Biodiversity patterns of epipelagic copepods in the South Pacific Ocean: Strengths and limitations of current data bases

Tmean) in the SPO are of significance in a lesser or greater degree at different zones of the study area; and, they have also been documented to be of importance on a macroscale basis for cope- pod abundance and body size [25]. Moreover, the GDM-based spatial analysis conducted for evaluating species composition showed that the stronger predictors of the observed dissimilar- ities were the same variables (S, O2, Tmean, Chla; S3 Fig). It has been found that chlorophyll strongly correlates with copepod abundance in the ocean [34, 86, 99, 105]. Oxygen is also a significant factor influencing copepod distribution in the ocean, because its availability is a major driver for temperature-size responses in aquatic organisms [106], and the variation in its concentration, for example, due to deoxygenation, can affect copepod physiology, with drastic impacts in both coastal [11, 107] and oceanic [108] species. Warmer water increases copepod growth, development and molting rates [107], with an associated increase in the oxygen demand. This can lead to a reduction in population growth by limiting the hatching success and naupliar growth in early life stages [11], as well as reducing egg production and somatic growth in adults [107], together with shifts in their depth distribution [108]. This involves physiological adaptations that may vary according to PLOS ONE Biodiversity of copepods in the South Pacific PLOS ONE | https://doi.org/10.1371/journal.pone.0306440 July 11, 2024 17 / 28

the species, their body weight and extent of motion [109]. Altogether, these combined drivers have a potential effect on copepod ecology and, consequently, on their biogeographic patterns. Salinity and its variation also affects copepod distribution [110], biomass [111], reproduc- tion [112, 113], growth [114, 115], body size [25] or fatty acid synthesis [115], with various ranges of tolerance that may or may not be favorable, according to their biology and location. This may be relevant since global warming in some areas of the ocean can lead to an increase in salinity (evaporation or greater circulation) or a reduction of the same (greater run-off of fresh water into the ocean, rainfall, or ice melt), producing osmotic stress that may be costly for their physiology. Temperature itself plays a fundamental role in the marine ecosystem, since it controls phys- iological rates [116], metabolism [116], body size [25], reproduction [10], mortality [117] and community structure [24, 118], even at large spatial scales [25, 35, 41, 119]. Thus, it may be thought that temperature also influences the distribution of organisms in the ocean, as it can substantially vary in its three dimensions. Over the meridional axis, temperature exhibits a strong gradient from the equator to the poles, with a similar gradient commonly present from coastal areas to central gyres, whereas in the vertical axis, it has a strong temperature gradient from the upper mixed layer down to the deep, cold waters. These gradients in the water col-

umn structure may contribute to the generation of large-scale spatial trends in the distribution of organisms such as copepods [120]. Although this study did not separate copepods into func- tional groups according certain traits more specifically linked to environmental variations, the fact that composition changed mainly due to oceanographic factors that influence the nesting of functional groups, highlights and support those findings [121], as beta diversity is a proxy of difference in number of species among ecosystems, thus reflecting changes or gradients in the environment as well. Species distribution and the structure of species assemblages may additionally obey some physical processes (e.g., large-scale circulation, currents, eddies, fronts, interaction with the atmosphere) and the effect of local conditions. In this sense, it is possible that in addition to more direct effects on copepod populations, temperature is reflecting variability in water mas- ses distribution controlled by large-scale circulation. The distribution of drifting plankton, including copepods, is largely affected by near-surface currents, allowing species radiation and colonization processes [122], and so influencing diversity patterns. Regarding the structure of the copepod community, we found that some dominant species were representing different current systems, and thus distinct plankton communities. How- ever, we could only assess most occurrences of species (not abundance) for each main surface current system of the SPO (S1 Table). The occurrences distribution of dominant species

seemed to be concordant with patterns documented for copepod diversity on a large scale, with cyclopoids showing greater diversity in the subtropics and calanoids in the temperate zone [25, 123]. Dominant species in the each of the major currents are mostly represented by small Calanoid copepods and the widely distributed Cyclopoid Oithona similis. Some of the recurrent species are Paracalanus parvus and P. indicus. Both species are possibly a single spe- cies traditionally known as P. parvus, but morphologically close to P. indicus, although most likely being a new, non-described species [124]. Lately, it has been referred as P. cf. indicus in the Eastern SPO [125]. The other recurrent species are Calocalanus spp. They are also known as small-sized copepods and mostly found in oceanic waters [126]. The high occurrence of small-sized oceanic copepods, and low presence of large-sized Calanoids, such as Calanus spp., Calanoides spp., and Pleurommama spp., which are abundant in coastal areas of the SPO [38, 93] may indicate that small copepods prevail, or are numerically more abundant than large- sized ones. Also, most data sets do not include harpacticoids or other cyclopoid copepods, such as Oncaea spp. which may be highly abundant in oligotrophic waters of the SPO [127]. PLOS ONE Biodiversity of copepods in the South Pacific PLOS ONE | https://doi.org/10.1371/journal.pone.0306440 July 11, 2024 18 / 28

The OBIS data base does not include the observed number of species in each of the surface cur- rent systems (see S1 Table) when compared to total species inventories. For example, the maxi- mum species richness available in OBIS for the East Australian Current (EAC) and the Humboldt Current System (HCS) is 148 and 91, respectively (see Table 1); that account for ca. a quarter of the species inventory compiled by the Banyuls Observatory data base on copepods for those areas [128]. In the other current systems, similar levels of differences between OBIS data and species inventories can be applied. Current data limitations over space and time Sampling biases must be considered when validating the robustness of the derived spatial pat- terns and the role of environmental drivers. For example, it is crucial to consider that due to spatial gaps of data (Fig 1, Table 1), our results may be strongly constrained be sampling cover- age [129, 130]. The spatial sampling effort was evaluated over the SPO through rarefaction/extrapolation curves by grouping the cells into the main surface current systems of the SPO (Fig 5A). It was found that species richness counts did not reach a clear asymptote at any of the defined current systems. Thus, the SPO is under-sampled for copepods. This aspect may have had an influence on Shannon-Wiener index values (Fig 3), as moderate to high values (from 1.7 on) were esti- mated only for a small area of the SPO. Subsequently, large areas exhibiting much lower values

of richness and biodiversity may result from lack of sampling, description, and digitization of copepod data rather than having a relatively low diversity of species on a significant part of its extension. Moreover, the GDM-based spatial analysis done for evaluating species composition was performed with ecological data based on occurrence instead of abundance, which requires the assumption that any location where the species has not been observed can be treated as absence; therefore, making under-sampled locations more problematic [72]. Although inverte- brates are the most common group documented after plants [131], and that an increasing number of data papers over time reflects progress in digitization and online platforms develop- ment for reporting observations that serve for biodiversity research [131, 132], it has been esti- mated that less than 10% of specimens’ registers are digitized [133], with even less data available online [134]. The lack of description and cataloguing of species and the incomplete knowledge regarding their geographic distribution (i.e., Linnean and Wallacean shortfalls, respectively) are among the most common setbacks for biodiversity research using databases [130], representing a practical limit for biodiversity knowledge that may lead to misidentifica- tion of ecological processes [130]. We further addressed this issue by estimating redundancy index, that indicated a middling sampling over the SPO (S6 Fig) with a lower coverage on its eastern side, that highlights the gaps on great extensions and depicts higher values for areas

where a greater sampling effort was seen. Moreover, through Moran’s I index, it is possible to infer that the distribution of richness is more spatially clustered than expected if underly- ing spatial processes were random, as biodiversity aspects of species and communities are linked to complex interactions between physical, chemical, and biological factors. Consider- ing this, and the developments in statistical modeling making possible to account for biases within heterogeneous data [135], we chose to use the RAC approximation [67] for the GAM analyses. This allows the incorporation of another variable into the models to take spatial autocorrelation into account and maintain strong predictive and inferential performance [67]. However, despite the usefulness of these methodologies, more effort is needed for improving the number and quality of distribution records used as primary data in macroeco- logical studies [136], with databasing focused on georeferencing the information obtained from data collections [129]. PLOS ONE Biodiversity of copepods in the South Pacific PLOS ONE | https://doi.org/10.1371/journal.pone.0306440 July 11, 2024 19 / 28

Copepod resilience to climate forcing Copepod diversity may have remained stable over a long time and large space due to a strong resilience to environmental variability. In this aspect and when dissecting species composition into turnover and nestedness, we observed that the former was higher in areas where hot spots were also identified (Figs 6 and 4, respectively), coinciding with the flow of cold currents (Humboldt current, Zeehan Current in southern Australia, and southern part of the East Aus- tralian current off New Zealand into the West Wind Drift). Therefore, basin-scale ocean cur- rents seem to be an important oceanographic driver explaining the distribution of copepods in the SPO. The potential of large-scale ocean currents generating distributional patterns at vari- ous levels in the ocean has recently been studied more across different taxa [137–141], showing results that support the hypothesis and suggest that it may be a general process transcending taxa and spatial scales [141]. All these large-scale circulation patterns may allow the recoloniza- tion and mixing of copepod assemblies, thereby acting as a buffer to cope with the local impact of warming, deoxygenation, or other altered environmental conditions driven by natural per- turbations or anthropogenic origin. This buffering effect may allow planktonic organisms such as copepods to exhibit a high resilience to large-scale changes in hydrographic conditions due to global warming. Conclusion The diversity distribution of copepods in the SPO follows general patterns observed in other

organisms in the ocean, although patterns over a basin scale with presence of hot and cold spots do not replicate observed patterns reported for the North Pacific or Atlantic Ocean. The key environmental correlates for such biodiversity patterns were found to be mean tempera- ture, chlorophyll-a concentration, salinity, oxygen concentration, and residuals autocovariate. A significant role for causing and maintaining copepod diversity patterns was also attributed to large-scale circulation processes, which may act as a buffer for changes in local conditions and allow species recolonization. Therefore, we suggest that planktonic copepods may exhibit a strong resilience to climate change impact. However, caution should be taken in interpreting our findings since our modeling approaches and spatial analyses were strongly constrained by spatial and temporal gaps in sampling efforts and data availability. Future analyses are expected to improve observed pat- terns with the completion of global biodiversity data bases. Supporting information S1 Fig. Kriging interpolation analysis done for the western and eastern side of the SPO with 394 cells accounting for number of genera. Transparent squares are the 1˚ sampled cells used for Kriging interpolation, whereas the grey dotted line delimits the South Pacific Ocean. Map projection is WGS 84/PDC Mercator (EPSG 3832). (JPG) S2 Fig. Maximum height of the spline function (hence the maximum value of the trans- formed predictors), indicating the stronger predictors of the observed dissimilarities for

S3 Fig. Spearman correlation matrices of environmental variables used in GAM models for (a) alfa diversity and (b) beta diversity. Positive correlations are displayed in red and negative correlations in blue color. Color intensity is proportional to the correlation coefficients. In the right side of the correlogram, the legend color shows the correlation coefficients and the corre- sponding colors. Their acronyms are: Tmean = mean temperature, Tstab = temperature stabil- ity, Tsd = standard deviation of temperature, S = salinity, Chla = chlorophyll-a concentration, O2 = dissolved oxygen concentration, MLD = mixed layer depth. (JPG) S4 Fig. Correlation between observed and predicted species richness. Pearson’s r coefficient: 0.77 (p-value<0.01). Negative residuals (below the reference line) indicate knowledge short- falls, whereas positive residuals (above the reference line) indicate underestimated species rich- ness. (JPG) S5 Fig. Correlation between observed and predicted species composition. Pearson’s r coeffi- cient: 0.84 (p-value<0.01). Negative residuals (below the reference line) indicate knowledge shortfalls, whereas positive residuals (above the reference line) indicate underestimated species composition. (JPG) S6 Fig. Redundancy index obtained for the western and eastern side of the SPO. Values close to 1 indicate good sampling, whereas values close to 0 indicate poor sampling. Transpar- ent squares are the 1˚ sampled cells used for Kriging interpolation, whereas the grey dotted

line delimits the South Pacific Ocean. Map projection is WGS 84/PDC Mercator (EPSG 3832). (JPG) S1 Table. Dominant copepod species in terms of occurrence for the main surface current systems of the South Pacific Ocean. CHC = Cape Horn Current; EAC = East Australian Cur- rent; HCS = Humboldt Current System; SEC = South Equatorial Current; SPSG = South Pacific Subtropical Gyre; WWD = West Wind Drift. (DOCX) S2 Table. Dominant copepod species of the South Pacific Ocean in terms of occurrence for all ranges values of beta diversity and its components turnover and nestedness. (DOCX) Acknowledgments We are grateful to all the scientific community providing data on marine biodiversity to OBIS portal, and special thanks to ESPOBIS, the Eastern South Pacific Node of OBIS. Two anony- mous reviewers greatly helped to improve our work. Author Contributions Conceptualization: Manuela Pe´rez-Arago´n, Ruben Escribano, Reinaldo Rivera. Data curation: Reinaldo Rivera, Pamela Hidalgo. Formal analysis: Manuela Pe´rez-Arago´n. Funding acquisition: Ruben Escribano. Investigation: Manuela Pe´rez-Arago´n, Ruben Escribano, Reinaldo Rivera, Pamela Hidalgo. PLOS ONE Biodiversity of copepods in the South Pacific PLOS ONE | https://doi.org/10.1371/journal.pone.0306440 July 11, 2024 21 / 28

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