Morphology and phytogeography of Laminaria appressirhiza and L. inclinatorhiza (Phaeophyceae) from the Sea of Okhotsk

The re-examination of morphological and anatomical characters of Laminaria appressirhiza and L. inclinatorhiza collected from different localities in the Sea of Okhotsk was performed. Despite their commercial and ecological importance to the region they have not been comprehensively reviewed since their first description in 1970. Our results show that some original diagnostic key characters (e.g., shape of holdfast, shape of sporangial sori, and dissection of blade) are not stable and have deviations from the type morphology when plants grow in different environments. In L. inclinatorhiza, the sporangial sori development occurred differently to the pattern indicated in original species description as they did not develop simultaneously on both sides of the blade. Instead, the sporangial sori outlines on both sides of the blade did not coincide at first and only became coincident later. Also, a deep-water population of L. inclinatorhiza with an unusual and interesting morphology, growing at depths of 15-25 m on opened rocky coasts in Taujskaya Bay (northern part of the Sea of Okhotsk) was found. The stable diagnostic key characters to distinguish these two species are the cone-like, multilayered, very thick and massive holdfast (in L. inclinatorhiza) and rolled margins of blades, lamellar rosette-like part of thallus, and sporangial sori developing only on one side of the blade (in L. appressirhiza). The ecological characteristics, distribution, and abundance of both species in the Sea of Okhotsk are discussed. Both species are perennial and widely distributed in the region. L. appressirhiza is more often found as a subdominant species among other kelps, forming maximum biomass and density of 7-9 kg and 8-25 plants per $1m^2$, respectively. L. inclinatorhiza sometimes forms local mono-species communities with maximum biomass and density of 10-12 kg and 10-15 plants per $1m^2$, respectively.