Corresponding author: Masato Hirose (
Academic editor: L. van Ofwegen
The status of
I examined specimens from Sagami Bay and surrounding areas collected by Ludwig Döderlein (1880–1881), Franz Doflein and Karl Haberer (1904–1905), Emperor Showa (1918–1971), and most recently by the National Museum of Nature and Science Tokyo (2001–2005); see
The author collected additional specimens from Sagami Bay by dredge from RV
Map showing the areas in Japan where species of
Specimens were observed by light microscope and scanning electron microscope (SEM). For SEM observation, part of each specimen was removed, soaked in a sodium hypochlorite solution to remove the soft tissue, rinsed in water, air dried, and mounted with double-sided adhesive tape or silver paste on an aluminum SEM stub. At Hokkaido University, mounted specimens were coated with Au in a Hitachi E-1030 sputter-coater and observed with a Hitachi S-3000N SEM at 15 kV accelerating voltage. At the SMF, specimens were coated with Pt-Pd and observed with a CamScan SEM. At ZSM, specimens were coated with Au in a POLARON SEM Coating System and observed with a LEO 1430VP SEM at 15 kV accelerating voltage. Fragments removed from specimens in the various collections, prepared and examined by SEM, and subsequently deposited in NMST are indicated in the text by the designation ‘NSMT Te’ (see Supplementary Table 1).
Measurements were taken from SEM images with ImageJ 1.37v software (Image Processing and Analysis in Java, Wayne Rasband, National Institutes of Health, USA;
Colony erect, rigid, dichotomously or irregularly branching: branches cylindrical, flattened, or plate-like, fan shaped. Zooidal frontal shield uniformly tessellated, with a few areolar pores near margin or offset centrally in secondarily calcified wall. Orifice deeply immersed, without teeth on distal periphery, without lyrula or condyles, slightly concave or straight proximally; oral spines absent. Secondary orifice at colony surface cormidial, formed by contributions of secondary calcification from distal and lateral zooids. Suboral avicularium lies at proximal margin of secondary orifice, directed proximally or laterally, sometimes enlarged and occupying about half of frontal shield; small, conical tooth associated with avicularium projecting into secondary orifice (
Orifices of three
Ovicells of
Excluding nominal
ZL, 0.595−1.334 (0.978±0.160); ZW, 0.214−0.840 (0.450±0.133); n=65. OrL, 0.131−0.208 (0.171±0.021); OrW, 0.123−0.219 (0.187±0.018); n=32. AvL, 0.078−0.198 (0.110±0.017); AvW, 0.055−0.148 (0.086±0.016); n=82. OvL, 0.205−0.391 (0.311±0.049); OvW, 0.286−0.439 (0.365±0.043); n=29. Additional measurements: large suboral avicularium (LAv) length, 0.619−0.766 (0.682±0.076); LAv width, 0.398−0.453 (0.426±0.027), n=3.
Colony erect, rigid, dichotomously branching, widely spreading, antler-like, terminal branches slender. Basal part of colony composed of both autozooids and kenozooids. Branches cylindrical, 1.39–4.76 mm wide (2.77±0.85 mm; n=25), with zooids opening all around, four or five zooids across in half-view (
Sagami Bay, Sagami Sea, Tokyo Bay, and off Kii Peninsula, at depths of 432–950 m. The collecting depth of the specimen (1921.11.7.9.) in NHMUK is 250–330 fathoms, which means 457–603 m; therefore, the depth given in
ZL, 0.767−1.150 (0.948±0.100); ZW, 0.468−1.050 (0.735±0.116); n=32. OrL, 0.152−0.211 (0.177±0.019); OrW, 0.189−0.245 (0.222±0.023); n=14. AvL, 0.106−0.271 (0.171±0.034); AvW, 0.096−0.193 (0.144±0.024); n=28.
Colony erect, rigid, dichotomously branching, widely spreading (
Eastern part of Sagami Bay, and the Sagami Sea southwest of Jogashima and west of Ōshima, at depths of 200–493 m.
Examination of Ortmann’s (1890) type specimens revealed this species belongs not in
Androsova’s type specimen (ZIN-1/3670) in Zoological Institute of the Russian Academy of Sciences (ZIN RAS), colony collected southwestern region of Sakhalin, Moneron Island (Kaibato), Sea of Japan, 36 m depth, (examined by micrographs); large erect colonies and fragments (NSMT Te-724 to Te-734; ZIHU 4130 and 4131), collected SE of Akkeshi Bay (
ZL, 0.558−0.921 (0.751±0.101); ZW, 0.408−0.882 (0.611±0.088); n=25. OrL, 0.135−0.223 (0.189±0.019); OrW, 0.130−0.226 (0.192±0.023); n=27. OvL, 0.124−0.444 (0.247±0.072); OvW, 0.104−0.395 (0.214±0.056); n=44.
Colony erect, rigid, robust, with thick, broad, strap-like branches at least 10 zooid widths across, or foliaceous, fan-shaped lobes; lobes or branches 0.86 to 8.04 cm wide (2.15±1.38 cm, n=25), multifurcate or irregularly lobed on distal margin; zooids open on both sides (
Moneron Island, SW Sakhalin, 36 m depth (
My material matches Androsova’s (1958) description of
To date, species of
Map showing the known distribution of the three
While
1a | Colony robust; branches thick and broad, fan-shaped distally; hypertrophied suboral avicularia often occur on the frontal shield | |
1b | Colony more delicate, branches cylindrical or flattened, slender distally; hypertrophied suboral avicularia rare or absent | 2 |
2a | Branches flattened; zooids rhomboidal; hypertrophied suboral avicularia absent | |
2b | Branches cylindrical; zooids rectangular, hypertrophied suboral avicularia present |
I thank Dr. Shunsuke F. Mawatari (Hokkaido University Museum) for valuable assistance in the study of the historical collections. I thank Dr. Bernhard Ruthensteiner and Mrs. Eva Lodde (Zoologische Staatssammlung München) for valuable advice and assistance in the study of the Doflein and Haberer Collections; Madame Marie-Dominique Wandhammer and Madame Marie Meister (Musée Zoologique Strasbourg) for kind help, assistance in observations, and the loan of the Döderlein Collection; Dr. Hiroshi Namikawa (Showa Memorial Institute of the National Museum of Nature and Science) for advice and assistance, and the loan of the bryozoan collection of Emperor Showa; and Mr. Hisanori Kohtsuka (Misaki Marine Biological Station, The University of Tokyo) for crucial support with collecting in Sagami Bay. Special thanks are due to Dr. Joachim Scholz and Mrs. Brigitte Lotz (Senckenberg Forschungsinstitut) for assistance during my stay in Germany. I thank Miss Mary Spencer Jones (Natural History Museum London) for valuable advice and assistance in the survey of the Harmer’s and Androsova’s material. I thank Mrs. Elly Beglinger (Naturalis, Leiden) for assistance in the survey of Harmer’s material. I thank Dr. Nina Denisenko (Zoological Institute of the Russian Academy of Sciences) for sending the photographs of Androsova’s type specimen. I thank Dr. Dennis Gordon (National Institute of Water & Atmospheric Research), Dr. Andrew Ostrovsky (University of Vienna), and two reviewers Dr. Björn Berning (Oberösterreichische Landesmuseen) and Dr. Kevin Tilbrook (Queensland Museum) for valuable advice and suggestions on the manuscript. I thank Professor Matthew Dick (Hokkaido University) for checking my English and providing crucial suggestions. This study was supported in part by the 21st Century COE Program “Neo-Science of Natural History” at Hokkaido University, funded by the Ministry of Education, Culture, Sports, Science and Technology (MEXT), Japan, and a Fellowship from the Japan Society for the Promotion of Science (No. 20-3856).
Specimens examined in this study. (doi: