The above methods provide assembly and installation instructions for two designs of Coral Arks systems. Prototypes for each design were assembled and field-tested in San Diego, USA, prior to long-term deployment to evaluate the drag characteristics and optimize the structural integrity based on modeled and empirical values of strength. The modeling efforts instrumental to the selection and refinement of both the Arks geometries presented here, including the results from wind tunnel testing, hydrodynamic simulations, and the in-water validation of the modeled values using prototype structures, are described in detail in Section 6 of Supplemental File 1. The results from the modeling and in-water testing of the "Shell" Arks design are shown here. Two structures of each design were then deployed at Caribbean field sites in Puerto Rico and Curaçao (four total Arks structures installed), and corals were translocated to the structures. Water quality, microbial community, and coral survival metrics associated with the "Shell" Arks design and two seafloor control sites were collected at several time points spanning 6 months to characterize and determine the changes in the environmental parameters and coral health associated with the Arks structures following natural recruitment and the addition of seeded ARMS.
Drag characteristics of Coral Arks
It is important to understand the drag characteristics of Coral Arks in order to design a structure and mooring that will survive the target environment. From a structural perspective, the hydrodynamic drag, in combination with the net buoyancy, imposes loadings within the structure, particularly on the mooring and its anchoring system. We conducted modeling and experimental measurements to estimate the drag characteristics of the Arks structures. The results of these tests for the "Shell" design of Arks structures are detailed below. Modeling was carried out by estimating the drag of the individual elements of the structure, summing these, and then combining the result into an effective drag coefficient as shown in equation (1) and equation (2):
(1)
(2)
where Dtotal is the total drag of the structure estimated from the sum of the Di element drags, CD is the overall structure drag coefficient, is the fluid density, U is the flow speed of the object relative to the fluid, and A is the frontal area of the structure. In these calculations, the elements were all assumed to be cylinders, with their orientation to the flow dictated by the upright geometry of the Ark structure. The modeling was performed for the same prototype "Shell" system (a 2V geodesic sphere) that was used for tow testing (described below) prior to the construction of the final field systems. The prototype had a total frontal area of approximately 2.10 m2, and the modeling results indicated an effective drag coefficient for the entire structure of approximately 0.12. The model-predicted drag of the structure as a function of velocity is shown in Figure 4.
Experimental estimates of the drag force of the structure that would be experienced under different flow velocities were obtained by towing the Ark structure behind a vessel with a load cell spliced in-line with the towing line and a tilt sensor to record the changes in the Ark's orientation relative to the vertical axis at a range of tow speeds. Prior to towing, the in-water weight of the structure was determined, and sufficient additional weight was added to the structure to simulate a net buoyancy of approximately 200 kg (an initial target for the system). Based on the tension in the tow cable and the inclination angle of the Ark, the drag (Dtow) at each speed was determined using equation (3):
(3)
where T is the measured tension from the load cell, and is the tilt angle relative to the vertical axis. The resulting drag versus speed relationship is shown in Figure 4. A best fit drag curve (of the form Dtow α U2; see Figure 4), combined with estimates of the frontal area and the water density, was then used to determine the empirical drag coefficient of 0.13.
The Reynolds number during the tow testing (and the range used for the modeling) was in the range of 105-106, generally in the turbulent flow regimes. Typical values of the drag coefficient for a sphere in this Reynolds number range are between 0.2 and 0.4. For comparison purposes, a plot of the drag curve for a sphere with a drag coefficient of 0.3 is shown in Figure 4. Thus, the modeled and experimental estimates of the drag coefficient are in the order of two to three times smaller than for a sphere, which is consistent with the more open character of the structure.
To validate these modeled results, we also conducted field measurements of the response of two "Shell" Arks structures to flow. To achieve this, the same load cell was installed temporarily in line with the Ark main mooring line, a tilt sensor was installed on the Ark, and a current meter was installed at the site to simultaneously monitor the water speed. The buoyancy and drag components of the tension were then calculated from the tilt angle and the load cell measurements (Figure 5). The current speeds during the measurement period were relatively stable at about 20 cm/s, and the data set was relatively short; hence, the data were averaged over the period and used to compare the field drag and velocity response to the modeled and experimental towing estimates. These results show that under expected conditions at the deployment site (flow speeds up to 1.3 m/s during a typical storm event), the drag force on the system is expected to be less than 300 kg.
Both "Shell" structures in Vieques, Puerto Rico, survived a direct hit from the Category 1 Hurricane Fiona in September 2022 with no apparent damage to the structures, mooring, or anchoring system, providing an in situ test that supports the design. A nearby buoy (CARICOOS) recorded current speeds of 1.05 m/s at a 10 m depth at the deployment site, corresponding to a drag force of approximately 160 kg on the mooring systems. The systems were designed to withstand 1,600 kg of force (considering the anchor capacity and component breaking strength) and, therefore, are not expected to fail under ambient or typical storm conditions.
Net buoyancy monitoring for Coral Arks
The same approach described for validating the drag characteristics of the Ark structures was also used to develop a method for monitoring the net buoyancy of the Arks. As long as the physical structure of the Ark remains constant, the net buoyancy provides a rough proxy for monitoring the overall community calcification and, thus, the coral growth, as well as a maintenance metric to determine if the system has sufficient positive buoyancy to compensate for biological growth over time. The buoyancy component (B) of the mooring tension was calculated using the strain gauge and tilt sensor data in equation (4):
(4)
where T is the measured tension from the load cell, and is the tilt angle. The resulting time series of the net buoyancy is shown in Figure 5. Under the relatively stable current conditions present during the field monitoring events, we found the two "Shell" Arks structures deployed in Vieques, Puerto Rico, to have similar net buoyancies of 82.7 kg ± 1.0 kg (Ark 1) and 83.0 kg ± 0.9 kg (Ark 2) when averaged over the monitoring period (± one standard deviation) after all the corals and seeded ARMS units were translocated to the structures 6 months after the initial structure deployment. The results show that short-term monitoring during relatively stable periods of water flow can be used to determine the net buoyancy in the field to within ~1 kg, which should prove useful over the long term for monitoring changes in biomass.
Water quality and microbial community dynamics
Metrics associated with water quality and water column-associated microbial communities were measured on two midwater "Shell" Arks, which were anchored in 55 ft of water with the top of the Arks at a 25 ft depth, offshore of Isla Vieques, Puerto Rico (Figure 6C). The water quality metrics, microbial and viral abundances, and average microbe size from two Arks were compared to the same metrics from two nearby seafloor "control" sites, which were also at a 25 ft depth but much closer to shore (Figure 6D). The measurements shown were collected immediately after the installation of the Arks with an initial batch of translocated corals (November 2021) and 6 months later after a second batch of corals and seeded ARMS were translocated to the Arks (May 2022); they were then averaged across both sites (Arks and control sites) for comparison. As the seeded ARMS were transferred to the Arks at 6 months post-deployment, the accumulation of biological communities on the structures during the first 6 month period was associated with biofouling and natural recruitment.
The Arks environment exhibited higher average daytime light intensities (Figure 6A), higher average flow speeds (Figure 6C), lower dissolved organic carbon concentrations (Figure 6F), and lower diel fluctuations in dissolved oxygen concentrations (Figure 6G) than the benthic control sites. The Arks also displayed microbial communities with higher virus-to-microbe ratios than the control sites (Figure 7A), driven by a higher abundance of free viruses (Figure 7C) and a lower abundance of microbes (Figure 7B) in the midwater Arks environment. The microbial communities on the Arks were composed of, on average, physically smaller cells than the microbial communities at the seafloor sites (Figure 7D). Differences in temperature between the Arks and the control sites were not significant (Figure 6E). All of the above trends are consistent with better water quality and healthier microbial communities on the Arks than at the control sites. These conditions persisted through the initial 6 months of the deployment, during which a nascent biological community developed on the Arks through both the translocation of coral nubbins and natural recruitment from the water column and experienced successional changes, as well as through the addition of seeded ARMS onto the structures at month 6.
Coral survival
A cohort of corals comprising eight species and various morphologies were distributed to the Arks and benthic control sites both following the installation of the Arks (month 0) and following the addition of the seeded ARMS at month 6. The original parent colonies of each species of coral were fragmented into nubbins (2-8 cm in a given dimension) and attached to limestone coral plates (four to five nubbins per 20 cm2 plate) that were distributed equally at both the Arks and control sites, ensuring that the same species and genotypes were represented at both the midwater Arks sites and control sites. The survival of these translocated corals was assessed every 3 months at the Arks and control sites. Nine months after the translocation of the first cohort of corals, more corals were still alive on the Arks (80%, Figure 8) compared to the control sites (42%, Figure 8).

Figure 1: Diagram showing the structural components of two fully installed Coral Ark structures. Left, "Shell" and "Two-Platform" (right) Coral Arks structures are shown, together with two methods for providing positive buoyancy and two methods for anchoring. Abbreviation: ARMS = Autonomous Reef Monitoring Structures. Please click here to view a larger version of this figure.

Figure 2: Design, deployment, and transfer of ARMS units. (A-D) PVC ARMS and (E-H) Limestone ARMS from seafloor seeding sites to Coral Arks. (A) Photo credit to Michael Berumen. (B) Photo credit to David Littschwager. Abbreviations: PVC = polyvinyl chloride; ARMS = Autonomous Reef Monitoring Structures. Please click here to view a larger version of this figure.

Figure 3: Images representing the deployment stages of Coral Arks, including transport to the site and full installation. (A-C) Shell type and (D-F) Two-Platform type systems. Please click here to view a larger version of this figure.

Figure 4: Drag characteristics of the "Shell" Ark structures based on modeling, experimental tow testing, and field validation relative to the drag of a sphere of the same approximate scale. "ARK1" and "ARK2" are identical "Shell" Ark structures installed at the same site in Vieques, Puerto Rico. Please click here to view a larger version of this figure.

Figure 5: Measured net buoyancy values for two "Shell" Arks in Vieques, Puerto Rico. Shown are the water velocity (right axis, medium colors), net buoyancy (left axis, light colors), and calculated drag/tension on the mooring line (left axis, dark colors) for "Shell" Ark 1 (blue) and "Shell" Ark 2 (green). Please click here to view a larger version of this figure.

Figure 6: Water quality metrics associated with the "Shell" Arks and seafloor control sites in Vieques, Puerto Rico, immediately following the installation and 6 months afterward. (A) Daytime light intensity, (B) current speed, (C,D) photos taken 6 months post installation, (E) temperature, (F) dissolved organic carbon, (G) changes in dissolved oxygen levels in the Arks versus control sites over 6 months. Please click here to view a larger version of this figure.

Figure 7: Metrics associated with the water column-associated microbial communities on the "Shell" Arks and seafloor control sites in Vieques, Puerto Rico immediately following installation and 6 months afterward. (A) Virus-to-microbe ratio, (B) bacterial cell abundance, (C) free virus abundance, and (D) average bacterial cell size. Please click here to view a larger version of this figure.

Figure 8: Proportion of surviving corals on the "Shell" Arks and seafloor control sites in Vieques, Puerto Rico during the first 9 months following translocation. The images represent the status of a single coral plate on the Arks (top) and on the benthic control sites (bottom) immediately following translocation (left) and 6 months after translocation (right). Please click here to view a larger version of this figure.
Table 1: ARMS construction and design considerations. Abbreviations: ARMS = Autonomous Reef Monitoring Structures; PVC = polyvinyl chloride. Please click here to download this Table.
Table 2: Coral Arks design considerations. Abbreviations: PVC = polyvinyl chloride; ARMS = Autonomous Reef Monitoring Structures; HDPE = high-density polyethylene. Please click here to download this Table.
Supplemental File. Please click here to download this file.