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The solitary cold-water scleractinian C. huinayensis is described here as a brooder. Although reproduction in scleractinian CWCs is still poorly known, no other temperate species has yet been described to brood larvae. The solitary temperate CWC D. dianthus12, as well as the temperate colonial CWC D. pertusum8,10,38, M. oculata8,10 and O. varicosa9 reproduce by broadcast spawning. Brooding has only been reported in subpolar and polar solitary CWCs from the Southern Ocean17,18.Although quantitative data on the number of larvae released in the four Southern Ocean brooders are lacking, a potential number of larvae released per polyp can be inferred from their maximum fecundity (Table 1). C. huinayensis appears to be in the lower range of larvae production (6.5 ± 11.4 month-1 larvae), when compared its larval size (750–1080 µm length) with Balanophyllia malouinensis larvae ( > 600 µm, Table 1).Table 1 Larval features of scleractinian CWC species.Full size tableThorson’s rule43,44 states that organisms at higher latitudes tend to produce larger and fewer offspring and are frequently brooders. However, the brooding C. huinayensis appears to defy this rule, as it occurs at mid-latitudes (36° and 48.5° S33,45). Though the phylogeography of C. huinayensis is not yet clear, six other solitary species of the genus Caryophyllia are endemic to Antarctica46, suggesting that the genus may have originated in the Southern Ocean, with the mid-latitude distribution of C. huinayensis making the downstream end dispersal via the cold Humboldt Current branching off the Southern Ocean. In our case, Thorson’s rule does not seem to be a good predictor of the macroevolutionary patterns and reproductive mode in Caryophyllia.A better explanation can be inferred from Kerr et al.47. Their phylogenetic study on scleractinians revealed that the change from spawning to brooding (or vice versa) is based on the sexuality of the corals (i.e., gonochoric or hermaphroditic) and not on latitudinal distribution. The main pathway is from gonochoric spawners to gonochoric brooders, then to hermaphroditic brooders, and finally hermaphroditic spawning, which is the dominant reproductive mode in shallow-water corals.The results of our study indicate that C. huinayensis reproduces throughout the year, albeit with large temporal variations in the number of larvae released. However, the fluctuations were not seasonal. This may be due to the fact that the aquarium for this experiment lacked external timing signals (zeitgebers) usually found in the field, i.e., there were no fluctuations e.g., in water temperature, food frequency, food quality, or salinity, which might otherwise have synchronized the corals’ internal clock. Although, it is not yet known if the local C. huinayensis population exhibits seasonality in their larval release, the lack of larvae in April 2021 could also be due to poor internal fertilisation success based on the quality and/or quantity of sperm released (which was never observed).If there is no seasonal release of larvae from the natural coral population, this may indicate that rapid recolonisation is possible throughout the year following a disturbance. Substrate alterations are usually observed in the Patagonian fjord region, where strong physical disturbances such as landslides occur48, due to precipitation and earthquakes49. Also, hypoxia events ( More

In this study, we provide the first description of the bacterial communities of cryoconite holes from South American glaciers, in particular from both small high-elevation glaciers of the Central Andes in the Santiago Metropolitan Region (Chile), and from the tongues of two large glaciers in Patagonian Andes that reach low altitudes. These pieces of information fill a large geographical gap in our knowledge of glacier environments because this is the first description of the microbial communities of supraglacial environments in South America, a continent with about 30,000 km2 covered by ice29. Results showed that the large Patagonian glaciers (Exploradores and Perito Moreno) had the highest oxygen concentrations, while Iver and East Iver had the lowest ones and Morado an intermediate value. This pattern could be related to the different altitudes of the glaciers. Indeed, since water temperature in cryoconite holes is always quite low and stable at all altitudes, oxygen solubility in these environments is related to the atmospheric partial pressure of oxygen that decreases at increasing altitude30. This result is consistent with [O2] values we found in our samples. Indeed, Exploradores and Perito Moreno are located in Patagonia at low altitudes ( 40%), whereas mining is also an additional important black carbon source50. Their similarity can therefore derive also from being exposed to the same general ecological conditions, including high UV radiation, oxidative stress, anthropic pressures, and probably, also from similar sources of bacteria. These results therefore highlight that correlative studies like the present ones can hardly disentangle the effects of geographical positions and ecological conditions on the structure of cryoconite hole bacterial communities, and further studies should be designed to add insight into this still open question.Analyses of alpha diversity indices indicated that cryoconite holes on Exploradores glacier showed the highest richness and evenness. Samples on the Exploradores were collected close to the glacier terminus, surrounded by a rich evergreen broadleaf vegetation, and in an area with abundant supraglacial debris and frequented by tourists. The higher biodiversity of this large, low-altitude glacier, compared to that of the small, high-altitude Iver and East Iver glaciers is not surprising, as the rich evergreen broadleaf forest that surrounds the tongue of the first glacier can be the source of a richer and more diverse bacterial community than the bare ground surrounding the other ones. However, it is more surprising that the alpha biodiversity of the large, low-altitude Perito Moreno was intermediate and similar to that of the Morado glacier. Interestingly, Perito Moreno was the southernmost glacier among those we collected, and was surrounded by a less diverse forest, dominated by southern beeches, Nothofagus ssp. than that of Exploradores, while Morado was the glacier where samples were collected at the lowest altitude among the three glaciers near Santiago. We may therefore speculate that a broad gradient related to altitude and general climate conditions of the area surrounding the glacier may somehow affect its biodiversity. For instance, among the most abundant orders, Cytophagales were more abundant on high than on low-elevation glaciers (Fig. 5b). A similar pattern was observed for the Micrococcales and Chitinophagales (Fig. 5c–k) with the only exception of Iver.In summary, we provide the first-ever description of the bacterial communities of cryoconite holes of glaciers in South America, specifically in the Southern Andes. This study thus fills an important gap of knowledge as almost no information was previously available on the cryoconite holes of this continent, and opens the possibility of future biogeography analyses including samples from almost every important glacial area of the world. The five glaciers we investigated are still a too small sample for thoroughly assessing the ecological processes that control cryoconite hole bacterial communities, and a larger set of environmental variables should also be considered, but we hope this study can be the basis for further investigations aiming at a deeper understanding of these extreme environments. More

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