chamber. Other crustaceans, particularly stomatopods, some peracarids, and ostracods, groom frequently. Some crustaceans avoid algal fouling by frequent molting, burrowing in abrasive sediments, or nocturnal behavior. In many species, appendages have also experienced specializations for reproductive purposes. The immobile sperm of crustaceans must be actively transferred by the male in crustaceans, and a variety of appendages have become modified for this task. The actual mechanics of reproductive appendages in crustaceans are still poorly known and remain a fertile topic for study.
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“During this molt, pleopods become modified, with more developed setae, which is related to their role in holding the eggs during the brooding period (Correa and Thiel, 2003; Bauer, 2013).”
From Is It First the Egg or the Shrimp? – Diversity and Variation in Microbial Communities Colonizing Broods of the Vent Shrimp Rimicaris exoculata During Embryonic Development (2019) · cites this paper as Bauer, 2013
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Baeza, J. A., Simpson, L., Ambrosio, L. J., Mora, N., Guéron, R., & Childress, M. J. (2016) · BMC Zoology · doi:10.1186/s40850-016-0006-6 · Cited by 36
“For instance, grooming of the brood mass by parental females increases embryo survival and hatching rates [14, 27].”
BackgroundMethou, P., Hernández-Ávila, I., Aube, J., Cueff-Gauchard, V., Gayet, N., Amand, L., Shillito, B., Pradillon, F., & Cambon-Bonavita, M.-A. (2019) · Frontiers in Microbiology · doi:10.3389/fmicb.2019.00808 · Cited by 30
“During this molt, pleopods become modified, with more developed setae, which is related to their role in holding the eggs during the brooding period (Correa and Thiel, 2003; Bauer, 2013).”
BackgroundTiny individuals attached to a new Silurian arthropod suggest a unique mode of brood care
Briggs, D. E. G., Siveter, D. J., Siveter, D. J., Sutton, M. D., & David, L. (2016) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1600489113 · Cited by 27
“Furthermore, the “host” arthropod was clearly living when the capsules became attached: it is unlikely to have tolerated the presence of so many drag-inducing epizoans, and head appendage 1 is long enough to have cleaned the trunk tergites (“general body grooming” as in some living crustaceans) (20).”
BackgroundBalázs, G., Biró, A., Fišer, Ž., et al. (2021) · Ecology and Evolution · doi:10.1002/ece3.8233 · Cited by 22
VanMaurik, L., & Wortham, J. L. (2014) · ZooKeys · doi:10.3897/zookeys.457.6292 · Cited by 17
Kamio, M., Furukawa, D., Wakabayashi, K., et al. (2014) · Journal of Experimental Marine Biology and Ecology · doi:10.1016/j.jembe.2014.11.008 · Cited by 13
Pescinelli, R. A., Davanso, T. M., Pantaleão, J. A. F., et al. (2017) · Journal of the Marine Biological Association of the United Kingdom · doi:10.1017/s0025315416001880 · Cited by 11
Self-citationBrix, S., Held, C., Kaiser, S., Jennings, R. M., Driskell, A. C., & Brandt, A. (2021) · Organisms Diversity & Evolution · doi:10.1007/s13127-021-00509-9 · Cited by 10
“Functionally, the first pereopod is used for feeding and grooming (Bauer, 2013; Hessler & Strömberg, 1989).”
BackgroundGregati, R. A., Fransozo, V., Greco, L. S. L., et al. (2014) · Invertebrate Biology · doi:10.1111/ivb.12067 · Cited by 10
“However, such information is still scarce compared with current knowledge on the same subjects in terrestrial arthropods (Bauer 1992, 2004, 2013).”
BackgroundSelf-citationThe role of the male copulatory organs in the colonization of the pelagic by shrimp-like eucarids
Lunina, A. A., & Vereshchaka, A. L. (2016) · Deep Sea Research Part II Topical Studies in Oceanography · doi:10.1016/j.dsr2.2016.11.018 · Cited by 10
Middlemiss, K. L., Urbina, M. A., & Wilson, R. W. (2015) · Helgoland Marine Research · doi:10.1007/s10152-015-0445-4 · Cited by 9
“Bauer (2013) suggests, however, that not all ‘passive’ grooming is purely linked to normal locomotory/feeding movements and that the act of ‘‘limb rocking’’ (movement caused by the animal actively rocking backward and forwards on the pereopods) may be a specific attempt to engage the setae.”
BackgroundInfluentialGrooming behaviors and setal morphology in smasher and spearer mantis shrimps (Stomatopoda)
Wortham, J. L., & Kostecka, L. G. (2018) · Journal of Crustacean Biology · doi:10.1093/jcbiol/ruy096 · Cited by 9
Baeza, J. A., Liu, X., Kostecka, L. G., et al. (2019) · Marine Biology · doi:10.1007/s00227-019-3579-0 · Cited by 7
“Previous studies in gonochoric decapods have demonstrated that APC not only provides benefits (Bauer 1989, 2013; Förster and Baeza 2001; Baeza and Fernández 2002; Bauer 2013) but also results in metabolic costs (Fernández et al.”
BackgroundInfluentialAhmad, M., Ismail, N., & Eldeen, I. M. S. (2020) · Marine Biology Research · doi:10.1080/17451000.2020.1840592 · Cited by 5
van Viersen, A. P., & Kloc, G. J. (2022) · Geologica Belgica · doi:10.20341/gb.2022.005 · Cited by 5
van Viersen, A. P., Poschmann, M., Langer, M. R., et al. (2025) · Geologica Belgica · doi:10.20341/gb.2025.010 · Cited by 2
Tan, M. M. J., Tudge, C. C., Penna-Díaz, M. A., et al. (2020) · Reproductive Biology · doi:10.1093/oso/9780190688554.003.0003 · Cited by 2
Dakin, M., & Wortham, J. L. (2023) · Journal of Crustacean Biology · doi:10.1093/jcbiol/ruad029 · Cited by 2
Triay‐Portella, R., González, J. A., Biscoito, M., et al. (2018) · Deep Sea Research Part I Oceanographic Research Papers · doi:10.1016/j.dsr.2018.10.005 · Cited by 2
“The real mechanism of reproductive appendages is still poorly known in crustaceans and remains a fertile topic for study (Bauer, 2013).”
BackgroundInfluentialWortham, J. L., & VanMaurik, L. (2019) · Journal of Crustacean Biology · doi:10.1093/jcbiol/ruz079 · Cited by 2
Size at sexual maturity of male Lucensosergia lucens (Decapoda: Sergestidae) in Suruga Bay, Japan
Doi, W., Hirofumi, W., & Suzuki, N. (2021) · Crustacean Research · doi:10.18353/crustacea.51.0_1 · Cited by 2
Multiple Matings and Sperm Competition
Becker, C., & Bauer, R. T. (2020) · Reproductive Biology · doi:10.1093/oso/9780190688554.003.0012 · Cited by 2
Self-citationDuffy, R., Hodgson, B., Quinn, A., et al. (2025) · Marine and Freshwater Research · doi:10.1071/mf25063 · Cited by 1
Petrova, M., & Bogomolova, E. (2025) · Marine Biology · doi:10.1007/s00227-025-04707-3 · Cited by 1
“These crustaceans carry embryo masses using specialised append-ages, and some also have external fertilisation (reviewed in Bauer 2013 ).”
InfluentialStephens, N. C., Behringer, D. C., Porter, R., Baker, A. M., & Baeza, J. A. (2025) · Marine Biology · doi:10.1007/s00227-025-04719-z · Cited by 2
“…parasites become established on their host, mitigation behaviors are expected to help mediate the negative effects of parasite infection and, thereafter, be maintained by selection for removing parasites when detected (Poulin 1995; Bauer 2013; Moore 2013; Vale et al. 2018; Gibson and Amoroso 2022).”
Shrimp Hygiene: Antifouling Adaptations
Bauer, R. T. (2023) · Fish & fisheries series/Fish and fisheries series (Print) · doi:10.1007/978-3-031-20966-6_4
Self-citationBobkov, Y. V., Strickler, J. R., & Derby, C. D. (2026) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.64898/2026.01.20.700623
Bíró, A., Balázs, G., Horváth, G., Fišer, Ž., Fišer, C., Sarbu, S. M., & Herczeg, G. (2026) · Frontiers in Ecology and Evolution · doi:10.3389/fevo.2026.1748446
“…on pereopod I (propodus length, propodus width, dactylus seta row length, dactylus length, and spine length) are involved in feeding and grooming (except spine length) and probably have a role in both interspeci fi c and intraspeci fi c aggressive interactions (Bertin et al., 2002; Bauer, 2013).”
Methou, P. (2019) · theses.fr (ABES) · doi:10.70675/bf7aab7fzc361z40c2z9ac6z34c7f7b317db · Cited by 1
Metadata
- Authors
- Bauer, R. T.
- Corresponding author
- Bauer, Raymond T.
- ORCID
- Bauer, Raymond T.
- Venue
- Functional Morphology and Diversity
- Publisher
- Oxford University Press
- Published
- 2013-01-16
- Pages
- 337-375
- OpenAlex
- W2492291302
- Cited by
- 29
- References
- 0
- Citations by year
- 2026: 2 · 2025: 4 · 2023: 2 · 2022: 1 · 2021: 3 · 2020: 3 · 2019: 4 · 2018: 2 · 2017: 1 · 2016: 3 · 2015: 1 · 2014: 3
- FWCI
- 5.47
- Type
- book chapter
- Language
- en
- Open access
- closed
- Locations
- Oxford University Press eBooks (publishedVersion)
- Topics
- Physical Sciences › Environmental Science › Ecology › Crustacean biology and ecology
- Physical Sciences › Earth and Planetary Sciences › Oceanography › Marine Biology and Ecology Research
- Health Sciences › Medicine › Anatomy › Comparative Animal Anatomy Studies
- Keywords
- grooming, antifouling, Decapoda, mantis shrimp, Peracarida, Ostracoda, molting, burrowing, nocturnal activity, sperm transfer
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