The “Cribrilina annulata" problem and new species of Juxtacribrilina (Bryozoa: Cheilostomata: Cribrilinidae) from the North Pacific

Abstract

Originally described from Greenland, Juxtacribrilina annulata (Fabricius, 1780) (previously known as Cribrilina annulata) has long been regarded as having a circumpolar, Arctic-boreal distribution. The genus Juxtacribrilina Yang, Seo, Min, Grischenko & Gordon, 2018 accommodated J. annulata and three related North Pacific species formerly in Cribrilina Gray, 1848 that lack avicularia, have a reduced (hood-like, cap-like, or vestigial) ooecium closely associated with modified latero-oral spines to form an ooecial complex, and produce frontally or marginally positioned dwarf ovicellate zooids. While the recently described NW Pacific species J. mutabilis and J. flavomaris, which have a vestigial ooecium like a short, flattened spine, clearly differ from J. annulata, the differences between J. annulata and other Pacific populations remained unclear. Here we provide descriptions for five species from the North Pacific region. We identified a specimen from the Sea of Okhotsk as J. annulata. Among the other four species, J. ezoensis n. sp. has a trans-Pacific distribution (abundant at Akkeshi, Hokkaido, Japan; also detected in the Commander Islands and at Ketchikan, Southeast Alaska); J. pushkini n. sp. was found only at Ketchikan; J. dobrovolskii n. sp. was found only at Shikotan Island in the Lesser Kuril Chain; and J. tumida n. sp. was found only at Kodiak, Gulf of Alaska. These four species all differ from J. annulata in having one or two frontal pore chambers on the proximal gymnocyst of most zooids; in budding frontal dwarf ovicellate zooids from these chambers rather than from basal pore chambers; in producing dwarf zooids more abundantly; and in having ooecia that are somewhat to markedly more reduced (cap-like rather than hood-like) and more closely integrated with the modified latero-oral spines. Furthermore, in the Pacific species, the ooecium in basal zooids arises from the roof of the distal pore chamber of the maternal zooid; ovicellate zooids can thus also bud a distal autozooid and are often arranged in columnar series with other zooids. In J. annulata, the hood-like kenozooidal ooecium budded from the maternal zooid replaces the distal autozooid, and ovicellate zooids are thus usually not embedded in a columnar series.