Osinga, R.

Highlights

• Fish abundance and biomass differed per artificial reef type.
• This difference is driven by the availability of small shelters.
• Artificial reef deployment can be more efficient by choosing better performing or cheaper designs.

In this study, we compared the early fish colonization of three types of artificial reefs deployed in the coastal waters of Saba and St Eustatius in the Caribbean: reef balls®, layered cakes and piles of locally obtained basaltic rock. As an indicator of performance, three fish assemblage parameters (abundance, biomass, species richness) were measured using underwater visual censuses at 11 months post-deployment and 4 months after restoration from hurricane damage. All artificial reef plots showed higher values for fish abundance, biomass and species richness than control plots covered by bare sand, which shows that artificial reefs can locally enhance the fish assemblage. However, the effect differed among artificial reef plots. Fish abundance was 3.8 times higher on the layered cake plots compared to the reef ball plots, while fish biomass was 4.6 times higher. Rock pile plots had intermediate values. Species richness did not differ significantly among different artificial reef plots. Three-dimensional modelling revealed that layered cakes had a smaller gross volume, shelter volume and total surface area than reef balls. The availability of multiple small shelters in the layered cake design appeared to be more relevant than other physical parameters, as the layered cake plots had higher fish abundance than the reef balls plots. We concluded that on Saba and St. Eustatius, layered cake plots performed better than reef ball plots after one year of colonization. Rock pile plots, made of local volcanic rock, showed an intermediate performance, and were 4–10 times cheaper to construct. If observed differences are consistent with other locations and persist during further colonization, current efforts to deploy reef balls could better be allocated to deploy artificial reef structures with a higher shelter density. 

Reef ball® , Layered cake, Fish abundance, Shelter availability, Habitat restoration, Artificial reef

Abtsract
This study describes in vivo cell turnover (the balance between cell proliferation and cell loss) in eight marine sponge species from tropical coral reef, mangrove and temperate Mediterranean reef ecosystems. Cell proliferation was determined through the incorporation of 5-bromo-2′-deoxyuridine (BrdU) and measuring the percentage of BrdU-positive cells after 6 h of continuous labeling (10 h for Chondrosia reniformis). Apoptosis was identified using an antibody against active caspase-3. Cell loss through shedding was studied quantitatively by collecting and weighing sponge-expelled detritus and qualitatively by light microscopy of sponge tissue and detritus. All species investigated displayed substantial cell proliferation, predominantly in the choanoderm, but also in the mesohyl. The majority of coral reef species (five) showed between 16.1±15.9% and 19.0±2.0% choanocyte proliferation (mean±SD) after 6 h and the Mediterranean species, C. reniformis, showed 16.6±3.2% after 10 h BrdU-labeling. Monanchora arbuscula showed lower choanocyte proliferation (8.1±3.7%), whereas the mangrove species Mycale microsigmatosa showed relatively higher levels of choanocyte proliferation (70.5±6.6%). Choanocyte proliferation in Haliclona vansoesti was variable (2.8–73.1%). Apoptosis was negligible and not the primary mechanism of cell loss involved in cell turnover. All species investigated produced significant amounts of detritus (2.5–18% detritus bodyweight−1·d−1) and cell shedding was observed in seven out of eight species. The amount of shed cells observed in histological sections may be related to differences in residence time of detritus within canals. Detritus production could not be directly linked to cell shedding due to the degraded nature of expelled cellular debris. We have demonstrated that under steady-state conditions, cell turnover through cell proliferation and cell shedding are common processes to maintain tissue homeostasis in a variety of sponge species from different ecosystems. Cell turnover is hypothesized to be the main underlying mechanism producing sponge-derived detritus, a major trophic resource transferred through sponges in benthic ecosystems, such as coral reefs.

Sponges have a remarkable capacity to rapidly regenerate in response to wound infliction. In addition, sponges rapidly renew their filter systems (choanocytes)

to maintain a healthy population of cells. This study describes the cell kinetics
of choanocytes in the encrusting reef sponge Halisarca caerulea during early regeneration (0–8 h) following experimental wound infliction. Subsequently, we investigated the spatial relationship between regeneration and cell proliferation over a six-day period directly adjacent to the wound, 1 cm, and 3 cm from the wound. Cell proliferation was determined by the incorporation of 5-bromo-20-deoxyuridine (BrdU). We demonstrate that during early regeneration, the growth fraction of the choanocytes (i.e., the percentage of proliferative cells) adjacent to the wound is reduced (7.0 ± 2.5%) compared to steady-state, undamaged tissue (46.6 ± 2.6%), while the length of the cell cycle remained short (5.6 ± 3.4 h). The percentage

of proliferative choanocytes increased over time in all areas and after six days of regeneration choanocyte proliferation rates were comparable to steady-state tissue. Tissue areas farther from the wound had higher rates of choanocyte proliferation than areas closer to the wound, indicating that more resources are demanded from tissue in the immediate vicinity of the wound. There was no diVerence in the number of proliferative mesohyl cells in regenerative sponges compared to steady-state sponges. Our data suggest that the production of collagen-rich wound tissue is a key process in tissue regeneration for H. caerulea, and helps to rapidly occupy the bare substratum exposed by the wound. Regeneration and choanocyte renewal are competing and negatively correlated life-history traits, both essential to the survival of sponges. The eYcient allocation of limited resources to these life-history traits has enabled the ecological success and diversification of sponges. 

Ever since Darwin’s early descriptions of coral reefs, scientists have debated how one of the world’s most productive and diverse ecosystems can thrive in the marine equivalent of a desert. It is an enigma how the flux of dissolved organic matter (DOM), the largest resource produced on reefs, is transferred to higher trophic levels. Here we show that sponges make DOM available to fauna by rapidly expelling filter cells as detritus that is subsequently consumed by reef fauna. This “sponge loop” was confirmed in aquarium and in situ food web experiments, using 13C- and 15N-enriched DOM. The DOM-sponge-fauna pathway explains why biological hot spots such as coral reefs persist in oligotrophic seas—the reef’s paradox—and has implications for reef ecosystem functioning and conservation strategies.