European Journal of Taxonomy 203: 1—41

http://dx.doi.org/10.5852/ejt.2016.203

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2016 • Hirose M.

Research article

urn:lsid:zoobank.org:pub:325E4EF8-78F9-49D0-82AF-4C358B24F7F8

Diversity and distribution of adeonid bryozoans

(Cheilostomata: Adeonidae) in Japanese waters

Masato HIROSE

Atmosphere and Ocean Research Institute, The University of Tokyo,

Kashiwanoha 5-1-5, Kashiwa, Chiba 277-8564, Japan.

Email: mhirose64@gmai1 com

urn:lsid:zoobank.org:author:C6C49C49-B4DF-46B9-97D7-79DE2C942214

Abstract. Adeonid bryozoans construct antler-like erect colonies and are common in bryozoan

assemblages along the Japanese Pacific coast. The taxonomy of Japanese adeonid species, however,

has not been studied since their original descriptions more than 100 years ago. In the present study,

adeonid specimens from historical collections and material recently collected along the Japanese coast

are examined. Eight adeonid species in two genera were detected, of which Adeonella jahanai sp. nov.,

Adeonellopsisparvirostrum sp. nov., and Adeonellopsis toyoshioae sp. nov. are described as new species

based on the branch width, size and morphology of frontal or suboral avicularia, shape and size of

areolar pores, and size of the spiramen. Adeonellopsis arculifera (Canu & Bassler, 1929) is a new record

for Japan. Lectotypes for Adeonellopsis japonica (Ortmann, 1890) and Adeonella sparassis (Ortmann,

1890) were selected among Ortmarm’s syntypes. Most species of Adeonellopsis around Japan have a

southern distribution from Sagami Bay to Okinawa, while A .japonica shows a more northern distribution

from Kouchi to Otsuchi. In contrast, Adeonellopsis arculifera was collected only from southwestern

Japan. A key to Japanese adeonid species is provided.

Keywords. Adeonella , Adeonellopsis , diversity, distribution, Japan.

Hirose M. 2016. Diversity and distribution of adeonid bryozoans (Cheilostomata: Adeonidae) in Japanese waters.

European Journal of Taxonomy 203: 1-41. http://dx.doi.org/10.5852/eit.2016.2Q3

Introduction

Order Cheilostomata in Class Gymnolaemata is the most speciose group of bryozoans in modern marine

enviro nm ents (Bock & Gordon 2013). While most cheilostome colonies are encrusting, forming an

essentially 2-dimensional layer on the substrate, there are also many types of erect morphology in

which the colony rises above the substrate in 3-dimensional space. Large, erect bryozoan colonies have

contributed to the formation of bryozoan reefs or thickets from the Paleozoic to the present (Cuffey

1974, 1977; Lombardi et al. 2014; McKinney & Jackson 1989; Scholz et al. 2005; Probert & Batham

1979; Carter et al. 1985; Batson & Probert 2000; Taylor & James 2013; Wood & Probert 2013). Adeonid

bryozoans (e.g., Adeonella Busk, 1884, Adeona Lamouroux, 1812, Adeonellopsis MacGillivray, 1886)

form large, erect colonies with many flat, narrow, dichotomous branches, or are cribrate in form, which

can be viewed as derived from dichotomously branched colonies, and contribute to assemblages of erect

bryozoans in Japan (Hirose et al. 2012).

1

European Journal of Taxonomy 203 : 1-41 ( 2016 )

The family Adeonidae consists of approximately 10 genera and 106 species (Bock & Gordon 2013). One

of the largest genera in the family, Adeonellopsis, comprises about 50 species that are almost globally

distributed but are absent from higher latitudes (or alternatively: boreal and arctic waters). Another large

genus, Adeonella, also comprises about 50 species, distributed in the Indo-West Pacific, Mediterranean,

and Atlantic. Adeonellopsis has been reported from Eocene deposits and is common throughout the

later Cenozoic to Recent. Adeonella has also been reported from Eocene deposits in Europe, but is

more common in the present-day fauna. Hayward & Cook (1979, 1983) and Hayward (1981, 1988)

described 27 new species of Adeonella from South Africa, where its diversity is remarkably high on the

east coast (Hayward 1983, 1988). Compared to Adeonella, Adeonellopsis is more common and diverse

in Australia and throughout the Indo-West Pacific. The Adeonidae have received a relatively great

amount of attention over the past decades, a number of new species were introduced, and several known

species revised (e.g., Cook 1982; Aristegui 1985; Hayward 1988; Amui 2005; Rosso & Novosel 2010;

Almeida et al. 2015). Moreover, owing to their complexity at zooidal and zoarial level, the ontogeny and

astogeny of the Adeonidae is fairly well known (e.g.. Cook 1973; Cheetham & Hayek 1983; Wass 1991;

Lidgard & Buckley 1994; Lidgard 1996; Bock & Cook 2000, 2004; Smith et al. 2001).

Busk (1884) first described Adeonella as a genus in the family Adeonidae. Gregory (1893) later

established the family Adeonellidae fox Adeonella and related genera. Some researchers followed Busk’s

original classification (Canu & Bassler 1920, 1923, 1929; Harmer 1957), and Cook (1973) reported

significant differences in frontal shield structure and development between Adeonella and the other

genera in Adeonidae, and regarded Adeonella as clearly belonging in Adeonellidae. Recently, however,

Adeonellidae has been considered as a junior synonym of Adeonidae (Bock & Gordon 2013).

There are some previous records of adeonid bryozoans in Japan. Ortmann (1890) reported three species

from Sagami Bay ( Adeonella tuberculata Busk, 1884, and two new species, Adeonella japonica

Ortmann, 1890 and Adeonella sparassis Ortmann, 1890). Ortmann’s type material, however, has not

been restudied in detail. Okada (1920) redescribed A. japonica from Sagami Bay and described a new

species, Adeonella hexangularis Okada, 1920, from the vicinity of the Miura Peninsula adjoining

Sagami Bay. Subsequently, Okada & Mawatari (1938) reported A japonica from Wakayama Prefecture.

Mawatari (1952) also reported A. japonica and A. hexangularis from Wakayama Prefecture, without any

descriptions. The taxonomy of the Japanese adeonid bryozoans has previously not been reviewed and is

still largely dependent on the limited descriptions in Ortmann (1890), and their diversity and distribution

around Japan remains poorly known.

For this study, I re-examined Ortmann’s (1890) type specimens and examined other material collected

in Japan over the past 130 years, as well as new material I collected personally. Here I review and

describe or redescribe eight adeonid species in two genera from Japan, of which Adeonella jahanai

sp. nov., Adeonellopsis parvirostrum sp. nov., and Adeonellopsis toyoshioae sp. nov. are described as

new species, and summarize the distributions of all eight species around Japan.

Material and methods

Material examined

I examined specimens from Sagami Bay and surrounding areas (Appendix 1; Fig. 1A-B) collected by

Ludwig Doderlein (1880-1881), Emperor Showa (1918-1971), and the National Museum of Nature

and Science Tokyo (2001-2005; 2006-2010); see National Museum of Nature and Science (2007),

Hirose (2010), and Spencer Jones et al. (2011) for historical overviews. The material is housed at the

Musee Zoologique Strasbourg (MZS) and the National Museum of Nature and Science Tokyo (NSMT),

which is now located in Tsukuba (Appendix 1). At the Seto Marine Biological Laboratory (SMBL) of

Kyoto University, I examined a specimen of Adeonella japonica collected from the vicinity of the Kii

2

HIROSE M., Japanese adeonid bryozoans

Peninsula and reported by Olcada & Mawatari (1938). I also examined adeonid specimens collected by

the R/V Albatross from Japan and Hawaii and now housed in the National Museum of Natural History

(USNM), Washington, D.C.

I collected additional specimens by dredge, grab, and beam trawl from several localities in Japan

(Fig. 1 A, C; Appendix 2), including Otsuchi (Iwate Prefecture), Maizuru (Kyoto Prefecture), the Sagami

Sea, north of Hachijo-jima (south of the Sagami Sea), southwest of Kochi (Kochi Prefecture), the Ariake

Sea, Koshiki Strait (west of Kyushu), and the Nansei Islands from Tanegashima to Okinawa. All of this

material has been deposited in the NSMT in Tsukuba.

Preparation and observation of specimens

Specimens were observed by light microscope and scanning electron microscope (SEM) at Hokkaido

University and at the Atmosphere and Ocean Research Institute of the University of Tokyo (AORI).

For SEM observation, part of each specimen was 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 at 15 kV accelerating voltage with a Hitachi S-3000N SEM;

at AORI, specimens were observed with a Hitachi Miniscope TM-1000 at 15 lcV accelerating voltage

without sputter-coating. All fragments removed from specimens in the various collections for SEM

observation were subsequently deposited in NMST.

Measurements were taken from SEM images with Image Jv. 1.37 software (Image Processing and Analysis

in Java, Wayne Rasband, National Institutes of Health, USA; http://rsb.info.nih.gov/ij/ ). Measurements

in the text are presented in micrometers, with the range followed by the mean and standard deviation (in

Z

co

o

O

CO

z

CD

CN

CO

CN

o

CD

CN

Fig. 1. Maps showing the localities where adeonid bryozoans were collected. A. Collecting localities

around Japan. Gray arrows indicate warm currents, the unfilled arrow indicates the cold current.

B. Enlargement from previous, showing the localities around Sagami Bay, Sagami Sea, and Hachijo-jima

Island. C. Enlargement from map A, showing the localities along the Nansei Islands, from Tanegashima

to Okinawa.

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European Journal of Taxonomy 203: \-A\ (2016)

parentheses). Sample sizes for measurements were n = 2-115, and measurements were generally taken

from more than one colony. Abbreviations used for characters measured are as follows:

FAvL, FAvW = frontal avicularium length and width

SAvL, SAvW = suboral avicularium length and width

SOrL, SOrW = secondary orifice length and width

SpL, SpW = spiramen length and width

ZL, ZW = zooid length and width

Results

Taxonomic descriptions

Order Cheilostomata Busk, 1852

Suborder Neocheilostomina d’Hondt, 1985

Infraorder Ascophorina Levinsen, 1909

Superfamily Adeonoidea Vigneaux, 1949

Family Adeonidae Busk, 1884

Remarks

This family consists of approximately 10 genera and 106 species (Bock & Gordon 2013). Most genera

form erect colonies with flat, dichotomous branches (e.g., Adeonella and Adeonellopsis) or are cribrate

in form (e.g., Adeona ), although some genera form encrusting colonies (e.g., Reptadeonella Busk,

1884). Adeonella and Adeonellopsis differ in the type of frontal shield: Adeonella has a lepralioid

shield in which a spiramen leads into the space above the sinus, and a distinct primary orifice, while

Adeonellopsis is thought to have an umbonuloid frontal shield and the spiramen leads into the space above

the frontal membrane (Cook 1973; Lidgard 1996; Berning et al. 2014). Adeonella and Adeonellopsis

are also different in the arrangement of the spiramen and frontal/suboral avicularia; Adeonella generally

has a monoporous spiramen and frontal avicularia in various positions, whereas Adeonellopsis has

a monoporous or multiporous spiramen, with a suboral avicularium situated distal to the spiramen,

directed distally or distolaterally.

Genus Adeonella Busk, 1884

Adeonella cf. lichenoides (Lamarck, 1816)

Figs 2-3

cf. Eschara lichenoides Lamarck, 1816: 176.

cf. Escharaplatalea Busk, 1854: 90, pi. 105, figs 1-3, pi. 108, fig. 4.

cf. Adeonella lichenoides -Harmer 1957: 799. —Busk 1854: 90, pi. 106, figs 1-3. —Hayward 1988:

126, figs 1C, 2-3.

cf. Adeonella platalea - Busk 1884: pi. 184, pi. 21, figs 4, 4a. — Harmer 1957: 809, pi. LIU, figs 2,

4-12. —Hayward 1983: 582, fig. 1A, C-D.

Adeona japonica - Mawatari 1952: pi. 12, fig. 1 [figured specimen].

Material examined

JAPAN: Izu, Sagami Sea, Emperor Showa collection (NSMT-BryR292, BryR297, BryR299, BryR303,

BryR304, BryR315, BryR317, BryR324 (SEM specimen only), BryR357 (SEM specimen only),

BryR358, BryR359, BryR360 (SEM specimen only), BryR365, BryR367, BryR377); Seto, Kii Peninsula

(SMBL-Brz.17, some fragments NSMT-Tel049), Aug. 1936, probably specimen studied in Olcada &

4

HIROSE M., Japanese adeonid bryozoans

Mawatari (1938); Sagami Bay and Sagami Sea, collected by NSMT (NSMT-TeS13, TeS14, TeS30

(SEM specimen only)); off Shimoda, Izu Peninsula (NSMT-Te891); Koshiki Strait (NSMT-Te752);

Ogasawara (NSMT-Te753, Te754, Te755, Te756).

Measurements

Autozooids. ZL: 298-526 (411±44); ZW: 203-317 (239±21); n = 39. SOrL: 44-92 (68±12); SOrW:

60-101 (77±10); n = 38. FAvL: 53-86 (68±9); FAvW: 25-47 (37±5); n = 32. SpL: 22-39 (30±5); SpW:

22-37 (28±6);n= 18.

Gonozooids.ZL: 456-521 (488±32);ZW: 290-334(318±24);n = 3. SOrL: 75-77 (76±1); SOrW: 93-122

(111±16); n = 3. FAvL: 61-67 (63±3); FAvW: 25-36 (29±5); n = 4. SpL: 34-38 (36±2); SpW: 35-81

(61±23); n = 3.

Description

Colony brown, erect, dichotomously branching, widely and irregularly spreading, forming a three-

dimensional bushy structure; up to 12 cm in diameter and 10 cm high (Fig. 2A). Branches flattened,

multiserial, with zooids opening on both sides, 1.9—4.1 mm wide (average 2.7 mm; n = 39); rounded at

end. Autozooids oval or rhomboidal, surrounded by distinct, deep marginal groove (Fig. 2B-C). Frontal

c

- A CJ

y%' Tl.

2 .

Fig. 2. Adeonella cf. lichenoides (Lamarck, 1816). A. Large, robust branching colony in Showa Emperor

Collection (NSMT-BryR299, Sagami Sea). B. Distal end of branch, showing vicarious avicularia

associated with a branch bifurcation (NSMT-BryR297, Sagami Sea). C. Autozooids (NSMT-BryR359,

Sagami Sea). D-E. Two types of lanceolate avicularia near branch bifurcations. D. Short type (NSMT-

BryR297). E. Long type (NSMT-BryR359, Sagami Sea). A = optical photograph; B-E = SEM images.

Scale bars: A = 2 cm; B = 500 pm; C-E = 200 pm.

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European Journal of Taxonomy 203: 1-41 (2016)

shield convex, entirely covered with minute granules, with about 10 small, circular areolar pores inside

each lateral margin, some additional pores in central region, and one large peristomial spiramen proximal

to orifice (Fig. 2C). Peristome deep, secondary orifice circular or transversely oval. Frontal avicularia

small, 1 to 4 in number, occurring near both margins on frontal shield and oriented inward (Fig. 2C) or

distally (Fig. 3C). Gonozooids (Fig. 3A) present in broad region at branch bifurcations and at periphery

of branches. Gonozooids larger than autozooids, with broader orifice and swollen, porous frontal shield;

peristomial spiramen transversely broad, with median projection from distal margin (Fig. 3A). Vicarious

avicularia occur near branch bifurcations, often replaced by gonozooids; elongate, as long as or sometimes

longer than autozooids (Fig. 2B), 316-495 pm (average 390 pm) long by 124-164 pm (average 140 pm)

wide (n = 6). Vicarious avicularian chamber large (Fig. 2D-E); 425-814 pm (average 572 pm) long by

228-340 pm (average 274 pm) wide (n = 6). Rostrum of vicarious avicularia lanceolate, acute, directed

distally; variable in form and length, up to 400 pm long. Large, triangular vicarious avicularia also occur

along branch edges. In basal part of branches and colony, autozooids often replaced by kenozooids (Fig.

3B); 342-460 pm (average 388 pm) long by 167-234 pm (average 192 pm) wide (n = 4).

Remarks

This species was previously known as Adeonella platalea (Busk, 1854) in Japan. Lamarck (1816) first

described Eschara lichenoides from the Indian Ocean, and Harmer (1957) transferred it to Adeonella.

Harmer (1957) discussed the high variability in A. lichenoides and A. platalea ; although he did not

synonymize these species, he noted three groups of A. platalea based on variant forms of vicarious

avicularia and mandible. Although Harmer’s concept of the nature of intra- vs interspecific variation was

different from that of more recent authors, Hayward (1988) restudied a syntype of Eschara lichenoides

and found this specimen to be nearly identical to the paratype of A. platalea , and thus considered

A. lichenoides to be a subjective senior synonym. Morphological variation within the extremely broad

recorded distribution of this species from East Africa to eastern Australia, however, might indicate the

presence of two or more unrecognized species.

Mawatari (1952) reported Adeona (Adeonellopsis) japonica from offMinabe and Shirahama, Wakayama

Prefecture, without any description but with a photograph of a single branch in which zooids have a

swollen frontal shield with a single spiramen, quite resembling Adeonella. The photograph in Mawatari

(1952) may be of a branch of A. cf lichenoides (Brz. 17) in the SMBL collection.

Fig. 3. Adeonella cf. lichenoides (Lamarck, 1816). A. Gonozooids at branch bifurcation (NSMT-Bry

R 365, Sagami Sea). B. Kenozooids with numerous frontal avicularia (NSMT-BryR360, Sagami Sea).

C. Autozooids with numerous frontal avicularia (NSMT-TeS14, Sagami Sea). SEM images. Scale bars:

A, C = 200 pm; B = 100 pm.

6

HIROSE M., Japanese adeonid bryozoans

Distribution

Adeonella lichenoides has previously been reported from the Indo-West Pacific (Philippines, Malay

Archipelago, Queensland coast, Torres Strait, northern and western coasts of Australia and Victoria, and

eastward to Zanzibar and East Africa) (Hayward, 1988). In Japan, Adeonella cf. lichenoides has been

collected from Sagami Bay, Sagami Sea, around the Izu Peninsula, off the Kii Peninsula, the west coast

of Kyushu (Koshiki Strait), and Ogasawara, at depths of 3-328 m. Although Adeonella cf lichenoides

was not collected from the eastern part of Sagami Bay by Doderlein or Doflein, it was very abundant

in the western Sagami Sea (in ES collection) such as exposed shallow rocky habitat (3—45 m) at the

southernmost part of the Izu Peninsula (Hirose et al. 2012).

Adeonella jahanai sp. nov.

urn:lsid:zoobank.org:act:36CA23B8-7008-4562-9E24-D29AD412BE0C

Figs 4-5

Diagnosis

Colony dichotomously branching; branches narrow, about 1.2-2.5 mm wide. Autozooids oval or

hexagonal, distinct, delineated by deep groove. Frontal shield entirely covered with minute granules,

with more than 10 circular or slit-like areolar pores. Peristome deep, secondary orifice circular, primary

Fig. 4. Adeonella jahanai sp. nov. A. Holotype colony with narrow branches (NSMT-Tel050, Okinawa).

B. Young autozooids at distal end of branch, showing primary orifices NSMT-Te889. C. Autozooids

with elongate frontal avicularia (NSMT-Te889, Okinawa). D. Young autozooids with a single triangular

avicularium abutting the proximal margin of the peristome (NSMT-Te889, Okinawa). A = optical

photograph; B-D = SEM images. Scale bars: A = 1 cm; B-D = 100 pm.

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European Journal of Taxonomy 203: 1-41 (2016)

orifice with a deep and broadly U-shaped sinus, condyles short but broad and blunt. Many irregular

frontal pores in central region. Frontal avicularia triangular, narrow, elongate; in center or at proximal

margin of frontal shield, directed inward or proximally. Another frontal avicularium abuts proximal

margin of peristome, rostrum slightly projected, directed laterally. Large, triangular vicarious avicularia

Fig. 5. Adeonella jahanai sp. nov. A. Branch bifurcation lacking vicarious avicularaia (NSMT-

Te889, Okinawa). B. Branch bifurcation with vicarious avicularium (NSMT-Te749, off Yakushima).

C. Vicarious avicularia on side of branch (NSMT-Te889, Okinawa). D. Old part of branch showing

autozooids and kenozooids (NSMT-Te889, Okinawa). E. Gonozooids at periphery of branch (NSMT-

Te889, Okinawa). F. Gonozooid at periphery of branch (NSMT-Te889, Okinawa). SEM images. Scale

bars: A- D = 200 pm; E = 300 pm; F = 100 pm.

8

HIROSE M., Japanese adeonid bryozoans

present along branch margins and at bifurcations. Gonozooids slightly larger than autozooids, with

porous frontal shield; secondary orifice broad, with slightly convex proximal margin.

Etymology

The species is named after Mr. Yoshikazu Jahana, the fisherman who collected the holotype specimen

off Motobu, Okinawa.

Material examined

Holotype

JAPAN: large colony collected 19Mar. 2014 north of Motobu, Okinawa, 26°42'54.72" N, 127°53'25.62"E,

60 m depth, by basket net (NSMT-Tel050).

Paratypes

JAPAN: several branches, same collection data as for holotype (NSMT-Te889); single small branch

collected 25 May 2009 southwest of Yalcushima, 30°05'08.40" N, 130°04'43.20" E, 87-88 m depth,

by dredge, T/V Toyoshio-maru (NSMT-Te749); many branches collected 23 May 2010 south of

Nagannu Island, Okinawa, 26°14'37.80" N, 127°32'28.20" E, 37—46 m depth, by dredge, T/V Toyoshio-

maru (NSMT-Te775); some fragments collected 21 May 2011 south of Nagannu Island, Okinawa,

26°14'39.00" N, 127°32'55.20" E to 26°14'43.20" N, 127°31'48.60" E, 47-52 m depth, by dredge,

T/V Toyoshio-maru (NSMT-Te751); some fragments collected 27 May 2012 south of Nagannu Island,

Okinawa, 26°14'34.68" N, 127°32T.26" E to 26°14'36.66" N, 127 0 3E54.18" E, 52-54 m depth, by

dredge, T/V Toyoshio-maru (NSMT-Te804); some fragments collected 27 May 2012 east of Maeshima

Island, Okinawa, 26°1E41.40" N, 127°27'20.34" E to 26°1E39.90" N, 127°27T8.36" E, 55-57 m depth,

by dredge, T/V Toyoshio-maru (NSMT-Te805); some fragments collected 28 May 2012 south of Iejima

Island, Okinawa, 26°39T4.28" N, 127 0 4E47.52" E to 26°39 , 19.38" N, 127°4E52.98" E, 94-95 m depth,

by dredge, T/V Toyoshio-maru (NSMT-Te806).

Measurements

Autozooids. ZL: 374-508 (437±39); ZW: 196-304 (242±31); n = 17. SOrL: 67-99 (82±13); SOrW:

78-92 (85±5); n = 7. FAvL: 71-121 (94±18); FAvW: 36-46 (39±3); n = 9. SpL: 24-35 (30±4); SpW:

24-32 (28±2); n = 7.

Gonozooids. ZL: 455-666 (515±79); ZW: 250-335 (285±37); n = 6. SOrL: 57-96 (71±14); SOrW: 95-

128 (103±12); n = 6. SpL: 19-27 (23±3); SpW: 38-49 (42±4); n = 6.

Description

Colony brown, erect, dichotomously branching, widely and irregularly spreading, forming a bushy

three-dimensional structure; up to 6 cm in diameter and 5 cm high (Fig. 4A). Branches slender, nearly

uniform in width, flattened, multiserial, with zooids opening on both sides, 1.1-2.4 mm wide (average

1.7 mm; n = 34); rounded at tip. Autozooids oval or hexagonal, surrounded by a distinct marginal groove

(Fig. 4C-D). Frontal shield convex, entirely covered with minute granules, with about 10-20 small,

circular or slit-like areolar pores inside each lateral margin, many additional irregular pores in central

region, and one large peristomial spiramen proximal to orifice (Figs 4C, 5B). Peristome deep, secondary

orifice circular or transversely oval. Primary orifice deeply imm ersed in peristome, visible only in early

ontogenetic zooids; with a deep and broadly U-shaped sinus and conspicuous condyles (Fig. 4B), 59-78

pm (average 72 pm) long by 71-90 pm (average 82 pm) wide (n = 5). Frontal avicularia triangular,

narrow, elongate, single or paired, occurring in center of frontal shield or toward proximal margin and

oriented proximally or inward (Fig. 4C). Another frontal avicularium abutting the proximal margin of

peristome, rostrum slightly projected, directed laterally (Fig. 4D). Gonozooids present at periphery of

branches (Fig. 5E). Gonozooids larger than autozooids, with broader orifice and porous frontal shield;

9

European Journal of Taxonomy 203: 1^1 (2016)

peristomial spiramen transversely broad, with median projection from distal margin (Fig. 5F). Vicarious

avicularia rarely present near branch bifurcations, often lacking; elongate, as long as or sometimes

longer than autozooids (Fig. 5B), 384-617 pm (average 500 pm) long by 126-158 pm (average 142 pm)

wide (n = 2). Rostrum of vicarious avicularia lanceolate, acute, directed distally; up to 410 pm long

(348±97, n = 2). Large vicarious avicularia also occur along branch edges (Fig. 5C), up to 579 pm long

(572±9, n = 2) by 142 pm wide (139±5, n = 2). In basal part of branches and colony, autozooids are

often replaced by kenozooids (Fig. 5D); 380-416 pm (average 400 pm) long by 252-304 pm (average

284 pm) wide (n = 5).

Remarks

Adeonella jahanai sp. nov. resembles A. lichenoides in having a granulated frontal shield, and in the

location of the small, triangular frontal avicularium, but differs from the latter in having markedly

narrower branches, numerous slit-like areolar pores on the frontal shield, elongate frontal avicularia,

and the frontal avicularia directed proximally. Autozooids of Adeonella jahanai sp. nov. also differ from

A. lichenoides in having another frontal avicularium, triangular, slightly projected, directing laterally,

abutting the proximal margin of the peristome. Adeonella jahanai sp. nov. resembles A. extensa Harmer,

1957 in having narrow branches, gonozooids along the margin of branches, and slightly longer frontal

avicularia, and in generally lacking vicarious avicularia near branch bifurcations, but differs from the

latter in the direction of the frontal avicularia, and in having gonozooids with a non-tubular peristome,

with the peristomial spiramen being very closely situated to the secondary orifice.

Distribution

Japan: Nansei Islands (near Yakushima and west of Okinawa), at depths of 37-95 m.

Genus Adeonellopsis MacGillivray, 1886

Adeonellopsis arculifera (Canu & Bassler, 1929)

Fig. 6

Adeona arculifera Canu & Bassler, 1929: 377, pi. 52, figs 1-2.

Adeonellopsis arculifera -Harmer 1957: 800, text-figs 82C, 84A, pi. LIU, figs 13-14.

Adeonella sub sulcata -Thornely 1905: 125.

Material examined

JAPAN: many branches collected from Hirase, southwest of Yakushima Island, 87-88 m depth (NSMT-

Te781, Te783, Te784, Te785, Te786, Te787, Te788, Te789); single colony collected from South of

Nagannu Island, 47-52 m depth (NSMT-Te782).

Measurements

Autozooids. ZL: 505-762 (590±62); ZW: 207-387 (288±53); n = 39. SOrL: 54-91 (71±10); SOrW:

62-103 (83±9); n = 30. SAvL: 76-147 (100±13); SAvW: 50-85 (64±7); n = 38. SpL: 12-27 (20±4);

SpW: 16-25 (20±3);n = 28.

Description

Colony small, slender, dichotomously branching (Fig. 6A); yellowish white. Branches subcylindrical

or flat (Fig. 6B); slender, on average 1 mm wide (0.9-1.3 mm, n = 8), multiserial, zooids opening

on both sides. Zooids oval, boundaries indistinct, delineated by shallow groove and opposing single

rows of tiny, irregular areolar pores (Fig. 6D). Frontal shield smooth, or finely granulate and slightly

10

HIROSE M., Japanese adeonid bryozoans

wrinlded. Zooids raised distally towards a low peristomial rim; primary orifice D-shaped, with straight

or convex proximal margin; secondary orifice broader than long. Spiramen small, uniporous, circular,

situated close to center of frontal shield proximal to orifice, often in depressed frontal zone (Fig. 6C).

Fig. 6. Adeonellopsis arculifera (Canu & Bassler, 1929). A. Colonies collected SW of Yakushima

Island (NSMT-Te789). B. Branch showing crenulate periphery (NSMT-Te781). C. Enlargement of

bifurcation showing autozooids with oblique avicularium and single small spiramen (NSMT-Te781).

D. Young autozooids showing peristomial rim and single small spiramen (NSMT-Te781). E. Old part

of branch, showing various stages in formation of kenozooids (NSMT-Te781). F. Old part of branch,

showing autozooids with depression containing avicularium and spiramen (NSMT-Te781). A = optical

photograph; B-F = SEM images. Scale bars: A = 5 mm; B = 1 mm; C-D = 300 pm; E = 500 pm; F

250 pm.

11

European Journal of Taxonomy 203: 1-41 (2016)

Suboral avicularium abutting proximal margin of peristome, smaller in area than secondary orifice,

rostrum slightly raised and sometimes hooked distally, slightly curved toward distal side, directed

distolaterally; no crossbar; mandible equilaterally triangular (Fig. 6C). No other types of avicularia

observed. Autozooids distal to branch bifurcations elongate, sometimes bearing up to three umbonate

bosses 47-76 pm in diameter (average 61 pm, n = 18), on one or both sides proximolateral to orifice and

toward proximal end of frontal shield (Fig. 6D). Older autozooids in basal part of branch broad, irregular

in outline; zooidal boundaries indistinct; frontal shield heavily secondarily calcified; spiramen often

lacking; suboral avicularium lies in depression in thickened frontal shield (Fig. 6F). Rarely, kenozooids

occur at basal bifurcations (Fig. 6E); same size as autozooids. Gonozooids not found.

Remarks

Canu & Bassler (1929) originally described Adeona arculifera from Jolo and Sulade Islands, Sulu

Archipelago, Philippines, on the basis of the single spiramen, although the branched colony resembles

the condition in Adeonellopsis. Harmer (1957) then transferred the species to Adeonellopsis on the

basis of the operculum, which is not widely sinuate. Adeonellopsis arculifera resembles A. subsulcata

(Smitt, 1873) in the arrangement of the suboral avicularia but differs from the latter in having a circular

spiramen in the center of the frontal shield.

Distribution

Adeonellopsis arculifera has previously been reported from the Philippines, Indonesia, and the Indian

Ocean, at depths of 38-53 m. In Japan, it has been detected only in the southwestern region, from

Okinawa and near Yakushima Island, at depths of 47-88 m. Although this is the first Recent record for

Japan, Hay ami (1971) reported the species from the Pliocene Shinzato Tuff on Okinawa Island.

Adeonellopsis japonica (Ortmann, 1890)

Figs 7-9, 10A

Adeonella japonica Ortmann, 1890: 54, pi. 4, fig. 11.

Adeonella japonica - Okada 1920: 628, text fig. 6, pi. 8, fig. 9. — Okada & Mawatari 1935: 140, pi. XI,

fig. 2. — Okada & Mawatari 1938: 460.

Adeona japonica - Canu & Bassler 1927: 34. — Mawatari 1952: 285 [figured specimen is not

A. japonica],

Adeonellopsisparvipuncta '-Harmer, 1957: 798, pi. LIII, fig. 16.

Adeonellopsis japonica - Cheetham et al. 1980: 335-369. —Hirose et al. 2012: 121-136, fig. 9.2.

non Adeonellopsis parvipuncta MacGillivray, 1886: 135, pi. II, fig. 4a.

Material examined

Lectotype [designated herein]

JAPAN: branched colony, Sagami Bay, 1882, 111-277 m depth, collected by L. Doderlein, MZS 3-3

(SEM specimen NSMT-Te741).

Paralectotypes [designated herein]

JAPAN: branched colonies, Jogashima, 1882, 185-370 m depth, collected by L. Doderlein, MZS 3-4

(SEM specimen NSMT-Te742); branched colony, Sagami Bay, 1882, collected by L. Doderlein, MZS

3-1; branched colonies, Sagami Bay, MZS 3-2 (SEM specimen NSMT-Te739), “15 August 1889/29 July

1890, 111 m depth, southeast of Jogashima, Sagami Bay” is written on a small label, but the collector

and the date are unclear).

12

HIROSE M., Japanese adeonid bryozoans

Fig. 7. Adeonellopsisjaponica (Ortmann, 1890). A. Lectotype from among Ortmann’s specimens in the

Doderlein Collection (MZS 3-3). B. Colonies collected at Otsuchi (NSMT-Te760). C. Autozooids with

large suboral avicularium and biporous spiramen (NSMT-Te741, from lectotype, MZS 3-3). D. Lateral

vicarious avicularia at branch bifurcation (NSMT-Te741, from lectotype, MZS 3-3). E. Young autozooids

at distal end of branch (NSMT-Te741, from lectotype, MZS 3-3). F. Young autozooids, with biporous

spiramen surrounded by thick, swollen rim bearing rows of granulation (NSMT-Te741, from lectotype,

MZS 3-3). G. Autozooids and gonozooids at branch bifurcation (NSMT-Te742, from paralectotype

MZS 3-4). H. Gonozooids showing large multiporous spiramen (NSMT-Te742, from paralectotype

MZS 3-4). A-B = optical photographs; C-H = SEM images. Scale bars: A = 2 cm; B = 1 cm; C-D, F,

H = 200 pm; E = 300 pm; G = 500 pm.

13

European Journal of Taxonomy 203: 1-41 (2016)

Other material examined

JAPAN: eastern side of Sagami Bay (NSMT-BryR36, Bry R71, Bry R193, BryR206, BryR225, BryR254,

BryR255, BryR259, BryR260, BryR261, BryR277, all in Emperor Showa collection); eastern side of

Sagami Bay, collected by NSMT (NSMT-TeS15 to TeS18); NE off Ohakozaki, Otsuchi, collected by

Fig. 8. Adeonellopsis japonica (Ortmann, 1890). A. Distal end of branch just starting to bifurcate,

showing young autozooids and gonozooids (NSMT-BryR277). B. Enlargement of young gonozooids,

showing peripheral rim with rows of granulation perpendicular to margin, and large multiporous

spiramen (NSMT-BryR277). C. Autozooids in middle of branch, showing imm ersed spiramen (NSMT-

Bry R 193). D. Basal part of branch, showing autozooids, and some kenozooids with closed orifice

(NSMT-BryR36). E. Young gonozooids, showing distinct marginal pores and multiporous spiramen

(NSMT-Te758). F. Distal end of branch, showing autozooid formation (NSMT-Te758). SEM images.

Scale bars: A, C = 1 mm, B, D-E = 300 pm; F = 500 pm.

14

HIROSE M., Japanese adeonid bryozoans

M. Hirose (NSMT-Te758, Te759, Te760); west-southwest of Jogashima, Sagami Bay, collected by H.

Koutsuka and M. Hirose (NSMT-Te795, Te798); south of Kanae-zaki, Tosashimizu, Kochi (NSMT-

Tel051); Seto, Kii Peninsula, Aug. 1936, probably specimen studied by Okada & Mawatari (1938)

(SMBL-Brz.17, some fragments NSMT-Tel052); Pacific Ocean, off Honshu Island, 7 May 1900,

Albatross Station D.3704, 107-275 m depth (USNM PAL 271600A-D).

HAWAFI: off Oahu Island, Albatross Station D.3916, 549-605 m depth, collected 6 May 1902 (USNM

PAL 271601); south coast of Oahu Island, Albatross Station D.3916 (USNM RB 8450).

Measurements

Autozooids. ZL: 404-750 (553±71); ZW: 184-419 (308±52); n = 115. SOrL: 69-109 (87±11); SOrW:

56-127 (102±14); n = 67. SAvL: 134-232 (176±20); SAvW: 62-133 (90±14); n = 81. PAvL: 75-138

(110±15); PAvW: 40-72 (59±8); n = 47. SpL: 16-92 (43±16); SpW: 20-75 (44±14); n = 70.

Fig. 9. Adeonellopsis japonica (Ortmann, 1890). A. Colony collected at Albatross Station D, off Oahu

Island, Hawaii (USNM 271601). B. Autozooids of Hawaii specimen, showing suboral avicularia and

biporous spiramen (USNM 271601, Hawaii). C. Autozooids and lateral vicarious avicularia at a branch

bifurcation (USNM 271601 paper box, Hawaii). D. Enlargement of lateral vicarious avicularia at branch

bifurcation (USNM 271601 paper box, Hawaii). A = optical photograph; B-D = photomicrographs.

Scale bars: A = 5 mm; B, D = 300 pm; C = 500 pm.

15

European Journal of Taxonomy 203: 1-41 (2016)

Gonozooids. ZL: 453-722 (612±60); ZW: 329-524 (415±53); n = 35. SOrL: 57 -125 (86±13); SOrW:

100-166 (134±14); n = 34. SAvL: 109-176 (135±17); SAvW: 52-104 (73±12); n = 25. SpL: 102-205

(146±24); SpW: 98-174 (134±21); n = 33.

Vicarious avicularia at branch margins. L: 358-399 (385±23); W: 141-172 (158±16); n = 3.

Description

Colony yellowish brown, erect, dichotomously branching, irregularly spreading; up to 10 cm high

(Fig. 7A). Branches flattened (Figs 7B, 9A), multiserial, with zooids opening both sides, rounded at

end; 1.6—4.6 mm wide (average 2.8 mm; n = 27). Autozooids oval or hexagonal, outlined by a distinct

deep marginal groove (Figs 7C, 8C, 9B), with 6-9 small pores along each lateral margin and two or three

small frontal pores just proximal to orifice. Frontal shield umbonuloid (Fig. 10A). Young autozooids at

growing margin of branch have concave frontal shields surrounded by broad, swollen rim with rows

of granulation arranged perpendicular to margin (Fig. 7E-F); rim width 41-76 pm (average 57 pm) (n

= 31). Mature frontal shield convex, entirely covered with minute granules. Spiramen abuts proximal

end of suboral avicularium; subdivided into two or three openings separated by narrow septa, or lobate

due to incomplete septa (Fig. 7C); with increased calcification, sometimes appearing as single opening

(Fig. 7D); reduced in size during ontogeny, associated with depression of avicularian rostrum, sometimes

occluded (Figs 7D, 8D). Secondary orifice nearly circular (Fig. 7C) or broader than long (Figs 7D,

8C); peristomial rim lacking; primary orifice submerged deep in peristome. Large, suboral avicularium

in center of frontal shield, extending from spiramen to proximolateral corner of orifice, or nearly so,

usually pointing distolaterally (Figs 7C, F, 9B); rostrum slightly raised distally, not hooked; mandible

elongate-triangular; no crossbar. Additional smaller frontal avicularium often present near proximal end

of zooid; identical in form to suboral avicularium; pointing in any direction (Fig. 1C). Gonozooids occur

at branch bifurcations (Fig. 8B, E); larger than autozooids, with broader, slightly curving orifice; bearing

suboral avicularium; smaller frontal avicularia either lacking (Fig. 8B, E) or up to three per zooid (Fig.

7G-H). Spiramen large, roughly circular in outline, divided into 6-10 pores (Figs 7G-H, 8B). Spiramen

lacking marginal denticles in both autozooids and gonozooids (Figs 7C, F, 8E), although incomplete

septa may appear as denticles. Vicarious avicularia at branch margins (Figs 7D, 8A, 9C-D) smaller

than autozooids; triangular, with mandible directed distally. Basal part of branch consists of kenozooids

with secondary orifice occluded by secondary calcification (Fig. 8D); about same size as autozooids,

486-631 pm long (average 541 pm) by 199-351 pm wide (average 314 pm) (n = 11). Kenozooids also

Fig. 10. Interior frontal shield. A. Adeonellopsis japonica (Ortmann, 1890) (NSMT-Tel052). B. A. parvi-

rostrum sp. nov. (NSMT-Te807). Arrowheads indicate ring scar. SEM images. Scale bars: A = 50 pm;

B = 25 pm.

16

HIROSE M., Japanese adeonid bryozoans

bearing several small avicularia budded from areolae, near proximal margin of zooid, associated with

other marginal pores (Fig. 8D), 101-152 pm long (average 121 pm) by 47-75 pm wide (average 59 pm)

(n = 15).

Remarks

Ortmann (1890) first described Adeonellopsis japonica as Adeonella japonica, based on Doderlein’s

specimens from Sagami Bay. Hayward (1988) discussed this species in his revision of Adeonella, though

he did not examine Ortm arm’s material; he argued that Adeonella japonica almost certainly belongs in

Adeonella because of the elongate avicularia along the edge of the branches, as described by Ortmann.

However, the frontal shield of A. japonica is umbonuloid, rather than the lepralioid frontal shield

characteristic of Adeonella ; therefore, A. japonica clearly belongs in Adeonellopsis. Harmer (1957)

reported a Japanese adeonid species resembling Adeonellopsisparvipuncta MacGillivray (1886), which

has two kinds of spiramina: a small, undivided, slightly denticulate single pore, or a much larger pore

region comprising many pores. These features of the spiramen completely correspond to the spiramina

of autozooids and gonozooids in A. japonica. Harmer (1957) noted that a few zooids in the middle of

the branches have a larger spiramen, which is the condition of gonozooids in A. japonica. Furthermore,

his sketch of the larger zooid also indicates a broader, slightly curved orifice. I thus consider Harmer’s

A. parvipuncta to be a synonym of A. japonica.

Distribution

Japan: Sagami Bay, Sagami Sea, Suruga Bay, Kii Peninsula, Kouchi, and Otsuchi; 48.7—493 m depth.

The current species has also been collected from Hawaii (off Oahu Island), 549-605 m depth, by the

R/V Albatross (USNM 271601, IZ cat 8450; RB 8450) (Fig. 9).

Okada & Mawatari (1938) reported this species from Kata, Seto, and off Wakayama, around the

Kii Peninsula; the specimens in the SMBF include both A. japonica and Adeonella cf. lichenoides.

Mawatari (1952) also reported A. japonica from Wakayama Prefecture, without a description, but with

an illustration (pi. XII, fig. 1), which apparently shows proximally swollen zooids and the absence

of suboral avicularia, and is more similar to A. cf. lichenoides. Adeonellopsis japonica is one of the

common erect species in the Sagami Sea (Hirose et al. 2012).

Adeonellopsispentapora Canu & Bassler, 1929

Fig. 11

Adeonellopsis pentapora Canu & Bassler, 1929: 382, pi. 53, figs 1-5.

Adeonella tuberculata - Ortmann 1890: 53, pi. 4, figs 9a-b. — Okada 1923: 231. —Kataoka 1961: 264,

pi. XXXVII, fig. 2.

Adeonellopsis pentapora - Sakakura 1935: 115. —Kataoka 1961: 264, pi. XXXII, fig. 10.

Adeonellopsisyarraensis -Harmer 1957: 799, pi. Fill, figs 18-19 [in part]. —Hirose 2010: 52, pi. 89,

figs A-D.

Material examined

JAPAN: colony in Doderlein Collection at MZS, from Sagami Bay, 1882 (MZS2-1, MZS2-2; SEM

specimen NSMT-Te748); colonies from east side of Sagami Bay and Sagami Sea, collected by NSMT

(NSMT-TeS19, TeS20, TeS21, TeS22); specimens from off Ohakozaki, Otsuchi, 70-77 m depth (NSMT-

Te761, Te762, Te765, Te766); Ariake Sea (NSMT-Te767, Te769); Wakasa Bay, Sea of Japan, 70-85.7 m

depth (NSMT-Te800); off Shimoda (NSMT-Te892); near Yakushima and Tanegashima, 108-122 m

depth (NSMT-Te763, Te764); off Iejima Island, 94-95 m depth (NSMT-Te801); Ogasawara, 136-141 m

depth (NSMT-Te768).

17

European Journal of Taxonomy 203: 1-41 (2016)

Measurements

Autozooids. ZL: 337-572 (434±54); ZW: 227-367 (291±35); n = 34. SOrL: 42-95 (58±10); SOrW:

65-120 (89±13); n = 34. SAvL: 36-84 (61±9); SAvW: 25-51 (38±7); n = 36. SpL: 48-153 (88±24);

SpW: 43-111 (75±18); n = 37.

Fig. 11. Adeonellopsispentapora Canu & Bassler, 1929. A. Left, colony in Doderlein Collection (MZS

2-1); right, colony collected by NSMT (NSMT-TeS22). B. Colony on hydroid stalk, from Sagami Bay

(NSMT-TeS20). C. Distal end of branch showing young autozooids with multiporous, denticulate

spiramen (NSMT-Te761). D. Young autozooids with umbonate processes on frontal shield (NSMT-

Te764). E. Mature autozooids with small suboral and other frontal avicularia (NSMT-Te761). F. Old

autozooids, showing deeply immersed spiramen and orifices (NSMT-Te762). A-B = optical photographs;

C-F = SEM images. Scale bars: A = 3 mm; B = 2 mm; C-D, F = 200 pm; E = 250 pm.

18

HIROSE M., Japanese adeonid bryozoans

Description

Colony yellow or white, small, fragile, dichotomously branching, almost two-dimensional (Fig. 11A-

B); up to 3 cm high. Branches flat, slender, 1.3-2.6 mm wide (average 1.9 mm; n = 16). Autozooids on

both sides of branches; oval or hexagonal, zooidal boundaries distinct, zooids delineated by groove and

suture line (Fig. 11E). Frontal shield minutely granulate, with marginal pores; sometimes bearing 2 or

3 low, umbonate processes along lateral or proximal margin (Fig. 11D), 40-80 pm in diameter (average

60 pm, n = 14). Spiramen large, circular, divided into 4-6 small denticulate pores (Fig. 11D). Peristome

present, broadly oval in outline, not tubular (Fig. 11C); orifice indistinct in frontal view, proximal margin

highly convex (Fig. 11C). One or two small suboral avicularia between orifice and spiramen (Fig. 11E);

if single, positioned near midline and pointing distally or angled slightly distomedially or distolaterally;

if paired, pointing distomedially; mandible short-triangular; crossbar incomplete. One to three frontal

avicularia similar in size and shape to suboral avicularia situated around proximal and proximolateral

margins of zooid (Fig. 11D). Basal part of colony consists of autozooids and kenozooids; 309-521 pm

long (average 415 pm) by 202-383 pm wide (average 244 pm) (n = 14). Basal autozooids heavily

secondarily calcified; peristome and spiramen depression confluent, forming single oblong depression

that also contains the suboral avicularia (Fig. 1 IF), common opening 112-203 pm long (average 146 pm)

by 66-108 pm wide (average 89 pm) (n = 10). Gonozooids lacking. Marginal vicarious avicularia not

observed.

Remarks

Ortmann (1890) reported this species as Adeonella tuberculata (Busk, 1884) from Sagami Bay based on

Doderlein’s specimens, as it has two small avicularia between the orifice and spiramen. Adeonellopsis

pentapora is quite small and fragile compared to the other adeonid species collected from Sagami Bay;

it also occupies unstable substrates such as hydroid stems (Fig. 11B) and gastropod shells (Fig. 11 A).

Salcakura (1935) also reported a colony encrusting a gastropod shell from Toyama Bay. A colony of

A. pentapora in the Doderlein collection is encrusted on a small shell of a dead gastropod inhabited by

a hermit crab, and a colony collected from Sagami Bay in 2002 is encrusted on a different species of

small shell of a dead gastropod. Canu & Bassler (1929) reported Adeonellopsis pentapora from a broad

area of the Indo-Pacific, including Japan (Tsugaru Strait). Harmer (1957) considered Adeonellopsis

pentapora and A. tuberculata to be junior synonyms of Adeonellopsis yarraensis (Waters, 1881) based

on having two small suboral avicularia and narrow branches. A. yarraensis , however, is currently

regarded as a strictly fossil species and A. tuberculata as a Recent species in Australia; therefore, I

consider A. pentapora to be a valid species.

Distribution

Philippines and Japan: detected in this study from Sagami Bay, Sagami Sea, Korea Strait, Wakasa Bay,

Toyama Bay, Otsuchi, Ariake Sea, near Tanega-shima and Yaku-shima, and Ogasawara, at depths of

29-274 m. Adeonellopsis pentapora was not as abundant as other adeonid species in any collections or

at any localities. Kataolca (1961) reported a fossil specimen from the Pleistocene Ryukyu Fimestone of

Kikai-jima, Kagoshima.

Adeonellopsis sparassis (Ortmann, 1890)

Figs 12-13

Adeonella sparassis Ortmann, 1890: 54, pi. 4, fig. 10.

Adeonella hexangularis Okada, 1920: 630, text fig. 7, pi. 8, fig. 10.

Adeonella hexangularis - Okada 1934: 17, pi. 2, fig. 2. — Mawatari 1952: 285.

Adeonellopsis sparassis - Hirose 2010: 52, pi. 88, figs A-G.

Reptadeonella sp. - Hirose 2010: 53, pi. 90, figs A-D.

19

European Journal of Taxonomy 203: 1-41 (2016)

Material examined

Lectotype [designated herein]

JAPAN: colony, Sagami Bay, 1882, collected by L. Doderlein, MZS 1-2 (SEM specimen NSMT-Te745).

Paralectotypes [designated herein]

JAPAN: colony, Yokohama, 1891, collected by Rolle, MZS 1-1 (SEM specimen NSMT-Te747); colony,

Sagami Bay, 1882, collected by L. Doderlein, MZS 1-3 (SEM specimen NSMT-Te740).

Fig. 12. Adeonellopsis sparassis (Ortmann, 1890). A. Left, lectotype (MZS 1-2); right, paralectotype

(MZS 1-1) from among Ortm arm ’s (1890) syntypes in the Doderlein Collection. B. Colonies collected

at Tanegashima (NSMT-Te774). C. Autozooids (MZS 1-2) (NSMT-Te745). D. Gonozooids (MZS

1-2) (NSMT-Te745). E. Young, encrusting colony in Showa Emperor Collection (NSMT-BryR362).

F. Enlargement showing young autozooids with small spiramen (NSMT-BryR362). A-B = optical

photographs; C-F = SEM images. Scale bars: A-B = 2 cm; C = 150 pm; D-E = 200 pm; F = 100 pm.

20

HIROSE M., Japanese adeonid bryozoans

Other material examined

JAPAN: colonies collected from Suzaki, Izu, Sagami Sea, in the Emperor Showa collection at NSMT

(NSMT-BryR362); colonies collected from Arialce Sea (NSMT-Te770), from near Yakushima and

Tanegashima (NSMT-Te772, Te773, Te774); from off Shimoda (NSMT-Te890); from south ofNagannu

Island (NSMT-Te750, Te803); from Ogasawara (NSMT-Te771).

Measurements

Autozooids. ZL: 358-659 (527±64); ZW: 186-379 (263±39); n = 75. SOrL: 46-91 (63±8); SOrW:

72-118 (89±9);n = 53. SAvL: 106-174 (139±15); SAvW: 48-105 (75±13); n = 75. SpL: 29-76 (54±11);

SpW: 32-83 (52±11); n = 74.

Gonozooids. ZL: 623-855 (722±54); ZW: 445-630 (534±54); n = 24. SOrL: 52 -112 (77±14); SOrW:

136-265 (223±26); n = 21. SAvL: 137-218 (162±22); SAvW: 88-117 (104±8); n = 21. SpL: 97-185

(129±20); SpW: 99-267 (159±33); n = 23.

Young autozooids in ancestrular complex. ZL: 320-396 (361±32); ZW: 198-283 (231±31); n = 6.

SOrL: 67-83 (77±6); SOrW: 93-109 (100±7); n = 6. SAvL: 56-75 (67±8); SAvW: 28-55 (46± 12);

n = 6. SpL: 18-31 (23±5); SpW: 31-56 (41±9); n = 6.

Vicarious avicularia at branch margins. L: 253-476 (327±69); W: 154-228 (174±26); n = 8.

Description

Colony brownish yellow, dichotomously branching, irregularly spreading, many proximal branches

anastomosing, forming compact three-dimensional structure (Pig. 12A); anastomosed basal region can

exceed 4 cm high (3.67^4.64 cm). Branches flat, ribbon-lilce, variable in width (Pig. 12B); 1.6-8.6 mm

wide (average 4.0 mm; n = 101); much wider basally. Autozooids on both sides of branch, oval or

hexagonal, outlined by deep marginal groove (Pigs 12C, 13B); younger zooids at end of branch shorter

(average 382 pm; n = 6) and broader (average 324 pm; n = 9) than older zooids in anastomosed basal

region. Prontal shield minutely granulate, with numerous marginal pores and small ascopore (Pigs 12C,

13B). Spiramen almost circular in younger zooids, divided into 3-5 denticulate pores (Pig. 13B, E);

slightly broader in older zooids in basal part of colony (average 66 pm; n = 12). Peristome transversely

oval, roughly semicircular in outline, not tubular; primary orifice submersed in peristome. Triangular

suboral avicularium in center of frontal shield, between spiramen and orifice, directed distally (Pigs 12P,

13B). Rarely autozooids have an additional, smaller frontal avicularium, budded from frontal areolae

near proximal margin of zooid, directed distally (Pig. 12C). Gonozooids (Pigs 12D, 13C, F) occur

at branch bifurcations (Fig. 13C, F); larger than autozooids, hexagonal; with single or double rows

of marginal pores (Fig. 12D); slightly crescentic orifice compressed, much wider than long; spiramen

large, rounded-triangular or oval in outline, divided into 6-10 small, denticulate pores (Figs 12D,

13C). Gonozooids in basal part of colony much broader (average 602 pm; n = 6), and with a slightly

larger spiramen (average 142 pm long by 199 pm wide; n = 5). Vicarious avicularia (Fig. 13A) often

along edges of branches; triangular, directed distally; about same length as autozooids; rostrum length

about half of autozooid length. One young colony observed (Fig. 13D); encrusting, circular, 1.2 cm

in diameter; ancestrular complex consisting of six zooids radiating from center. Autozooids in young,

encrusting colonies (Figs 12E-F, 13D) oval, surrounded by distinct shallow groove; frontal shield with

few marginal pores and 4-6 knobs (Fig. 12F); secondary orifice broadly semicircular; spiramen circular,

divided into 2-4 small, denticulate pores; triangular suboral avicularium much smaller than that of

mature autozooids; between spiramen and orifice, directed distally, often partly overlapping proximal

orificial margin (Fig. 12F).

21

European Journal of Taxonomy 203: 1-41 (2016)

Remarks

Ortmann’s specimens comprise only the basal, anastomosed parts of colonies (Fig. 8A), and zooidal

characters in these specimens are identical to those in the basal parts of the more-complete Adeonellopsis

sparassis colonies that I studied. Adeonellopsis sparassis resembles A. sulcata (Milne Edwards, 1836)

in having a multiporous spiramen and small, distally directed suboral avicularia, but differs from

the latter in having a smaller, circular spiramen. MacGillivray’s (1880: plate 48, fig. 7) illustration

of Eschara mucronata (MacGillivray, 1868) (= A. sulcata ) appears to represent the basal part of a

Fig. 13. Adeonellopsis sparassis (Ortmann, 1890). A. Autozooids and marginal vicarious avicularia

(NSMT-Te772). B. Enlargement of autozooids (NSMT-Te772). C. Autozooids and gonozooids (NSMT-

Te774). D. Young, encrusting colony (NSMT-Te770). E. Young autozooids, showing circular spiramen

and large areolar pores (NSMT-Te774). F. Young autozooids and gonozooids at branch bifurcation

(NSMT-Te774). SEM images. Scale bars: A, C = 300 pm; B = 150 pm; D-E = 200 pm; F = 300 pm.

22

HIROSE M., Japanese adeonid bryozoans

colony of Adeonellopsis that resembles A. sparassis. However, as I did not examine MacGillivray’s type

specimen, I do not include it in the synonymy of A. sparassis.

I identified an encrusting colony in the Emperor Showa collection as a young colony of A. sparassis

based on the small, round multiporous spiramen and the triangular suboral avicularium between the

ascopore and orifice. Hayward & McKinney (2002) described a similar ancestrular complex consisting

of six zooids in the closely related encrusting genus Reptadeonella. Bock & Cook (2000) also reported

a similar encrusting ancestrular complex in Adeonellopsis , but they described a remarkably different

bipolar ancestrular colony in the articulated genus Adeona. I conclude the ancestrular complex in the

Emperor Showa collection to be A. sparassis rather than Reptadeonella or the other genera, based on the

orifice shape and the multiporous spiramen. The knobs on the frontal shield of zooids in the encrusting

colony appear to be a character common to young zooids in several adeonid species.

Okada (1920) described another adeonid bryozoan, Adeonella hexangularis Okada, 1920, from

Misaki (Sagami Bay) and the Kagoshima Gulf (southern Kyushu), which surely is Adeonellopsis

according to his description of a multiporous spiramen. Due to the limited original description and

apparent loss of the type specimens, the identity of A hexangularis is unclear, but Okada’s description

of an anastomosed colony morphology, frontal shields with a single or double rows of marginal

pores, and gonozooids located at branch bifurcation and having a broad, proximodistally compressed

orifice indicate that it is conspecific with A. sparassis and constitutes a junior synonym of the latter.

Mawatari (1952) reported nominal A. hexangularis from off Wakayama and Minabe, Kii Peninsula,

without a description; according to this record, A. sparassis is probably also distributed around the

Kii Peninsula.

Distribution

Japan: detected in this study from Sagami Bay, Tokyo Bay (Yokohama), Sagami Sea (Izu Peninsula),

Ariake Sea, Nansei Islands (near Yakushima and west of Okinawa), and Ogasawara, at depths of 24-

141 m. It was collected from eastern Sagami Bay and western Sagami Sea by Doderlein and Emperor

Showa, but was not found recently in those areas by NSMT.

Adeonellopsisparvirostrum sp. nov.

urn:lsid:zoobank.org:act:E46E9CDl-84F7-41DC-9AE9-9164B5262E5E

Figs 10B, 14-15

Adeona n. sp. - Hirose 2010: 50, pi. 86, figs A-D.

Diagnosis

Colony dichotomously branching; branches flat, roughly 3-4 mm wide. Autozooids oval, distinct,

delineated by a deep groove; with a single row of marginal pores. Frontal shield smooth or slightly

granulated. Peristome deep, oval. Spiramen monoporous. Two to six tiny pores in frontal wall proximal

to orifice. Suboral avicularia triangular, small, almost same size as spiramen; in center of frontal shield,

distal to spiramen, directed distally. Triangular marginal vicarious avicularia present. Gonozooids

slightly broader than autozooids; orifice broad, proximodistally compressed, with slightly convex

proximal margin; spiramen and avicularia almost same size as in autozooids.

Etymology

The specific name derives from the Latin parvus (small) and rostrum (used as a noun in apposition),

referring to the small rostrum of the suboral avicularia.

23

European Journal of Taxonomy 203: 1-41 (2016)

Material examined

Holotype

JAPAN: two branches of single colony collected by NSMT, north-northwest of Hachijo-jima Island,

33°20'54.48" N, 139°41'11.16 M E to 33°21'4.32" N, 139°40'30.84" E, 185-213 m depth, with CB dredge

from R/V Tansei-maru , 26 Nov. 2007 (NSMT-TeS25).

Paratypes

JAPAN: branch of colony collected by NSMT, same collection data as for holotype (NSMT-TeS27); four

dried colony fragments collected by NSMT, NE off Hachijo-jima Island, 33°34'6.00" N, 140°15'54.00" E

to 33°34'24.00" N, 140°16'0.00" E, 179-182 m depth, from R/V Shinyo-maru , 23 Oct. 2003 (NSMT-

TeS26); small branches of colony collected by NSMT, N of Hachijo-jima Island, 33°26'48.00" N,

139°42'42.00" E to 33°27'0.00" N, 139°42'24.00" E, 200-211 m depth, from R/V Shinyo-maru , 21 Oct.

2003 (NSMT-TeS28); several branches of colony collected by NSMT, southwest of Boso Peninsula,

~34°51' N, 139°40' E, 135-172 m depth, from R/V Shinyo-maru , 25 Oct. 2002 (NSMT-TeS29); several

branches of colony, Oshima Shin-sone, 28°52'24.60" N, 129°33'5.40" E to 28°53'1.20" N, 129°33'5.40" E,

169-172 m depth, with biological dredge from T/V Toyoshio-maru , collected 24 May 2011 (NSMT-

Te780); several branches of colony, Oshima Shin-sone, 28°52.5H' N, 129°33.233' E to 28°52.631' N,

Fig. 14. Adeonellopsis parvirostrum sp. nov. A. Colony branches from Oshima Shin-sone, N of

Amami Oshima (NSMT-Te780). B. Distal end of branch (NSMT-TeS26). C. Young autozooids and

some gonozooids (four of them are indicated with asterisks) at distal end of branch (NSMT-TeS26).

D. Gonozooids with broad, curved orifice, small triangular avicularia, and small spiramen (NSMT-

TeS26). A-B = photomicrographs; C-D = SEM images. Scale bars: A = 1 cm; B-D = 500 pm.

24

HIROSE M., Japanese adeonid bryozoans

129°33.313' E, 165-200 m depth, with biological dredge from T/V Toyoshio-maru , collected 29 May

2012 (NSMT-Te807).

Measurements

Autozooids. ZL: 593-865 (743±89); ZW: 283-366 (323±26); n = 21. SOrL: 54-112 (81±14); SOrW:

84-114 (102±8); n = 25. SAvL: 40-82 (58±10); SAvW: 21-46 (34±6); n = 51. SpL: 23-45 (33±6);

SpW: 23-40 (33±4);n = 38.

Gonozooids. ZL: 566-865 (659±91); ZW: 382-463 (420±19); n = 19. SOrL: 60-104 (78±15); SOrW:

118-132 (123±5);n= 11.

Vicarious avicularia at branch margins. L: 189-224 (209±14); W: 111-125 (118±6); n = 7.

Additional small frontal avicularia in middle and basal parts of branch. L: 99-114 (106±6); W: 46-63

(55±5); n = 10.

Description

Colony yellowish white, dichotomously branching. Branches flat, slender, 2.607M.433 mm wide

(average 3.3 mm; n = 10); (Fig. 14A). Autozooids on both sides of branches. Autozooids oval, distinct,

delineated by deep groove (Fig. 14B-C). Frontal shield umbonuloid (Fig. 10B). Frontal shield smooth or

Fig. 15. Adeonellopsis parvirostrum sp. nov. A. Middle of branch, showing autozooids lacking suboral

avicularium (NSMT-Te780). B. Kenozooids in old part of branch (NSMT-Te780). C. Autozooids and

marginal vicarious avicularia at periphery of branch (NSMT-Te780). D. Lateral view of vicarious

avicularium at branch bifurcation (NSMT-Te780). SEM images. Scale bars: A, C-D = 200 pm; B =

300 pm.

25

European Journal of Taxonomy 203: 1-41 (2016)

with minute granules or wrinkles; with single row of marginal pores. Peristome deep, secondary orifice

transversely oval. Spiramen in center of frontal shield; monoporous, circular (Figs 14D, 15A), rarely

divided into two or three non-denticulate pores. Two to six tiny, irregular frontal pores proximal to orifice;

roughly same size as marginal pores (Figs 14C-D, 15A). Small suboral avicularium (Figs 14D, 15A, C)

just distal to and often abutting spiramen; rostrum short, with triangular mandible; separated from orifice

by distance up to several times greater than mandible length; directed distally. Additional small frontal

avicularia lacking in younger part of branch (Fig. 14C). Vicarious avicularia present (Fig. 15C-D);

marginal, almost three times length of suboral avicularia; with triangular mandible. In middle and basal

parts of colony, autozooids somewhat irregular in shape; borders indistinct; 572-774 pm long (average

665 pm) by 238-328 pm wide (average 274 pm) (n = 12); frontal shields heavily secondarily calcified;

orifice nearly circular, 76-93 pm long (average 85 pm) by 73-115 pm wide (average 90 pm) (n = 8). In

basal autozooids, spiramen consists of small single pore, 26-33 pm (average 30 pm) long by 23-31 pm

wide (average 27 pm) (n = 4); suboral avicularia slightly larger than in non-basal zooids, 107-122 pm

long (average 112 pm) by 41-47 pm wide (average 44 pm) (n = 5); spiramen and suboral avicularia

submerged in common depression in heavily calcified frontal shield. Autozooids in middle and basal parts

of branch bear additional small frontal avicularium, often on proximal margin of zooid (Fig. 15A, C);

directed proximally or proximolaterally. Basal part of colony consists of irregularly shaped lcenozooids

that bear several small frontal avicularia (Fig. 15B). Gonozooids (Fig. 14C-D) slightly broader than

autozooids; orifice crescentic, much wider than long, with convex proximal margin; spiramina and

suboral avicularia roughly same size as in autozooids.

Remarks

Adeonellopsis parvirostrum sp. nov. fits the original description of Adeonellopsis japonica in having a

small, circular spiramen and small suboral avicularium, but its avicularium is much smaller than in the

latter. Young autozooids of A. parvirostrum sp. nov. lack the striation perpendicular to the margin seen

on the rim of young zooids of A. japonica. Adeonellopsis parvirostrum sp. nov. resembles A. lichenoides

in the basal part of the colony (compare Fig. 5D with Fig. 15B), but differs from the latter in the

presence of a distally directed suboral avicularium between the orifice and spiramen in older parts of

branches. Some small frontal avicularia are sparsely surrounded by the remains of associated areolae of

original autozooids and/or kenozooids (Fig. 15A-C). Some of the specimens of A. parvirostrum sp. nov.

I observed were pinkish, although this may have resulted from artificial staining of another organism, as

other bryozoan colonies in the same bottle were also pinkish.

Distribution

Japanese Pacific waters: north of Hachijo-jima Island, Sagami Sea, and near Amami Oshima, at depths

of 135-213 m.

Adeonellopsis toyoshioae sp. nov.

urn:lsid:zoobank.org:act:B7DA81C8-D492-4AEA-A953-CB19EDF8EF45

Fig. 16

Adeonellopsis subteres - Hayami 1971: 87, pi. 11, fig. 4.

Diagnosis

Colony dichotomously branching; branches flat, or nearly cylindrical, slender, roughly 0.7-1.4 mm

wide. Autozooids oval, distinct, delineated by deep groove; with single row of small marginal pores.

Frontal shield entirely with granules and wrinkles, forming thickened ridge around spiramen. Peristome

raised, transversely broad. Spiramen large, equal to or greater than secondary orifice in size, variable in

26

HIROSE M., Japanese adeonid bryozoans

shape, divided into 4-8 stellate pores. Suboral avicularia triangular, large, distal to spiramen; directed

distally or distolaterally, rostrum tip extending over proximal margin of orifice. Gonozooids unknown.

Etymology

The specific name refers to the T/V Toyoshio-maru of Hiroshima University.

Fig. 16 . Adeonellopsis toyoshioae sp. nov. A. Colonies from Amami Oshima (NSMT-Te776). B. Branch

showing autozooids with broad multiporous spiramen (NSMT-Te776). C. Branch showing crenulate

periphery, with tubular peristomes and acute avicularia (NSMT-Te776). D. Branch bifurcation lacking

vicarious avicularia (NSMT-Te776). E. Distal end of branch showing young autozooids with large

multiporous spiramen (NSMT-Te776). F. Old autozooids with immersed spiramen (NSMT-Te776). A =

optical photograph; B-F = SEM images. Scale bars: A = 5 mm; B, E = 300 pm; C, F = 500 pm; D =

200 pm.

27

European Journal of Taxonomy 203: 1-41 (2016)

Material examined

Holotype

JAPAN: small colony collected at Takatori, southwest of Otsuki, Kochi Prefecture, ~32°30' N, 132°50' E,

by coral fishing net, 118-120 m depth, F/V Yuryo-maru, 29 Sep. 2010 (NSMT-Te777).

Paratypes

JAPAN: small colony, same collection data as for holotype (NSMT-Te779); small branches of colony,

north of Amami Oshima, 28°52'38.40" N, 129°33'13.80" E, 178 m depth, by biological dredge,

T/V Toyoshio-maru , 24 May 2009 (NSMT-Te776); several branches of colony, collected at Oshima

Shin-sone, 28°52 , 24.60" N, 129°33 , 5.40" E to 28°53T.20" N, 129°33'5.40" E, 169-172 m depth, by

biological dredge, T/V Toyoshio-maru , 24 May 2011 (NSMT-Te778); colony collected by NSMT, north

of Hachijo-jima Island, 33°26'48.00" N, 139°42'42.00" E to 33°27'0.00" N, 139°42'24.00" E, 170—

176 m depth, RV Shinyo-maru, 21 Oct. 2003 (NSMT-TeS23); colony collected by NSMT, north of

Hachijo-jima Island, 33°25'60.00" N, 139 o 41'54.00" E to 33°26’6.00” N, 139 o 4E36.00” E, 160-190 m

depth, RV Shinyo-maru , 21 Oct. 2003 (NSMT-TeS24); several branches of colony, Oshima Shin-sone,

28 0 52.51P N, 129°33.233' E to 28°52.63P N, 129°33.313' E, 165-200 m depth, by biological dredge,

T/V Toyoshio-maru , 29 May 2012 (NSMT-Te808); several branches of colony, north of Tanegashima

Island, 30°53.317' N, 131°02.565' E to 30°53.336' N, 131°02.747' E, 165-200 m depth, by biological

dredge, T/V Toyoshio-maru , 30 May 2012 (NSMT-Te802).

Measurements

Autozooids. ZL: 437-624 (530±43); ZW: 224-312 (255±20); n = 37. SOrL: 36-76 (56±8); SOrW:

63-102 (75±10);n = 28. SAvL: 91-174 (136±19); SAvW: 52-97 (72±9);n = 56. SpL: 62-262 (114±46);

SpW: 35-116 (72±16);n = 61.

Additional small frontal avicularia. L: 83-119 (97±10); W: 37-62 (48±6); n = 13.

Description

Colony yellowish white, dichotomously branching, small, up to 3 cm high. Branches flat or nearly

cylindrical, slender (Fig. 16A); 0.7-1.4 mm wide (average 1.0 mm, n = 15). Autozooids all around

branches; oval, distinct, delineated by deep groove, with single row of small marginal pores (Fig. 16B).

Frontal shield raised from margin, entirely textured with small granules and wrinkles, forming a thickened

ridge around spiramen; ridge often bears up to four low, smooth umbos 40-78 pm in diameter (average

60 pm, n = 52) (Fig. 16D). Peristome raised, thick-walled distally and laterally, transversely broad;

secondary orifice semicircular to transversely oval (Fig. 16B, D). Primary orifice hidden in peristome

in frontal view. Spiramen large, size roughly equal to or greater than area of secondary orifice; variable

in shape, divided into 4-8 stellate pores (Fig. 16D-E). Single large, triangular suboral avicularium

between orifice and spiramen, directed distally or distolaterally (Fig. 16B, E); rostrum acute, tip often

extending over proximal margin of orifice (Fig. 16D); mandible elongate-triangular; opesial opening

oval; crossbar lacking. Some zooids have additional, smaller frontal avicularium budded, near proximal

margin of zooid, pointing in any direction. Marginal vicarious avicularia triangular, directed distally;

312 pm long by 134 pm wide (n =1). Gonozooids unknown.

Remarks

Adeonellopsis toyoshioae sp. nov. resembles A. subteres (Romer, 1863) in the number of alternate zooidal

rows, raised margin of the frontal shield, and a large ascopore occupying a broad area of the frontal shield

(Romer, 1863: pi. XXXV, fig. 6). However, A. subteres was originally reported as a fossil from Germany

and occurs in the Eocene (Canu 1907; Braga 1963) and Oligocene (David & Pouyet 1968) deposits of

Europe. I therefore concluded that A. toyoshioae sp. nov. represents a new species. Moreover, the Recent

28

HIROSE M., Japanese adeonid bryozoans

Adeonellopsis distoma (Busk, 1858) from the eastern N Atlantic is closely related to both species;

Adeonellopsis distoma resembles A. toyoshioae sp. nov. in having a subtubular peristome and a large

area of multiporous spiramen consisting of about seven pores. Adeonellopsis distoma , however, differs

from A. toyoshioae sp. nov. in having an elongated spiraminal area, with usually two rows of pores, in

having larger suboral avicularia and fewer zooidal rows comprising the branches, and in lacking the

marginal umbos. Adeonellopsis toyoshioae sp. nov. also resembles A. sparassis in having a multiporous

ascopore with more than four denticulate pores, and a single suboral avicularium extending over the

proximal margin of the orifice. However, the ascopore in A. toyoshioae sp. nov. is not circular as in

A. sparassis and occupies a much larger area of the frontal shield. In addition, A. toyoshioae sp. nov. has

branches less than 2.0 mm wide, whereas those in A. sparassis are usually wider than 2.5 mm. Finally,

the suboral avicularium is often directed slightly distolaterally in A. toyoshioae sp. nov., whereas it

points directly distally A. sparassis.

The marginal umbos evident on younger zooids in A. toyoshioae sp. nov. are identical in size to those in

A. pentapora ; these umbos are also a characteristic feature of younger zooids in these species.

Distribution

Japanese Pacific waters: north of Hachijo-jima Island, southwest of Kochi Prefecture, and the northern

Nansei Islands (near Amami Oshima and Tanegashima); at depths of 118-200 m. Hay ami (1971)

reported this species from fossiliferous sandstone in the Pliocene Shinzato Tuff (Shinzato Formation)

on Okinawa Island, which is close to the Recent distribution in Japan.

Taxonomic key to Japanese adeonid species

1. Orifice circular; single peristomial spiramen proximal to orifice; no avicularia between orifice and

spiramen; frontal shield with many small pores and several avicularia.2.. .Adeonella Busk, 1884

- Orifice oval; spiramen single or multiporous; single or paired avicularia between orifice and spiramen;

frontal shield normally bearing only marginal pores.3 ...Adeonellopsis MacGillivray, 1886

2. Branch width reaches more than 3 mm; generally large vicarious avicularia present at branch

bifurcations; generally more than two avicularia on frontal shield; areolar pores nearly uniform

and circular; F/W ratio of frontal avicularia generally less than 2.0.

. Adeonella cf. lichenoides (Famarck, 1816)

- Branch width less than 2.5 mm; often lacking a large vicarious avicularium at branch bifurcations;

generally single frontal avicularium on frontal shield; areolar pores irregular in shape and often slit-

lilce; F/W ratio of frontal avicularia generally greater than 2.0. Adeonella jahanai sp. nov.

3. Autozooidal spiramen small, 1-2 openings; openings not denticulate .4

- Autozooidal spiramen large, multiporous with more than four openings; openings denticulate.6

4. Branches less than 2.0 mm wide; suboral avicularium abutting orifice, separate from spiramen,

slightly curved, directed distolaterally. Adeonellopsis arculifera (Canu & Bassler, 1929)

- Branches greater than 3.0 mm wide; suboral avicularium abutting spiramen, rostrum straight, directed

laterally or distolaterally.5

5. Suboral avicularium large, rostral width equal to or greater than orifice width; rostrum extending

from spiramen to orificial margin. Adeonellopsis japonica (Ortmann, 1890)

- Suboral avicularium small, rostrum narrower than orifice; avicularium distant from orifice

. Adeonellopsis parvirostrum sp. nov.

6. Suboral avicularia small, single or paired. Adeonellopsis pentapora Canu & Bassler, 1929

- Suboral avicularium large, single.7

29

European Journal of Taxonomy 203: 1-41 (2016)

7. Branches usually greater than 2.5 mm wide; suboral avicularium directed distally; peristome

not tubular; autozooidal spiramen width less than orifice width; spiramen circular in marginal

autozooids . Adeonellopsis sparassis (Ortmann, 1890)

- Branches usually less 2.0 mm wide; suboral avicularia directed distolaterally; peristome tubular;

autozooidal spiramen width equal to or greater than orifice width; spiramen oval or irregular in marginal

autozooids . Adeonellopsis toyoshioae sp. nov.

Discussion

This study detected eight adeonid bryozoan species in Japanese waters, including a new record for

Japan and three new species. It is the first review of Japanese adeonid bryozoans since Ortmann (1890)

published the first monographic treatment of Japanese bryozoans. In a taxonomic key to Japanese

cheilostome bryozoans published in Japanese, Okada (1918) established another adeonid bryozoan

species, Adeonella vulgaris Okada, 1918, which I have not treated herein; the only description was

that it has flat, antler-like branches with autozooids on both sides. Okada (1934) later provided a more

detailed description based on material from Shimoda (western Sagami Sea); Adeonella vulgaris is surely

not an adeonid because Okada (1934) mentions a globose ovicell perforated by a longitudinal slit. This

species belongs in some other group that includes erect bryozoans, such as Cigclisula Canu & Bassler,

1927 in the family Colatooeciidae Winston, 2005.

The six species of Adeonellopsis described herein show clear-cut differences in the size and shape of

the spiramen and suboral avicularium (Fig. 17). The direction of the suboral avicularium and character

of denticulation in the spiramen have been regarded as important diagnostic characters for species in

this genus. Although they are often immersed in old zooids, the spiramen and suboral avicularia seem

to be strongly constrained in size and shape from early ontogeny, and their morphology provides good

diagnostic characters throughout development.

Further studies on adeonid taxonomy are needed. Adeonid larvae settle rapidly, after only a few hours in

the water column, and are unlikely to disperse broadly. However, some adeonid species are supposedly

broadly distributed, having been reported across several oceans: A. lichenoides , for example, is cited to

occur from East Africa to the eastern Pacific and from southern Australia to Japan. It will be informative

to examine these broadly distributed species using molecular data, to determine whether they actually

represent species complexes that include species previously undetected through morphology, and to

further clarify their taxonomy, ecology, and biogeography.

Fig. 17. Schematic drawings of autozooids for the six species of Adeonellopsis MacGillivray, 1886 in

Japan. A. A. arculifera (Canu & Bassler, 1929). B. A. japonica (Ortmann, 1890). C. A. parvirostrum

sp. nov. D. A. pentapora Canu & Bassler, 1929. E. A. sparassis (Ortmann, 1890). F. A. toyoshioae

sp. nov.

30

HIROSE M., Japanese adeonid bryozoans

Adeonellopsis pentapora is also thought to be widely distributed, occurring in the eastern Pacific

from Tasmania to Japan. Mature colonies of A. pentapora are small and often occur as epibionts on

substrata such as hydroid stems and gastropod shells inhabited by living hermit crabs. Some hydroids

can potentially inhabit soft bottoms, providing substrate for other organisms, and hermit crabs are at

least more mobile than sessile bryozoan colonies, which might explain^, pentapora having the broadest

distribution in Japan among the adeonids treated in this study (Fig. 18). Adeonellopsis japonica also

appears to be widely distributed, occurring along the Pacific coast of Japan and having been reported

from Hawaii, more than 6000 km distant; I did not detect any morphological differences between

Japanese and Hawaiian specimens, but genetic studies are needed to clarify whether these populations

are cryptic or invasive species.

In Japan, adeonid bryozoans have not been found north of Tsugaru Strait, lying between Hokkaido and

Honshu (Fig. 18). This pattern may be correlated with the warm Kuroshio Current and its branches

(the Tsushima and Tsugaru Currents), and limited in the north by the cold Oyashio Current (Fig. 1).

The lack of records from western Hokkaido despite the extension of the Tsushima current might be

due to decreasing effects of the current and colder temperatures, but might also be due to a general lack

of collecting in this area, especially in deeper water. Although the Pacific coast of Tohoku (northern

Honshu) is influenced by the Oyashio Current, two species (A. japonica and A. pentapora) occur at

Otsuchi; this is presumably due to the effects of the Kuroshio and Tsugaru currents, which normally

affect nearshore areas more than does the Oyashio Current.

A.

japonica

£

f

z

P> Jf

/

§

Z

1

0

Eo

co

Z

/

4

1

o

ro

A

toyoshic

-ae—

•

P

1

d

r

fj?

*

r

9

i

130" E 135’E 140’E 145’E 130" E 135" E 140" E 145" E

Fig. 18. Maps showing the distribution of the eight adeonid species in Japanese waters. Solid circles

indicate sites where a species was detected in this study and unfilled circles indicate records from

Canu & Bassler (1929) and Sakakura (1935) for A. pentapora Canu & Bassler, 1929, Okada (1920)

and Mawatari (1952) for A. sparassis Ortmann, 1890, and Okada & Mawatari (1935) for A. japonica

Ortmann, 1890.

31

European Journal of Taxonomy 203: 1-41 (2016)

Adeonellopsis pentapora shows the broadest distribution in Japan, from south of Yakushima to Otsuchi

(this study) and Tsugaru Strait (Canu & Bassler 1929); the paucity of records of A. pentapora from the

Sea of Japan is probably due to low sampling effort in this area. Most other adeonid species are distributed

in southern Japan, from central Honshu to some point in the Nansei Islands. Adeonella jahanai sp. nov.

and Adeonellopsis arculifera , detected only from the Nansei Islands, show the narrowest distributions

(Fig. 18).

In the present study, Adeonellopsis toyoshioae sp. nov. was collected from both temperate and subtropical

habitats, in areas strongly influenced by the Kuroshio Current. Hay ami (1971) described a typically Indo-

Pacific bryozoan fauna from the Miocene to Pliocene Shinzato Tuff in Okinawa, where she reported

fossil colonies of A. toyoshioae sp. nov., indicating a temperate to subtropical paleoenvironment in this

area at the time.

Adeonellopsis japonica is distributed from Kouchi to Otsuchi on the Pacific side of Shikoku and Honshu,

but unlike the other species is conspicuously absent from Kyushu and the Nansei Islands (Fig. 18).

Although A. japonica has also been reported from Hawaii (Cheetham et al. 1980: USNM 271601, IZ cat

8450; RB 8450), the records there are from deep water (549-605 m). This species thus appears to be

distributed in cooler water than the other adeonid species.

Acknowledgments

I thank Mrs Marie-Dominique Wandhammer and Mrs Marie Meister (Musee Zoologique Strasbourg) for

assistance in observations, the loan of the Doderlein Collection, and other help; and Dr Hiroshi Namikawa

(Showa Memorial Institute of the National Museum of Nature and Science) for valuable assistance

and the loan of the bryozoan collection of Emperor Showa; Dr Joac him Scholz and Mrs Brigitte Totz

(Senckenberg Forsuchung Institute) for assistance during my stay in Germany; Ms. Jo Ann Sanner and

Ms. Kathy A. Hollis (Smithsonian Institution, the National Museum of Natural History) for assistance

in observation of the USNM bryozoan collection during my stay in Washington; Dr Shigeyuki Yamato

(Seto Marine Biological Taboratory, Kyoto University) and Dr Shinta Fujimoto (Kyoto University) for

assistance in observing specimens at SMBL studied by Okada and Mawatari; Prof. Susumu Ohtsuka and

the crew of T/V Toyoshio-maru (Hiroshima University) for crucial support in field collecting; Dr Asako

Matsumoto (Reitaku University) and Mr Masataka Kurosawa, Mr Kouichi Morita, and Mr Masaaki

Hirano (International Coastal Research Center, The University of Tokyo) for support in collecting in

Otsuchi; Mr Yoshihiko Niiya for collecting and providing bryozoan specimens from Kochi Prefecture;

Mr Hisanori Kohtsuka (Misaki Marine Biological Station, The University of Tokyo) for support in

collecting in Sagami Bay; Dr Yoshiaki Kai (Maizuru Fisheries Research Station, Kyoto University) for

support in collecting in Wakasa Bay; Mr Yoshikazu Jahana for collecting bryozoan specimens around

Okinawa; Takuo Higashiji and Hiroko Takaoka (Okinawa Churaumi Aquarium) for providing specimens

from Okinawa; Professor Shunsuke Mawatari (Hokkaido University) for much crucial support of this

study; Dr Matthew Dick (Hokkaido University) for reviewing and editing drafts of the manuscript;

and Dr Bjorn Beming and an anonymous reviewer for highly constructive comments and suggestions

during peer review. This study was supported in part by Grants-in-Aid from the Japan Society for

the Promotion of Science (Nos 20-3856 and 23-3410), a Grant-in-Aid from the Kuroshio Biological

Research Foundation in 2010, and the research program Tohoku Ecosystem-Associated Marine Science

(TEAMS) subsidized by the Ministry of Education, Culture, Sports, Science and Technology (MEXT),

Japan.

32

HIROSE M., Japanese adeonid bryozoans

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Manuscript received: 2 September 2015

Manuscript accepted: 15 February 2016

Published on: 8 June 2016

Topic editor: Rudy Jocque

Desk editor: Kristiaan Hoedemakers

Printed versions of all papers are also deposited in the libraries of the institutes that are members of the

EJT consortium: Museum national d’Histoire naturelle, Paris, France; Botanic Garden Meise, Belgium;

Royal Museum for Central Africa, Tervuren, Belgium; Natural History Museum, London, United

Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Natural History Museum of

Denmark, Copenhagen, De nm ark

36

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