Habitat complexity is an important mediator of processes spanning levels of biological organization from organisms to ecosystems (Shumway et al. 2007, Soukup et al. 2022). This complexity, which can be biogenic (e.g., foundation species; Bracken et al. 2007, Ellison 2019) or abiotic (e.g., substrate rugosity; Kovalenko et al. 2012), shapes processes ranging from individual foraging behavior (Michel and Adams 2009) to species’ interactions to food-web structure and biogeochemical rates (Langellotto and Denno 2006, Larsen et al. 2021, Soukup et al. 2022). For example, in the presence of simulated aquatic vegetation, predatory diving beetle larvae shift from active foraging to sit-and-wait predation, reducing activity and prey encounter rates (Michel and Adams 2009).

In this contribution, Edeline et al. (2023) present a detailed perspective on the role of habitat complexity in shaping populations of a freshwater fish (medaka, Oryzias latipes, Adrianichthyidae), including survival, age-class diversity, population growth rate, and density-dependence in the stock-recruitment relationship associated with changes in carrying capacity. Importantly, changes in these population demographic attributes and rates were associated with the role of habitat complexity in mitigating cannibalism – consumption of juvenile O. latipes by conspecific adults. Whereas this is not unexpected – Langelotto and Denno (2006) showed that habitat complexity reduces cannibalism in wolf spiders – the careful work of Edeline et al. (2023) to link changes in habitat complexity to multiple population-level attributes provides a uniquely detailed description of the role of submerged aquatic vegetation in mediating population diversity (e.g., higher age-class diversity) and productivity (e.g., population growth rate).

In many ways, this work by Edeline et al. (2023) provides population-level parallels to perspectives on the role of habitat complexity in determining community-level diversity and productivity. Structurally complex habitats, such as those provided by foundation species (Bracken et al. 2007, Ellison 2019) and substrate heterogeneity (Fairchild et al. 2024), are associated with higher species diversity and abundance at the community level. Edeline et al. (2023) extend these perspectives to the population level, highlighting the importance of habitat complexity across levels of biological organization. Their work highlights within-population diversity and interactions, including cannibalism and competition, illustrating often-neglected aspects of food-web complexity (Polis and Strong 1996).

References

Matthew E. S. Bracken, Barry E. Bracken, Laura Rogers-Bennett (2007) Species diversity and foundation species: potential indicators of fisheries yields and marine ecosystem functioning. California Cooperative Oceanic Fisheries Investigations Reports 48: 82-91. https://calcofi.org/downloads/publications/calcofireports/v48/Vol_48_Bracken.pdf

Eric Edeline, Yoann Bennevault, David Rozen-Rechels (2023) Habitat structural complexity increases age-class coexistence and population growth rate through relaxed cannibalism in a freshwater fish. bioRxiv, ver.4 peer-reviewed and recommended by PCI Ecology https://www.biorxiv.org/content/10.1101/2023.07.18.549540v4

Aaron M. Ellison (2019) Foundation species, non-trophic interactions, and the value of being common. iScience 13: 254-68. https://doi.org/10.1016/j.isci.2019.02.020

Tom P. Fairchild, Bettina Walter, Joshua J. Mutter, John N. Griffin. (2024) Topographic heterogeneity triggers complementary cascades that enhance ecosystem multifunctionality. Ecology 105: e4434. https://doi.org/10.1002/ecy.4434

Katya E. Kovalenko, Sidinei M. Thomaz, Danielle M. Warfe (2012) Habitat complexity: approaches and future directions. Hydrobiologia 685: 1-17. https://doi.org/10.1007/s10750-011-0974-z

Gail A. Langellotto, Robert F. Denno. (2006) Refuge from cannibalism in complex-structured habitats: implications for the accumulation of invertebrate predators. Ecological Entomology 31: 575-81. https://doi.org/10.1111/j.1365-2311.2006.00816.x

Annegret Larsen, Joshua R. Larsen, Stuart N. Lane (2021) Dam builders and their works: beaver influences on the structure and Function of river corridor hydrology, geomorphology, biogeochemistry and ecosystems. Earth-Science Reviews 218: 103623. https://doi.org/10.1016/j.earscirev.2021.103623

Matt J. Michel, Melinda M. Adams. (2009) Differential effects of structural complexity on predator foraging behavior. Behavioral Ecology: 313-17. https://doi.org/10.1093/beheco/arp005

Gary A. Polis, Donald R. Strong (1996) Food web complexity and community dynamics. American Naturalist 147: 813-46. https://doi.org/10.1086/285880

Caroly A. Shumway, Hans A. Hofmann, Adam P. Dobberfuhl (2007) Quantifying habitat complexity in aquatic ecosystems. Freshwater Biology 52: 1065-76. https://doi.org/10.1111/j.1365-2427.2007.01754.x.

Pavel R. Soukup, Joacim Näslund, Johan Höjesjö, David S. Boukal (2022) From individuals to communities: habitat complexity affects all levels of organization in aquatic environments. Wiley Interdisciplinary Reviews: Water 9: e1575.  https://doi.org/10.1002/wat2.1575

The slippersnail (Crepidula fornicata), originally from the eastern coast of North America, has invaded European coastlines from Norway to the Mediterranean Sea [1]. This species is capable of achieving incredibly high densities (up to several thousand individuals per square meter) and likely has major impacts on a variety of community- and ecosystem-level processes, including alteration of carbon and nitrogen fluxes and competition with native suspension feeders [2].

Given this potential for competition, it is important to understand the diet of C. fornicata and its potential overlap with native species. However, previous research on the diet of C. fornicata and related species suggests that the types of food consumed may change with age [3, 4]. This species has an unusual reproductive strategy. It is a sequential hermaphrodite, which begins life as a somewhat mobile male but eventually slows down to become sessile. Sessile individuals form stacks of up to 10 or more individuals, with larger individuals on the bottom of the stack, and decreasingly smaller individuals piled on top. Snails at the bottom of the stack are female, whereas snails at the top of the stack are male; when the females die, the largest males become female [5]. Thus, understanding these potential ontogenetic dietary shifts has implications for both intraspecific (juvenile vs. male vs. female) and interspecific competition associated with an abundant, invasive species.

To this end, Androuin and colleagues evaluated the stable-isotope (d13C and d15N) and fatty-acid profiles of food sources and different life-history stages of C. fornicata [6]. Based on previous work highlighting the potential for life-history changes in the diet of this species [3,4], they hypothesized that C. fornicata would shift its diet as it aged and predicted that this shift would be reflected in changes in its stable-isotope and fatty-acid profiles. The authors found that potential food sources (biofilm, suspended particulate organic matter, and superficial sedimentary organic matter) differed substantially in both stable-isotope and fatty-acid signatures. However, whereas fatty-acid profiles changed substantially with age, there was no shift in the stable-isotope signatures. Because stable-isotope differences between food sources were not reflected in differences between life-history stages, the authors conservatively concluded that there was insufficient evidence for a diet shift with age. The ontogenetic shifts in fatty-acid profiles were intriguing, but the authors suggested that these reflected age-related physiological changes rather than changes in diet.

The authors’ work highlights the need to consider potential changes in the roles of invasive species with age, especially when evaluating interactions with native species. In this case, C. fornicata consumed a variety of food sources, including both benthic and particulate organic matter, regardless of age. The carbon stable-isotope signature of C. fornicata overlaps with those of several native suspension- and deposit-feeding species in the region [7], suggesting the possibility of resource competition, especially given the high abundances of this invader. This contribution demonstrates the potential difficulty of characterizing the impacts of an abundant invasive species with a complex life-history strategy. Like many invasive species, C. fornicata appears to be a dietary generalist, which likely contributes to its success in establishing and thriving in a variety of locations [8].

References

[1] Blanchard M (1997) Spread of the slipper limpet Crepidula fornicata (L. 1758) in Europe. Current state dans consequences. Scientia Marina, 61, 109–118. Open Access version : https://archimer.ifremer.fr/doc/00423/53398/54271.pdf

[2] Martin S, Thouzeau G, Chauvaud L, Jean F, Guérin L, Clavier J (2006) Respiration, calcification, and excretion of the invasive slipper limpet, Crepidula fornicata L.: Implications for carbon, carbonate, and nitrogen fluxes in affected areas. Limnology and Oceanography, 51, 1996–2007. https://doi.org/10.4319/lo.2006.51.5.1996

[3] Navarro JM, Chaparro OR (2002) Grazing–filtration as feeding mechanisms in motile specimens of Crepidula fecunda (Gastropoda: Calyptraeidae). Journal of Experimental Marine Biology and Ecology, 270, 111–122. https://doi.org/10.1016/S0022-0981(02)00013-8

[4] Yee AK, Padilla DK (2015) Allometric Scaling of the Radula in the Atlantic Slippersnail Crepidula fornicata. Journal of Shellfish Research, 34, 903–907. https://doi.org/10.2983/035.034.0320

[5] Collin R (1995) Sex, Size, and Position: A Test of Models Predicting Size at Sex Change in the Protandrous Gastropod Crepidula fornicata. The American Naturalist, 146, 815–831. https://doi.org/10.1086/285826

[6] Androuin T, Dubois SF, Hubas C, Lefebvre G, Grand FL, Schaal G, Carlier A (2021) Trophic niche of the invasive gregarious species Crepidula fornicata, in relation to ontogenic changes. bioRxiv, 2020.07.30.229021, ver. 4 peer-reviewed and recommended by Peer Community in Ecology. https://doi.org/10.1101/2020.07.30.229021

[7] Dauby P, Khomsi A, Bouquegneau J-M (1998) Trophic Relationships within Intertidal Communities of the Brittany Coasts: A Stable Carbon Isotope Analysis. Journal of Coastal Research, 14, 1202–1212. Retrieved May 4, 2021, from http://www.jstor.org/stable/4298880

[8] Machovsky-Capuska GE, Senior AM, Simpson SJ, Raubenheimer D (2016) The Multidimensional Nutritional Niche. Trends in Ecology & Evolution, 31, 355–365. https://doi.org/10.1016/j.tree.2016.02.009

Understanding processes at species’ range limits is of paramount importance in an era of global change. For example, the boreal kelp Laminaria digitata, which dominates low intertidal and shallow subtidal rocky reefs in northwestern Europe, is declining in the equatorward portion of its range [1]. In this contribution, Migné and colleagues [2] focus on L. digitata near its southern range limit on the coast of France and use a variety of techniques to paint a complete picture of the physiological responses of the kelp to environmental changes. Importantly, and in contrast to earlier work on the species which focused on subtidal individuals (e.g. [3]), Migné et al. [2] describe responses not only in the most physiologically stressful portion of the species’ range but also in the most stressful portion of its local environment: the upper portion of its zone on the shoreline, where it is periodically exposed to aerial conditions and associated thermal and desiccation stresses.
The authors show that whereas L. digitata possesses mechanisms to protect it from irradiance stress at low tide, these mechanisms are not sufficient to prevent damage to photosynthetic pathways (e.g., reduction in optimal quantum yields of photosystem II). This species experiences severe heat stress associated with mid-day low tides during the summer, and the cumulative damage associated with these stresses is likely associated with the range contraction that is currently underway. Given the important role that L. digitata plays as food and habitat for other organisms, its loss will have cascading impacts on community structure and ecosystem functioning. Understanding the mechanisms underlying these declines is essential to understanding the impacts of climate change on species, communities, and ecosystems.

References

[1] Raybaud, V., Beaugrand, G., Goberville, E., Delebecq, G., Destombe, C., Valero, M., Davoult, D., Morin, P. & Gevaert, F. (2013). Decline in kelp in west Europe and climate. PloS one, 8(6), e66044. doi: 10.1371/journal.pone.0066044
[2] Delebecq, G., Davoult, D., Menu, D., Janquin, M. A., Migné, A., Dauvin, J. C., & Gevaert, F. (2011). In situ photosynthetic performance of Laminaria digitata (Phaeophyceae) during spring tides in Northern Brittany. CBM-Cahiers de Biologie Marine, 52(4), 405. doi: 10.21411/CBM.A.C9EE91F
[3] Migné, A., Delebecq, G., Davoult, D., Spilmont, N., Menu, D., Janquin, M.-A., and Gevaert, F. (2019). Photosynthesis of Laminaria digitata during the immersion and emersion periods of spring tidal cycles during hot, sunny weather. Hal, 01827565, ver. 4 peer-reviewed and recommended by PCI Ecology. hal-01827565