Abstract: Aim Delimiting recently diverged species is challenging. During speciation, genetic differentiation may be distributed unevenly across the genome, as different genomic regions can be subject to different selective pressures and evolutionary histories. Reliance on limited numbers of genetic markers that may be underpowered can make species delimitation even more challenging, potentially resulting in taxonomic inconsistencies. Rockhopper penguins of the genus Eudyptes comprise three broadly recognized taxa: northern (E. moseleyi), southern (E. chrysocome) and eastern rockhopper (E. filholi). Their taxonomic status has been controversial for decades, with researchers disagreeing about whether E. chrysocome and E. filholi are distinct species or conspecific. Our goal is to evaluate genome-wide patterns of divergence to evaluate genetic differentiation and species delimitation in rockhopper penguins, and to assess which mechanisms may underlie previous discordance among nuclear versus mitochondrial analyses. Location Sub-Antarctic and temperate coastal regions of the Southern Hemisphere. Methods We generated reduced-representation genomic libraries using double digest restriction-site associated DNA (ddRAD) sequencing to evaluate genetic differentiation, contemporary migration rates and admixture among colonies of rockhopper penguins. Results The extent of genetic differentiation among the three taxa was consistently higher than population-level genetic differentiation found within these and other penguin species. There was no evidence of admixture among the three taxa, suggesting the absence of ongoing gene flow among them. Species delimitation analyses based on molecular data, along with other lines of evidence, provide strong support for the taxonomic distinction of three species of rockhopper penguins. Main conclusions Our results provide strong support for the existence of three distinct species of rockhopper penguins. The recognition of this taxonomic diversity is crucial for the management and conservation of this widely distributed species group. This study illustrates that widespread dispersive seabird lineages lacking obvious morphological differences may nevertheless have complex evolutionary histories and comprise cryptic species diversity.

Abstract: Tides are one of the basic types of ocean water motions. Previous studies have reported that the M2 constituent exhibits seasonal variations (annual cycles) in some regions. However, based on the newly proposed method of modified two-step harmonic analysis (HA) and its application at 240 global tide gauges, we find that the M2 constituent as well as the S2 and K1 do not have significant seasonal variations at these tide gauges. The seasonal variations of the M2 constituent reported in previous studies are caused by its satellites, the H1 and H2 constituents, which are not resolved in these studies due to the short time window (one month or three months) used in HA. Because the frequency of the H1 (H2) constituent is equal to that of the M2 minus (plus) the frequency of annual cycles, the superposition of the M2, H1 and H2 constituents with constant amplitudes is equivalent to the M2 constituent with seasonally varying amplitudes. Compared with the new method, some adaptations to traditional HA aiming to capture variations in amplitudes and phase lags of constituents have some limitations, because they either neglect some satellites of the major constitutes or introduce spurious fluctuations resulting from an unreasonably large number of independent points. The nodal modulations of predominant constituents are also explored in this study. On the global scale, the nodal modulations of the M2, K1 and O1 constituents agree with the theoretical predictions, except a cold spot region with reduced nodal modulation in the Gulf of Maine and a hot spot region with enhanced nodal modulation in the South China Sea for the M2. Nodal modulation is also found for the S2 constituent (in theory, the S2 has no nodal modulation), which is 0.8% averaged at 164 tide gauges where the S2 is not too weak.