New Collections of Ascidians from the Western Pacific and Southeastern

Home , Biaro Island, Didemnidae, Exostoma, Leptoclinides, Ring of Red, Sigillina

habitats that were investigated were the reef flat accessible on foot, and bottoms bottoms and foot, accessible on reef flat the were investigated were that habitats

Siboga Siboga Naval Officers interested in natural history and the specimens were deposited in in deposited were specimens the and history natural in Officers interested Naval

large European Museums. Museums. European large

limited areas. The first descriptions of ascidians from this part of the world (Heller (Heller of world the this part from of ascidians first The descriptions areas. limited

Pacific. Many of the described species were the result of sporadic collections from collections species sporadic of described result the of were the Pacific. Many

regions. regions.

pattern is likely for ascidians, as they are most abundant on coral reefs reefs tropical in coral on abundant most are they as is likely ascidians, for pattern

from the western Pacific adjacent to the Inda-Malayan region which which is region considered Inda-Malayan the to Pacific western the adjacent from

to be the center of their evolution, to the central Pacific (Wells (Wells Pacific central the to evolution, their of center the be to

tebrates

acters obviously favorable to the development of a varied fauna. For all inver all For fauna. varied a of development the to favorable obviously acters

Studies of scleractinian corals, for example, have shown a decreasing diversity diversity decreasing a shown have example, for corals, scleractinian of Studies

content of nutrients carried in sea water from nearby land masses, are all char all are masses, land nearby from water sea in carried nutrients of content

igins of the present day sea floor, the rather uniform temperature, and the high high the and temperature, uniform rather the floor, sea day present the of igins

1878, 1878, numerous islands and reefs, and an extremely diverse coast-line. The diverse or diverse The coast-line. diverse extremely an and reefs, and islands numerous

Micronesica Micronesica

Most of our knowledge of ascidians of this region is due to the the to due is region this of ascidians of knowledge our of Most

Previously, ascidians had not been systematically collected in the western western the in collected systematically been not had ascidians Previously,

The tropical western Pacific Ocean has a large area of shallow waters with with waters shallow of area large a has Ocean Pacific western tropical The

provinces in this large region. region. large this in provinces

general statements, and presently it is not possible to delimit faunistic faunistic delimit to possible not is it presently and statements, general

indicated for each species, but the sampling is not sufficient to allow allow to sufficient not is sampling the but species, each for indicated

abundance of the Didemnidae family. The geographical distribution is is distribution geographical The family. Didemnidae the of abundance

species, species, a with peculiar 34 of which forms new. predominate are Colonial

mens from Palau, Chuuk, the Philippines, Indonesia, and Papua New New Papua and Indonesia, Philippines, the Chuuk, Palau, from mens

Guinea. A large part of them were collected by the Coral Reef Research Research Reef Coral were the by them collected of large part A Guinea.

Abstract-Recent Abstract-Recent Foundation, based in Palau. Descriptions and figures are given for for given are figures and Descriptions Palau. in based Foundation,

pce nw fo teId-et aii ein eeaie speci examined We region. Pacific Inda-West the from species known

and southeast Asian islands have significantly increased the number of of number the increased significantly have islands Asian southeast and

von Drasche Drasche von

Expedition and the works of Sluiter Sluiter of works the and Expedition

, ,

the western Pacific is considered a major source of taxonomic diversity. diversity. taxonomic of source major a Pacific is considered western the

29(2): 133 279, 1996 1996 279, 133 29(2):

New Collections Collections New of Ascidians from the Western Pacific

Biologie des des Biologie

1884

FRANl;OISE MONNIOT AND CLAUDE MONNIOT MONNIOT CLAUDE AND MONNIOT FRANl;OISE

CNRS CNRS

lnvertebres lnvertebres

collections of ascidians from several western Pacific Pacific western several from ascidians of collections

, ,

Traustedt Traustedt

D 0699 D

e-mail: monniot@cimrs/.11111/mfr monniot@cimrs/.11111/mfr e-mail:

and and Southeastern Asia

Marins, Marins,

, ,

55 55

Introduction Introduction

1885) 1885)

me me

M11se11m M11se11m

B11ffon

were made from specimens specimens by collected from were made

, ,

(1904, 1909). 1909). (1904,

75005 75005

National National

Paris

Histoire Histoire

, ,

France France

The only two kinds of of kinds two only The

Nall/re/le, Nall/re/le,

1988). 1988).

1899

The same same The

99 99

- 1900 1900

Alba

collections. collections. Many type speci

These These figures demonstrate how little

. .

(MNHN) (MNHN)

Expedition Expedition (Herdman 1882, 1886), the

Micronesica Micronesica 29(2), 1996

Challenger Challenger

The The present study includes collections made by the Coral Reef Reseach Foun

Nevertheless Nevertheless ascidians remain poorly known in the tropics. Besides the tech

Cruise Cruise (Van Name but 1918), they did not collect extensively. Later, samples

The The recent popularity of SCUBA diving and the presence of specialists in the

The The ascidian fauna is bettter known on each side of this wide tropical region.

Other Other oceanographic expeditions to the western Pacific from which ascidians

Monniot Monniot to 1987 1991, F. Monniot to 1987 1994).

. .

Japan, Japan, over a period of years, 50 Tokioka, and later, Nishikawa have described

and and the need for future continued work on this group.

new new species, and redescriptions of other poorly known species. This contribution

of of the shallow water fauna of the western Pacific by giving descriptions of many clearly clearly shows the great diversity of the western Pacific coastal fauna of ascidians

geographic geographic distributions can be given among the different archipelagos and only

very general very general conclusions can drawn. be The present work improves our knowledge

species species was preserved and no intraspecific variation can be addressed. Often, a

here here do not constitute a complete inventory. Very often only one specimen each of species previously previously species collected from an area was not recollected elsewhere. No precise

mens mens from several Museums observed were for comparisons; the described species

Museum Museum National d'Histoire Naturelle

leagues leagues at Scripps Institute of Oceanography, and specimens deposited in the

dation dation which is based in Palau, Caroline Islands, a few samples collected by col

quires quires fully mature specimens, fixation in formalin, dissection and staining before

nical nical difficulties of collecting, especially colonial animals, a taxonomic study re any any microscopic observation.

(Monniot (Monniot & Monniot 1987) and 202 species from southern New Caledonia

(1984) (1984) recorded 62 species, excluding Didemnidae, in Micronesia (Chuuk, Pohn

pei, pei, and Majuro). We have recorded about 100 species from French Polynesia

For For example Kott (1981) described 60 species on the Fiji reef flat and Nishikawa

600 600 species. field have have field increased the number of ascidians collected and identified considerably.

more more than 400 species. The ascidian fauna of Australia is estimated to be 500-

and and excluding the didemnids, which are not yet published, this fauna numbers

In In cidians cidians were extensively collected and described by Kott in 1985, and 1990 1992,

more more than 300 species. Around Australia, mostly on the Great Barrier Reef, as

is known is known about the western Pacific tunicate fauna.

only only about species from 100 the Philippines

fura fura sea (Tokioka 1952), Palau Islands (Tokioka 1950, 1955), and Philippines dicate dicate that about 250 ascidian species have been recorded from Indonesia, and

(Tokioka (Tokioka have 1970) also increased our knowledge. Other short publications in

tross tross sonian sonian Institution were studied by Tokioka (1967). Millar (1975) described some

from from the Zoological Museum in Copenhagen. Smaller collections from the Ara

from from sporadic collections were deposited in several Museums. Those of the Smith

134 134 were were collected were the

common common in virtually all shallow marine habitats which shows how limited sam

below below 40 m depth explored with dredges from a ship. Ascidians, however, are pling pling was in the early 20th century.

are visible as pink spots through a translucent tunic and are arranged in double double in arranged are and tunic translucent a through spots pink visible as are

Holotype

Aplidium multiplicatum: multiplicatum: Aplidium

Aplidium ca/ifornicum: ca/ifornicum: Aplidium

134°26.82' E, l m l 134°26.82' E,

Description: The colonies form cushions or balls, balls, 2 3 or The cushions colonies to form cm zooids Description: The across.

iue I Figure

: :

larva. larva.

MNHN: Al Apl B 312 B Apl Al MNHN:

. .

Aplid

1'';-'", 1'';-'",

. .

-..'. -..'.

um um

Aplidium controversum controversum Aplidium

F. Monniot 1987: 518, fig. 8 c-g, New Caledonia 1987: Caledonia 518, New c-g, 8 fig. Monniot F.

contro

Kott 1992a: 1992a: 567, fig. Australia 90, Kott

Monniot Monniot

l'e

Order Aplousobranchia Aplousobranchia Order

rsum rsum

Family Polyclinidae Polyclinidae Family

& &

Fig. 1, Pl. IA IA 1, Pl. Fig.

Monniot: New Ascidians Ascidians New Monniot:

. .

Palau, Koror State, marine lake, 7°17.81' N; N; 7°17.81' lake, marine State, Koror Palau,

.: .:

, ,

oi; B zooid;

new species species new

s ,

cin f th of ection

0,5 0,5

0.5 0.5

e e

mm mm

stomach

; ;

C C 135 135

A. A.

Aplidium Aplidium

Aplidium Aplidium cali

has not been located. has located. not been

stomach stomach has to 14 deep 15

is registered at the Zoologisch at the Zoologisch registered is

The The

A. A. californicum

sie1e. sie1e.

1911) and and 1911) Kott (1963) to synonymize

A. A. multiplicatum

have have been noted by ascidian taxonomists for a

Micronesica Micronesica 29(2), 1996

from from British Columbia the colonies form rounded cush

A. A. multiplicatum

Sluiter, 1909 by Kott by Kott Sluiter, the except which synonyms gave, she (1992a) 1909

and and

(Ritter, (Ritter, from New Caledonia 1900) by F. Monniot and (1987)

A. A. californicum

In In

The The external appearance and numerous similar anatomical characters in

Remarks: Remarks: This resembles species the specimens identified as

The zooids reach 15 mm in total length. The thorax, 2 mm long when relaxed, relaxed, when total long thorax, mm The in length. mm 2 reach zooids The 15

controversum. controversum.

. .

the the ovary rather close to the gut loop, followed by a regular double row of testis

and and cacea at 2 the beginning the of rectum. The post abdomen elongated, is with

folds, folds, some of them interrupted or irregular. There is an annular post-stomach

languets languets are strongly displaced to the left

branchial branchial sac has 8 to 11 rows of stigmata, usually 9 or The 10. dorsal rapheal

forms forms a short tube and has a triangular short languet on its dorsal rim. The

ions, ions, yellow or pale orange in color. The zooids are arranged in double rows zooids zooids have 6 oral lobes, tentacles 12 in 2 orders of size. The cloacal aperture

converging converging to common cloacal openings at the upper part of the colony. The

Only newly-collected specimens of of the specimens latter Only from newly-collected British species Columbia have A

been studied. We provide a brief redescription of each species for comparison comparison for with species each redescription of provide brief a We studied. been Museum of Amsterdam, of Amsterdam, the type Museum of however

the question. The type specimen of of the question. The specimen type

californicum californicum the the two however species, descriptions insufficient complicated have successively

long long time. This caused Hartmeyer (1909

successive confusion confusion authors of successive several about and this species its relatives.

have to be verified by reexamining the specimens. However, we consider the Palau Palau the consider we However, the specimens. reexamining by verified have to be

colonies are colonies different and the name represent the a reflecting species species, new

begin begin to differentiate.

multiplicatum multiplicatum

have have 3 flat adhesive papillae. On each side there is a row of about 30 round

Several larvae larvae Several are incubated the cavity. They inside measure cloaca! mm 0.9 and epidermal vesicles along along the epidermal anterior vesicles half of the perimeter of the trunk (Fig. IC).

lowed lowed by a double row of extending down testis follicles to the heart (Fig. IA). The The sensory organs are located posteriorly. In some larvae 2 rows of stigmata

post-abdomen post-abdomen very long. is The ovary distance some lies from the abdomen, fol

(Fig. (Fig. IA). The anus opens at two thirds of the cavity. way The down the cloaca}

ment, by ment, a narrow followed joining caeca tube the well-marked 2 rectum between

post-stomach post-stomach has an annular rod. The intestine with begins an olive-shaped seg

is plicated is plicated in 20 to 24 longitudinal folds (Fig. IB), of few them incomplete. The

fornicum fornicum

of small lateral swellings. The thoracic muscles are mostly longitudinal. longitudinal. mostly are The thoracic The bran muscles lateral swellings. small of displaced displaced to the left The side. abdomen is shorter than the thorax. The stomach

gated gated a in stout, short, pointed languet (Fig. IA). The latter has a sometimes pair chial chial has rows sac to of stigmata. 20 IO 15 The rapheal languets are slightly only

136 136 pigment The pigment cells. cloacal aperture forms a short, narrow tube, dorsally prolon

is is wide (Fig. IA). The oral siphon has 6 lobes; its basal part encircled by white

color color pale (Pl. is yellow IA). The is soft, consistency but there is no mucus. lines lines along cloacal channels that open at the top of side the colony. The general

The cloaca] aperture is widely opened (Figs. 2A, B, C), uncovering a large part part large a uncovering C), B, 2A, (Figs. opened widely is aperture cloaca] The

of the branchial sac. Its posterior margin has very small denticules. Dorsally, the the Dorsally, denticules. small very has margin posterior Its sac. branchial the of

lines, the test forms white swellings. The general consistency is not gelatinous but but gelatinous not is consistency general The swellings. white forms test the lines,

thickness. The oral siphon is wide and ends in 6 pointed lobes (Figs. 2A, B, C). C). B, 2A, (Figs. lobes pointed 6 in ends and wide is siphon oral The thickness.

rather firm. firm. rather

which correspond to the internal cloaca] channels. These furrows are easily seen seen easily are furrows These channels. cloaca] internal the to correspond which

transparent tunic; the species name refers to these lines (Pl. IB). Between the zooid zooid the Between IB). (Pl. lines these to refers name species the tunic; transparent

in living colonies, yellow in color from the pigmented zooids visible through the the through visible zooids pigmented the from color in yellow colonies, living in

surface is bare. The oral apertures of zooids open at both sides of depressed lines lines depressed of sides both at open zooids of apertures oral The bare. is surface

reef. It was encrusting, about 1 cm thick and 6 6 and 1 thick cm about encrusting, was It reef.

Holotype: Holotype:

A. multiplicatum multiplicatum A.

379), and Renganathan & Monniot (1984: 259). 259). (1984: Monniot & Renganathan and 379),

Tokioka (1953, 1954a, 1967), Millar (1963), Kott (1963, 1992a), Kott Kott 1992a), (1963, Kott (1963), Millar 1967), 1954a, (1953, Tokioka

following identifications of of identifications following

& &

body (1982), Nishikawa (1980, 1984, 1990), Rho (1975), Rho Rho (1975), Rho 1990), 1984, (1980, Nishikawa (1982), body

cies represented by the type, as they have an elongated post-abdomen with testes testes with post-abdomen elongated an have they as type, the by represented cies

11°45.lO'N; 124°58.87'E, 33 m. m. 33 124°58.87'E, 11°45.lO'N;

in 2 rows, a different stomach or a different cloaca] siphon. In our opinion the the opinion our In siphon. cloaca] different a or stomach different a rows, 2 in

constriction. The post-abdomen is short and contains many testis follicles in a a in follicles testis many contains and short is post-abdomen The constriction.

The post-abdomen is not continuous with the abdomen, but is separated by a a by separated is but abdomen, the with continuous not is post-abdomen The

bunch. bunch.

diately posterior to the cloaca] aperture, at the level of the second stigmata row. row. stigmata second the of level the at aperture, cloaca] the to posterior diately

nopee Tee sa anlr otsoah n cea Te ns pn imme opens anus The caeca. and post-stomach annular an is There incomplete.

agus is very short. The stomach has at least 20 longitudinal folds, few of them are are them of few folds, longitudinal 20 least at has stomach The short. very is agus

The oral siphon has 6 lobes. The cloaca] siphon is narrow, not quite tubular, but but tubular, quite not narrow, is siphon cloaca] The lobes. 6 has siphon oral The

a os fsimt. h admn smc salr hn h hrx Te esoph The thorax. the than smaller is much abdomen The stigmata. of rows 9 has

rather opening in a vertical slit with a horse-shoe shaped rim opening dorsally. A A dorsally. opening rim shaped horse-shoe a with slit vertical a in opening rather

and both lack larvae. They are cushion like with zooids arranged in double rows. rows. double in arranged zooids with like cushion are They larvae. lack both and

triangular languet is inserted on the dorsal rim of the aperture. The branchial sac sac branchial The aperture. the of rim dorsal the on inserted is languet triangular

h ooye n aohr ooy olce na tetp lclt, eerte dam rather were locality, type the near collected colony another and holotype the

papilla. papilla.

aged. However, it could still be determined that they are similar in all characters, characters, all in similar are they that determined be still could it However, aged.

of small ectodermal vesicles. Two odd median vesicles lie between each adhesive adhesive each between lie vesicles median odd Two vesicles. ectodermal small of

in the cloaca] cavity are small with 3 adhesive papillae, and an irregular half circle circle half irregular an and papillae, adhesive 3 with small are cavity cloaca] the in

follicles occupying the whole length of the post-abdomen. The larvae, incubated incubated larvae, The post-abdomen. the of length whole the occupying follicles

Lee (1989). The only descriptions that correspond well to the type specimen of of specimen type the to well correspond that descriptions only The (1989). Lee

The zooids are perpendicular to the colony surface, extending throughout its its throughout extending surface, colony the to perpendicular are zooids The

Description: The colony was collected on the deepwater slope of a fringing fringing a of slope deepwater the on collected was colony The Description:

Many specimens identified as as identified specimens Many

The preserved specimens of of specimens preserved The

MNHN: MNHN:

are those of Millar (1975: 247), Nishikawa & Tokioka (1976: (1976: Tokioka & Nishikawa 247), (1975: Millar of those are

A 1 Apl B 311-Indonesia, north Sulawesi, Manado area, area, Manado Sulawesi, north 311-Indonesia, B Apl 1 A

Aplidium Aplidium lineatum

Monniot & Monniot: New Ascidians Ascidians New Monniot: & Monniot

A. multiplicatum multiplicatum A.

Fig. 2, Pl. 1 1 Pl. B 2, Fig.

A. multiplicatum multiplicatum A.

new species species new

are erroneous: Van Name (1918), (1918), Name Van erroneous: are

x x

4.5 cm across. The mammillated mammillated The across. cm 4.5

that were examined, including including examined, were that

o o crepn t te spe the to correspond not do

& &

Huh (1984), Rho Rho (1984), Huh

& &

Good 137 137

gut; gut;

the the

of of

detail detail

D, D,

thoraces; thoraces;

C, C,

996 996

B, B,

; ;

zooi

A, A,

: :

n. n.

Micronesica 29(2), 29(2), Micronesica

stomach. stomach.

lineawm lineawm

of of the

ion ion

Aplidium Aplidium

sect

2. 2.

E, E,

Figure Figure

not not present.

ied colony. The The colony. ied ovary located is at the beginning of the posterior half of the post

intestine intestine with begins marked caeca at the bottom of the gut loop (Fig. 2C). The abdomen, followed by numerous testis follicles in two irregular lines. Larvae Larvae irregular two in abdomen, were lines. numerous by follicles followed testis

rarely rarely interrupted (Figs. 2D, E). The post-stomach makes a ring. The posterior

(Figs. 2C, D). D). The (Figs. 2C, with an longitudinal cylindrical stomach average 20 of folds, is elongated elongated post-abdomen contains (Fig. 2A) poorly-developed gonads in the stud

anterior anterior part; it not is perforated along the dorsal The line. gut about is the same

gitudinal. gitudinal. The branchial sac has 12 or 13 rows of stigmata, per 18 row in the

length length as the thorax. in The a esophagus swollen ring part middle its in slightly is

divided divided in (Figs. deeply cut 3 2A, Jobes C). The thoracic are muscles mostly lon

margin margin of the aperture prolonged is by a short languet, (Fig. either or 2B) simple 138 138

Aplidium tabascum tabascum Aplidium

shipping channel shipping

Material Examined: MNHN: Al Apl B 314-Papua New Guinea, Madang, main main Madang, Guinea, New 314-Papua B Apl Al MNHN: Examined: Material

Palau Islands. Islands. Palau

western Pacific (New Caledonia-Eastern Australia) and is now extended to the the to extended now is and Australia) Caledonia-Eastern Pacific (New western

preserved, this specimen is almost certainly certainly almost is specimen this preserved,

Wales, loaned by the Australian Museum in Sydney, and this specimen specimen obviously this Sydney, and in Museum Australian the by loaned Wales,

(1963) (1963) as

belongs to the genus genus the to belongs

cloacal cavity of one zooid, it was not yet differentiated into a tadpole. tadpole. a into differentiated yet not was it zooid, one of cavity cloacal of of

nies. nies.

colonies studied here. Only one embryo was found in the posterior part of the the of part posterior the in found was embryo one Only here. studied colonies

tectans tectans

merous testis follicles along its entire length entire its follicles testis along merous

The stomach is cylindrical with 5 folds. The post-abdomen is very long with nu with long is very post-abdomen The folds. 5 with is cylindrical stomach The

cm in length in cm

erally ending in 3 lobes 3 in ending erally

cloacal aperture is far anterior; its dorsal rim is elongated in a short languet gen languet short a in is elongated rim dorsal its anterior; far is aperture cloacal

not reach the endostyle the reach not

and retained its color when preserved. The zooids had red thoraces and yellow yellow and thoraces red had zooids The preserved. when color its retained and

abdomens. abdomens.

cm with a thickness of 3 of thickness a cm with

ytm rud omn laa apertures cloacal common around systems cushion attached to the substrate by a narrow part. The largest diameter is 6.5 6.5 is diameter largest The part. narrow a by substrate the to attached cushion

round hollows in which the oral apertures of the zooids are arranged in circular circular in arranged are zooids the of apertures oral the which in hollows round

Aplidium /ongithorax /ongithorax Aplidium Barrier Reef. Reef. Barrier

land., Australia Australia land.,

Aplidium protectans: protectans: Aplidium

Material Examined: MNHN: Al Apl B B Apl 331-Palau 317 and Al MNHN: Examined: Material

2" 71.2N 134°29 7°19.52'N; "29",

particularly large cloacal apertures cloacal large particularly

its zooids lining depressed cloacal channels, the numerous stomacal folds and the the and folds stomacal numerous the channels, cloacal depressed lining zooids its

Amaroucium protectans protectans Amaroucium

egahc Distribution Geographic

These characters are similar to those of specimens specimens of identified as those to similar are characters These

l hrces r ut iia otoe bevdi e aeoin colo Caledonian New in observed those to similar quite are characters All

Even though strongly contracted, the zooids are large, some exceeding 1.5 1.5 exceeding some large, are zooids the contracted, strongly though Even

The colonies are not very well preserved. They have the shape of a flattened flattened a of shape the have They well very preserved. not are colonies The

Remarks: Remarks:

(Herdman, 1899) by Kott (1992a). However, we examined the type colony type colony the (1992a). we 1899) However, examined Kott by (Herdman,

A. protectans, protectans, A.

. .

The oral aperture generally has 8 lobes, but occasionally 7 or 9 7 or occasionally 8 but lobes, generally has aperture oral The

Aplidium lineatum lineatum Aplidium

, ,

5°12.40'S; 145°48.85'E, 30 145°48.85'E, m. 5°12.40'S;

Kott

Aplidium /ongithorax /ongithorax Aplidium

Po/ycitor

Kott, 1963: 102, New South Wales, and 1992: 1992: 579, Queens and Wales, 1963: South 102, New Kott,

F. Monniot, 1987: 1987: 525 Monniot, F.

. .

lent by the Australian Museum, and while not very well well very while not and Museum, Australian the by lent

The branchial sac has 20 to 22 rows of stigmata that do do that stigmata of rows 22 to 20 has sac branchial The

5 cm for the middle of the colony. The surface surface is flat The with colony. of the 5 middle the for cm

Aplidium tabascum tabascum Aplidium 1992a: 589, fig.103, Queensland, Australia. Australia. 1992a: 589, fig.103, Queensland,

There is no constriction between thorax and abdomen. abdomen. and thorax between constriction no is There

62'E, 12 m; and Mutremdiu, 7°16.51'N; 134°31.55'E, 134°31.55'E, 7°16.51'N; Mutremdiu, and m; 12 62'E,

Herdman, 1899, collected in Port Jackson, New South South New Jackson, 1899, Port in collected Herdman,

Monniot Monniot

: :

The species was previously known from the south south the from known species previously was The

. .

We also examined the colony identified by Kott Kott by identified colony the examined also We

& &

n. sp. differs from other other differs from sp. n.

. .

Monniot: New Ascidians Ascidians New Monniot:

Pl. ID ID Pl.

Pl. IC IC Pl.

F. Monniot, 1987 1987 Monniot, F.

piim longithorax Aplidium

. .

Kott

, ,

The ovary was not developed in the the in developed not was ovary The

The test was yellow in life (Pl. 1 1 (Pl. life in yellow was test The

fig. C-G, pl. II F, New Caledonia. Caledonia. New F, II pl. fig. C-G,

, ,

1992 1992

Aplidium Aplidium

, ,

Rock Islands, Island Island Islands, Rock

. .

Aplidium pro Aplidium

species species to due

. .

The The

C) C) 139 139

is is

It It

in the the color, in and

A. A. tabascum

has has 5 which folds, are shorter

It It

sp. sp. A

F. F. Monniot, from 1987 New Caledonia,

eel/is eel/is

indicated indicated Kott by The number ofrows (1992a).

Aplidium Aplidium

Fig. 3A, B, Pl. 1 E E Fig. B, Pl. 3A, 1

A. A.

Micronesica Micronesica 29(2), 1996

mm mm

3 3

Geographic Geographic Distribution: The species was previously only recorded from

This This differs species from

Remarks: These colonies correspond correspond Remarks: to colonies These closely

The The zooids are perpendicular to the colony surface. The oral aperture has 6

The The colonies are encrusting, centimeters in several length and cm in thick 1

Kavieng, 2°45.06'S; 150°42.84'E, 20 20 m. Kavieng, 2°45.06'S; 150°42.84'E,

Material Material Examined: MNHN: Al Apl 313-Papua B New Guinea, New Ireland,

10 mm, is larger larger than is mm, the 10

Queensland. Queensland.

ter, a short post-abdomen and male follicles just just posterior a short ter, post-abdomen The to ovary. larvae the and follicles male

are slightly different. different. are slightly

which has a colorless tunic, a small simple cloaca} languet, languet, no has strong a tunic, which cloaca} colorless oral a small simple sphinc

always present present always in our specimens.

same except that that except same the interruption not was in middle of the the vesicles of crescent

of of stigmata, languet the cloaca} and the stomach are similar. The tadpole is the

arrangment arrangment of the zooids. But the of the size zooids, varying here between and 5

terrupted terrupted in its middle either on one or side both of sides the larva. The sensory

vesicle contains contains an vesicle and ocellus an otolith.

each each is side an anterior crescent of numerous button-like sometimes in vesicles,

cavity. Only Only cavity. one larva matures at a time and protrudes in a body wall dilatation

ovary ovary is visible, it is far behind the gut. The larvae are incubated in the cloaca)

which is not not is pedunculated. which The tadpole papillae on has adhesive thin 3 stalks. On

always longer longer always than the rest of the zooid and particularly long when the ovary is

starts starts with caeca. well-developed The post-abdomen is variable in length.

not not developed. The testis are follicles far posterior next to the heart. When the

stomach stomach is in the anterior half of the abdomen.

on on each side of the typhlosolis. No annular post-stomach was seen ; the rectum

stigmata. stigmata. The thorax narrow is and the number stigmata of not could counted. be

on on the anterior portion of the siphon. The branchial sac has 14 rows of long

The esophagus is short, short, The esophagus and conical is connects thinner it to the stomach. where The

and and 6th rows of stigmata. The cloaca) languet is on the dorsal rim of the siphon

and can be simple, bilobed or deeply cut into 3 lobes. The oral tentacles are located are located The oral tentacles lobes. cut into 3 deeply or bilobed simple, and be can

size, either a either in short size, tube or uncovering part the of branchial the 4th between sac

dentate dentate lobes above a strong sphincter. The cloaca) aperture is low, variable in

is particularly is particularly abundant around the oral siphon and above the endostyle.

times, at at times, junction the of canals, several are large common cloacal apertures. The

heights. The The with no red internal heights. sand around is tunic pigment The red the zooids. 140 140

colony colony surface is uneven because patches without sand have zooids of various

free channels channels along the free which oral apertures open in double (Pl. Some rows 1D).

areas with the tunic densely impregnated impregnated sand, with the tunic areas separated with densely sand depressed, by ness. The The colony divided uneven into because ness. is surface compartments of swollen Monniot & Monniot: New Ascidians 141

E E .....

Figure 3. Aplidium sp. (A), A: zooid; B, detail of the gut. Aplidium sp. (B): C, zooid.

The colonies (Pl. IE) are fleshy, cone-like with tapering but rounded ends, 3 to 6 cm long and I cm in diameter . The color is translucent brown, the zooid apertures are lined along deep long furrows which converge into circular common cloacal openings . Brown pigment cells are irregularly distributed in the thickness of the test and are absent in the zooids. The zooids are thin and elongated (Fig. 3A). The oral siphon has 6 digitiform lobes. The cloaca! siphon is narrow , with a muscular ring at its rim. A cloaca) 142 Micronesica 29(2), 1996 languet is located at some distance from the aperture (Fig. 3A). It is tapered at the free extremity or cleft in 2 or 3 teeth. The branchial sac has 13 to 15 rows of stigmata. The abdomen comprises a long esophagus, a long cylindrical stomach with 5 deep folds (Fig. 3B), an oval posterior stomach, a mid-intestine slightly dilated and a posterior intestine beginning with caeca. The post-abdomen is par ticularly long. The gonads are located at the very posterior extremity (Fig. 3A) with a short testis, with few follicles in two rows and an ovary immediately an terior to it. Some embryos were incubated in the cloacal cavity, but at an early stage of development. Remarks: This species differs from A. caelestis F. Monniot, 1987, which is encrusted with sand and has gonads occupying the entire abdomen. A. eel/is F. Monniot, 1987 has a short post-abdomen and an atrial languet on the rim of the cloacal aperture. A. clivosum Kott, 1992a has a different colony shape, a tubular cloacal siphon. A. crateriferum (Sluiter, 1909) has more rows of stigmata and gonads that do not reach the heart. A. lenticulum Kott, 1992a is encrusted with sand, has more rows of stigmata and a different abdomen. Aplidium longithorax F. Monniot, 1987 has zooids in rosettes, an atrial languet at the rim of the cloaca} aperture and more rows of stigmata. A. nadaense Nishikawa, 1980 is densely embedded with sand.

Aplidium sp. B Fig. 3C, Pl. 1F Material Examined: MNHN: Al Apl B 330-Palau, Koror State, Risong Lake, 7°18.74'N; 134°28.04'£, 1 m. The colonies are spherical, somewhat translucent. The oral siphons are seen as small red spots, lined in double rows converging into a single cloaca} opening, also of a deep red, at the top of each colony (Pl. lF). The colonies are grouped in clusters. They are soft and measure no more than 2 cm in diameter.The red color of the zooids completely fades in formalin. Fine sand particles are scattered throughout the soft tunic. The zooids are very short. The oral siphons have 6 round lobes. The cloaca} aperture is narrow with a sphincter and a tridentate languet at its upper rim (Fig. 3C). There are 10 longitudinal muscles on each side of the thorax. The branchial sac has 7 rows of stigmata with 10 to 15 stigmata in the first row (Fig. 3C). The abdomen and post-abdomen are very short and not clearly separated (Fig. 3C). The gut loop is narrow. The stomach has 5 deep folds. A cluster of testis follicles ( 10 to 12) is located just posterior to the gut loop, some are located inside the loop. The ovary was not observed and no larvae were pres ent.

Aplidiopsis oce/latus new species Fig. 4, Pl. 1G Holotype: MNHN: Al Apl A 15-Indonesia, north Sulawesi, west of Manado, l 0 23.50'N; 124°44.60'£, 7 m.

The cloaca! aperture is deeply cut between two lips (Figs. 4A, 4A, (Figs. lips two between cut deeply is aperture cloaca! The

exists below the posterior lip. The branchial sac has 17 rows of about 30 30 stigmata 17 about of rows sac has branchial lip. The exists posterior the below

margin with 5 distinct teeth (Fig. 4A). A small button-like protrusion sometimes sometimes protrusion button-like small A 4A). (Fig. teeth distinct 5 with margin

parallel. The edge of the oral siphon is undulated by contraction, but is lobed. lobed. is but contraction, by undulated is siphon oral the of edge The parallel.

ach is voluminous. The fusiform post-abdomen emerges from the middle of the the of middle the from emerges post-abdomen fusiform The voluminous. is ach gelatinous. gelatinous.

abdomen is shorter than the thorax, with the typical typical the with thorax, the than is shorter abdomen

displaced to the left side are wide at their base but with sharp extremities. The The extremities. sharp with but base their at wide are side left the to displaced

separated by high transverse membranes that lack papillae. The rapheal languets languets rapheal The papillae. lack that membranes transverse high by separated

is produced into a languet with an undulating tip; the posterior lip has a round round a has lip posterior the tip; undulating an with languet a into produced is

brown. The superficial test is tougher than the internal tunic which is soft and and soft is which tunic internal the than tougher is test superficial The brown.

yellow and olive pigment totally disappeared in formalin and the colonies turned turned colonies the and formalin in disappeared totally olive pigment yellow and

are 1 to 3 circular or stellate systems of zooids in each lobe of the colony. The The colony. the of lobe each in zooids of systems stellate or circular 3 1 to are

and some of the oral siphons are circled in yellow, hence the species species There yellow, circled hence the in name. are siphons oral the of some and

brown, which is the internal color of the colony. The common cloaca! openings openings cloaca! common The colony. the of color internal the is which brown,

ations (Pl. 1 G). The oral apertures and common cloaca! openings appear in dark dark in appear cloaca! openings common and 1 (Pl. apertures oral ations The G).

bv acmo bs.Te etsraei lv re,sot, ihu encrust without smooth, green, olive is surface test The base. common a above

The zooids are perpendicular to the colony surface with their post-abdomens post-abdomens with their surface colony the to perpendicular are zooids The

Description

Figure 4. 4. Figure

other zooid. zooid. other

Aplidiopsis ocellatus ocellatus Aplidiopsis

: :

The colonies have several clavate lobes, 1 to 2 cm high, raised raised high, cm 2 1 to lobes, clavate several have colonies The

Monniot Monniot

B B

n. sp.: A, B, zooid; C, thorax and abdomen of an of abdomen and thorax C, zooid; B, A, sp.: n.

& &

Monniot: New New Ascidians Monniot:

Aplidiopsis Aplidiopsis

:a :a

3 3

C); C);

shape. The stom The shape.

the dorsal lip lip dorsal the

C C 143 143

Mon

, ,

e e

. .

"' "'

N N

0 0

in some some in

; ;

but but

in in two parts

the the ovary was present

. .

zooid zooid

, ,

Only Only

larva

: :

, ,

; ;

larvae larvae in the colony studied,

brood brood pouch

Fig. Fig. 5

western western Australia and Queensland.

reserve reserve material.

Monniot Monniot & Monniot, 1987

, ,

empty empty

. .

Microncsica 29(2), 1996 1996 29(2), Microncsica

Monniot Monniot & Monniot, 1987

developed developed

Monniot, Monniot, 84 fig. L987a: 30 BC, Polynesia

& &

1992:463

abundant abundant

, ,

with with

Kott

Polyclinum Polyclinum pute

of of a zooid with an

Monniot Monniot

were were no fully

Polyclinum pule pule Polyclinum

New New Caledonia

. .

long, long,

. .

thorax thorax

514, 514,

There There

Figure Figure

1987: 1987:

c/inum c/inum tsutsui:

This This species is particularly characterized by the shape and color of the col

Pol

niot, niot,

Polyclinum Polyclinum pute

144 144

cloaca) cloaca) aperture

onies, onies, numerous rows of stigmata and the very peculiar shape of the lips of the

cloaca! cloaca! cavity.

zooids, zooids, a single young embryo was being incubated in the posterior part of the

gut gut loop; it is (Fig. (Fig. 4B).

ada Island, east side, side, 4°06 east Island, ada

Holotype: MNHN: Al Pse 31-Papua New Guinea, Duke of York Islands, Mak Islands, of York Duke Guinea, New Pse 31-Papua Al MNHN: Holotype:

Japanese species. The colonies from Philippines and Palau had no larvae, but in in but larvae, no had Palau and Philippines from colonies species. The Japanese

tified by Kott from Australia should be considered to be be to considered be should Australia from Kott by tified

pedunculate pouch. We also examined colonies Tokioka (1967) ascribed to his his to ascribed (1967) Tokioka colonies examined also We pouch. pedunculate

apparently lacks a pedunculate incubatory pouch. Therefore, the specimens specimens iden the Therefore, pouch. incubatory pedunculate a lacks apparently

embryos incubated in the cloaca! cavity, and inside the oviduct. There was no no was There oviduct. the inside and cavity, cloaca! the in incubated embryos

the largest of the 2 colonies from Gilbert islands zooids remained with young young with remained zooids islands Gilbert from colonies 2 the of largest the

proposed by Kott (1992a) cannot be be justified (1992a) cannot Kott by proposed

present in the oldest tadpoles of the New Caledonian specimens. specimens. Caledonian New the of tadpoles oldest the in present

trace of a protruding or pedunculate diverticulum, so Tokioka's species species so Tokioka's diverticulum, pedunculate or protruding a of trace illustrates a zooid with an embryo located in the cloaca! cavity with no trace of a a of trace no with cloaca! cavity the in located embryo an with zooid a illustrates

developed and there are no embryos. However, in pl. However, embryos. no are there and developed

been examined, but the zooids are not fully mature, the post-abdomens are poorly poorly are post-abdomens the fully not are mature, the zooids but examined, been

part that contains the gonads. There are between 10 to 20 testis follicles and the the follicles testis and 20 10 to between are There gonads. the contains that part

seen in polarized light. light. polarized in seen

Inside the visceral mass 1 to 3 characteristic crystals of calcium oxalate can be be can oxalate calcium of crystals characteristic 3 to 1 mass visceral the Inside

the two patches of vesicles in the larva, which were not described but were also also were but described not were which larva, the vesicles in of patches two the dermal ampullae anteriorly. A patch of small vesicles lies dorsally, on each side. side. each on vesicles small lies of dorsally, patch A anteriorly. ampullae dermal

oNwCldna colonies Caledonian New to anterior adhesive papillae on a thin and short stalk. There are 4 pairs of ecto of pairs 4 are There stalk. short and thin a on papillae adhesive anterior

The dorsal part is produced in a languet, generally long and narrow, ending in a a in ending narrow, and long generally languet, a in is produced part dorsal The

vr scnrl h aveaesaladrudd (Fig rounded small and are larvae The is central. ovary

They are separated by transverse sinuses bordered by 12 to 14 14 papillae 12 elongated to by sinuses bordered transverse by separated are They

has a long peduncle, that may reach 3 times the length of its enlarged posterior posterior enlarged its of length the times 3 reach may that peduncle, long a has

the thorax. The post-abdomen is inserted in the middle part of the abdomen; it it abdomen; the of part middle the in inserted is post-abdomen The thorax. the

point or numerous small teeth. A button-like protrusion is located dorsally under under dorsally is located protrusion button-like A small teeth. numerous or point

(Fig. SA). The abdomen has the typical shape of the genus and is folded under under folded is and genus the of shape typical the has abdomen SA). The (Fig.

in the middle part of the thorax, a stalked brood pouch contains up to 6 6 embryos to up contains pouch brood stalked a thorax, the of part middle the in

on each side. The body wall has strong muscular longitudinal bundles. Dorsally, Dorsally, bundles. longitudinal muscular strong has wall body The side. each on

Ngermutidech, 7°18.64'N; 134°31.12'E, 0-1 m, seagrass bed. bed. seagrass m, 134°31.12'E, 0-1 7°18.64'N; Ngermutidech,

the cloacal siphon (Figs. (Figs. siphon cloacal the

mon cloacal openings. The tunic is rubbery, internally devoid of sand and soft. soft. and sand of devoid internally rubbery, is tunic The openings. cloacal mon

eesars n ecutage h oisfr crua ssesaon com around systems circular form zooids The algae. encrust and across meters

at their base and in small patches on the surface. The colonies are several centi several are colonies The surface. the on patches small in and base their at

of the branchial sac, almost at the same level as the oral siphon (Figs. (Figs. siphon oral the as level same the at almost sac, branchial the of

6 sharp oral lobes (Figs. (Figs. lobes oral 6 sharp

Black pigment cells are present everywhere. The zooids have a large thorax with with thorax large a have zooids everywhere. The present cells are pigment Black

Material Examined: MNHN: Al Pol SO-Palau, Airai State, S.W. Airai Channel, Channel, S.W. Airai State, Airai SO-Palau, Pol Al MNHN: Examined: Material

Geographic Distribution: Palau, Queensland, New Caledonia, and Polynesia. Polynesia. and New Caledonia, Queensland, Palau, Distribution: Geographic

All characters correspond closely to the type specimen from Polynesia and and Polynesia from specimen type the to closely correspond characters All

Remarks: The synonymy of of synonymy The Remarks:

The colonies are black, somewhat greenish internally. Some sand is is Some sand greenish internally. encrusted somewhat black, colonies are The

Pseudodistoma coronatum coronatum Pseudodistoma

SA, SA,

8S 152°25.45'E 28'S; Monniot & Monniot & Monniot

. .

B). The cloacal siphon opens horizontally at the top top the at horizontally opens siphon cloacal B). The

The only difference with the original description is is description original the with difference only The

B). There are 12 to 14 rows of 20 to 25 stigmata. stigmata. 25 to 20 of 14 rows 12 to are There B).

Figs. 6A-E, Pl. SH SH Pl. 6A-E, Figs.

. .

pute pute

with with

: :

New Ascidians Ascidians New

, ,

. .

15 15 m.

The type specimen specimen of type The

Polyclinum tsutsui tsutsui Polyclinum

new species species new

XVIII XVIII

. .

SC). SC). possess 3 wide They

. .

fig. 2, Tokioka (1954) (1954) fig. 2, Tokioka

pute

. .

Tokioka, 1954b 1954b Tokioka,

. .

tsutsui tsutsui

P. tsutsui tsutsui P.

SA, SA,

has has

B). B). 145 145 146 Micronesica 29(2), 1996

Figure 6. Pseudodistoma corona/um n. sp.: A, B, zooids; C, detail of the gut; D, section of the stomach; E, larva. Pseudodistoma mega/arva n. sp.: F, G, H, di.fTerentzooids ; I, detail of the gut; J, larva.

joined cushions 2 to 5 cm across. They had a translucent test through which which bright test through translucent a 5 2 to had They cushions cm across. joined

musculature is weak. The digestive tract makes a tight, straight loop (Fig. 61) 61) (Fig. loop straight tight, a makes tract digestive The weak. is musculature

rapheal languets are short and clearly displaced on the left side. The thoracic thoracic The side. left the on displaced clearly and short are languets rapheal

rounded lobes (Figs. 6G, H). There are 3 rows of about 20 long stigmata. The 2 2 The stigmata. long 20 about of rows 3 are There H). 6G, (Figs. lobes rounded

hrx n admn oehr (Figs together abdomen and thorax

sediment. sediment.

orange zooids could be seen (Pl. 2A). The colony surface was smooth, without without smooth, was surface colony The 2A). (Pl. seen be could zooids orange

2°08.48'N

ooye MH: l s 27 Pse Al MNHN: Holotype:

ranged in circular systems and the cloaca! siphon has a characteristic shape. shape. characteristic a has siphon cloaca! the and systems circular in ranged

mata which are already pierced (Fig pierced already are which mata

of ectodermal ampullae. They have an ocellus and an otolith and 3 rows of stig of rows 3 and otolith an and ocellus an have They ampullae. ectodermal of

The tadpoles are small, 500 µm, with 3 very short adhesive papillae and 4 pairs pairs 4 and papillae adhesive short very 3 with µm, 500 small, are tadpoles The

the posterior half of the post-abdomen (Fig. 6B). The embryos are incubated all all incubated are embryos 6B). The (Fig. post-abdomen the of half posterior the

from the gut loop. When the testis is developed, few male follicles are located in in located follicles are few male developed, is testis the When loop. gut the from

along the oviduct, and only the mature tadpoles reach the thoracic cloaca! cloaca! cavity. thoracic the reach tadpoles mature the only and oviduct, the along

peared (Fig peared

the abdomen. When the ovary is present, the male gonads have generally disap generally have gonads male the present, is ovary the When abdomen. the

begins with 2 caeca. The post-abdomen is not separated by a constriction from from constriction a by separated not is post-abdomen The caeca. 2 with begins

97 hc a smlrgt n gnd. u tepeet pce a zod ar zooids species has present the But gonads. and gut similar has 1957) which

left side. The abdomen is longer than the thorax even in relaxed zooids (Figs. 6A, 6A, (Figs. zooids relaxed even in thorax the than is longer abdomen side. left The

the abdomen length. It is asymmetrical with 4 swellings swellings 4 with is It asymmetrical (Figs length. abdomen the

have 20 stigmata or less. The dorsal languets are short, obviously displaced to the the to displaced obviously short, are languets less. dorsal or The 20 stigmata have

B). The esophagus is long and cylindrical, the short stomach is placed at 2/3 of of 2/3 at placed is stomach short the cylindrical, and long is esophagus The B).

are 8 longitudinal muscle bands on each side of the thorax. The first row of of row first The thorax. the of side each on bands muscle longitudinal 8 are

much shorter when the zooids are breeding with a regressed post-abdomen. There There regressed post-abdomen. a with breeding are zooids the when shorter much

stigmata is dorsally curved and contains about 22 perforations 22 about contains and curved dorsally is stigmata

surface, long and thin when the post-abdomen is fully developed with gonads, but but is gonads, fully with developed post-abdomen the when thin and long surface,

are oblique at the tip of the tubes. The zooids are perpendicular to the colony colony the to perpendicular are zooids The tubes. the of tip the at oblique are

cloaca! openings are grouped in the center of these circles. circles. these of center the in grouped are openings cloaca!

asymmetrical with 3 dorsal lobes longer than the 3 ventral ones, so the apertures apertures the so ones, 3 ventral the than longer lobes 3 dorsal with asymmetrical

yellow in formalin and the fixative takes a yellow color. The oral siphons open in in open siphons oral The yellow a color. fixative takes the and yellow formalin in

rings (hence the species name) at the smooth upper surface of the colony. The The colony. the of surface upper smooth the at name) species the (hence rings

the tunic is translucent, pale yellow mottled with white (Pl. lH), the zooids light light zooids the lH), (Pl. white with mottled yellow pale translucent, is tunic the

cm across, on rock or sponges. The consistency is firm and rubbery. When living, living, When rubbery. is consistency firm and The sponges. or rock on across, cm

orange and visible where the tunic is somewhat transparent. The zooids remain remain zooids The transparent. somewhat is tunic the where visible and orange

The zooids are long and thin, the post-abdomen is much longer than the the than longer much is post-abdomen the thin, and long are zooids The

Description

Remarks: This species is closely related to to related closely is species This Remarks:

The oral apertures of zooids have 6 rounded lobes. The cloaca! siphons are are siphons cloaca! The lobes. rounded 6 have zooids of apertures oral The

Description

; ;

125° 21.15'E, 12 12 m 125° 21.15'E,

. .

6A) and the post-abdomen is shorter, the ovary lying lying ovary some the distance at is shorter, the post-abdomen 6A) and

: :

Collected on the shaded side of a patch reef, the colonies were were colonies the reef, patch a of side shaded the on Collected

The colonies are encrusting sheets, 0.5 to 1 cm thick, up to 10 10 to up thick, 1 cm to 0.5 sheets, encrusting are colonies The

Pseudodistoma megalarva megalarva Pseudodistoma

Monniot & Monniot: New Ascidians Ascidians New Monniot: & Monniot

. .

Figs

Indonesia, North Sulawesi, Biaro Island, Island, Biaro Sulawesi, North Indonesia,

. .

6F, G, H). Both apertures are short with 6 6 with short are apertures Both H). G, 6F,

6E). 6E).

J, Pl. 2A 2A Pl. J,

new species species new

Pseudodistoma Pseudodistoma aureum

. .

6C, D). The rectum rectum The D). 6C,

. .

The other rows rows other The

(Brewin, (Brewin, 147 147

embryos embryos remain in the

.The .The

new species new species

Fig. Fig. 7, Pl. 2B

Federated Federated States of Micronesia, Chuuk State,

Micronesica Micronesica 29(2), 1996

The cloacal siphon is short. short. The The siphon is rows cloacal branchial has 3 sac

The The gut is a cylindrical tube interrupted half way down

Pseudodistoma Pseudodistoma poculum

C). C).

Al Al Pse 28

C). C).

MNHN: MNHN:

The The zooids are very A). long (Fig. Both siphons 7 have 6 short lobes. The

Description: The colonies (Pl. 2B) form form a pedunculate 2B) ramified (Pl. of bush The lobes Description: colonies

Remarks: Remarks: characterized is This species by the shape and of the size tadpoles

tremity tremity with no more than testis and 10 follicles an ovary containing one large

incubates incubates at a time in the cloaca! cavity. It is oval, the tail shorter than a half

is extremely is extremely long and A). thin The (Fig. gonads 7 are limited to the cardiac ex

oocyte oocyte (Fig. The 7E). larvae are small, the trunk µm 400 long. Only one tadpole

the the abdomen by a rounded stomach without folds (Fig. 7D). The post-abdomen

of about about of stigmata. The abdomen 10 and thorax are of equal length and the same

the colony (Figs. 7B, 7B, colony (Figs. the

diameter diameter (Figs. 7B,

post-abdomens. post-abdomens. They also have epibionts diverse on their surface.

distance distance the between siphons varies, according to the location of each zooid in

red red spots. The stalks are up to 3 cm long with a maximum diameter of 3 mm.

siphons siphons the tunic contains fine sediment particles giving a white design between

open open independently. The oral apertures are regularly distributed on the external

They are isodiametrical, ramified, completely encrusted encrusted with sand and completely contain the They ramified, are isodiametrical,

side side of the head, circled by a ring of red tunic, without inclusions. the Between

diameter, diameter, with a top hollow, resembling a common aperture. cloaca} On the in

ternal ternal side of this "cup" (from which the is named), species the cloacal siphons

Kuop Kuop E, 30 m. Atoll, reef, oceanside 7°00.00'N; 151°56.05

arising arising from a narrow base. Each lobe ends in a spherical head, up to 7 mm in

Holotype: Holotype:

after after fixation. The deep orange-red pigment soluble highly in is alcohol.

papillae papillae were not in a straight line, but this may have been due to contraction

and and their incubation in the post-abdomen. It should be noted that the adhesive 3

median median are ones undivided.

pillae pillae on each side, the dorsal and ventral ones are divided (Fig. 6J), while the

papillae papillae are close to each other and also elongated. There are 4 ectodermal pa

sorial vesicle, with an ocellus and and an an ocellus with posteriorly. otolith, located adhesive The is 3 vesicle, sorial

trunk trunk can reach mm in 1.6 length (Fig. justifying 6J) the name. species The sen

cylindrical cylindrical shape. The tail forms a sharp angle before encircling the trunk. The

oviduct, oviduct, below the abdomen (Fig. 6F), they are large, compressed into a long

6F, 6F, develop along the post-abdomen H). The oocytes

61). The post-abdomen The post-abdomen varies 61). depending on the gonadal maturation When stage.

half half of the post-abdomen (Figs. 6G, H). The ovary appears far posterior (Figs.

148 148

the testis follicles are well developed, they lie in two irregular irregular two in the posterior in rows lie they developed, well are follicles the testis

segment segment at the bottom of the gut loop. The pyloric gland curved has tubules few

converging to to converging a thin duct opens into which the posterior part the of stomach (Fig.

beginning with with a long and beginning narrow esophagus enters which dorsally into an asym metrical, metrical, smooth-walled stomach (Fig. The 61). rectum begins with an enlarged

-;p.I ;F Mnit 1990 Monniot, g; F. IV pl. c-i;

Synonymy: see Kott, 1980: 1980: 1982: 10; 111; Monniot see Kott, Synonymy: cloaca} cavity, which does not exist in in exist not cloaca} does which cavity,

colony shape, the deep depressions at the top of the lobes resembling a common common a resembling lobes the of top the at depressions deep the shape, colony

already has 3 rows of stigmata. In formalin, the tadpoles are pigmented in in orange pigmented are tadpoles the formalin, In 3 rows of has stigmata. already

ectodermal vesicles (Fig. 7F). Ventrally some digitations are also present (Fig. (Fig. present also are digitations some Ventrally 7F). (Fig. vesicles ectodermal

perimeter of the trunk (Fig. 7F). There are 3 adhesive papillae and 4 pairs of of pairs 4 and papillae adhesive 3 are There 7F). (Fig. trunk the of perimeter

brown, they were probably red when living. living. when red probably were they brown,

7F). The sensory vesicle is proportionally large. The branchial sac of the oozooid oozooid the of sac branchial The large. vesicle sensory is The proportionally 7F).

Remarks: This species differs from all other other all from differs species This Remarks:

A A

Figure 7. 7. Figure

different zooids; D, detail of the gut; E, detail of the gonads; F, larva. larva. F, gonads; the of detail E, gut; the of detail D, zooids; different

B B

Pseudodistoma poculum poculum Pseudodistoma

Trididemnum cyclops cyclops Trididemnum

Monniot Monniot

0 0

3 3

Vl Vl

Family: Didemnidae Didemnidae Family:

: :

520, New Caledonia 520, New

& &

Monniot: New New Monniot: Ascidians

. .

n. sp.: A, zooid; B, C, thorax and abdomen of of abdomen and thorax C, B, zooid; A, sp.: n.

' '

,. ,.

' '

, ,

Pseudodistoma arborescens arborescens Pseudodistoma

Michaelsen, 1921 1921 Michaelsen,

Vl Vl

,_, ,_,

0 0

3 3

D D

Pseudodistoma Pseudodistoma

& &

. .

Monniot, 1987a: 1987a: 2 fig. 20, Monniot,

Vl Vl

0 0

3 3

Millar, 1967. 1967. Millar,

species by the the by species

E E 149 149

mm mm

0,25 0,25

The The cloacal aperture

. .

8A)

. .

B B

new new species

A, A, thorax; B,abdomen, C, detail of the

: :

n. n. sp.

Palau, Palau, Koror State, outside 7°10.7'N; Krist reef,

Fig. Fig. 8, Fig. 9A,

Micronesica Micronesica 29(2), 1996

A2 A2 Tri and 106 A2 Tri 100-Palau, Koror State,

Trididemnum polyorchis polyorchis Trididemnum

MNHN: MNHN:

A3 Tri Tri A3 111

and and other algae, this shallow, likes species lit habitats. well It is

Trididemnum Trididemnum po/yorchis

. .

gut. gut.

MNHN: MNHN:

Figure Figure 8

Halimeda Halimeda

The The zooids are extremely small, 200 µm for the thorax and µm 300 for the

Description: Description: The colony pinkish was brown and cream with in a life, "leop

Geographic Geographic Distribution: This has species a wide distribution in all tropical

The The pad-like colonies have varied colors according to the abun respective

134°22.1 'E, 'E, 8 m. 134°22.1

areas. areas. The lateral thoracic organs form large cups above the second row of stig-

sac. There are 3 rows of stigmata of stigmata There rows the anterior are between sac. 3 and posterior unperforated

is sessile and and circular sessile is and a exposes part small of the dorsal of side the branchial

throughout throughout the entire tunic except at the uppermost surface layer of the colony

gut gut loop. The oral siphon is tubular and narrow (Fig

to to break, with a rounded outline. There are irregular of zooids, systems with oral

1°34.lO'N; 124°47.93'E, 1 m. m. 1°34.lO'N; 124°47.93'E, 1

ard" ard" design, several cm across and 2 mm thick. This is species encrusting, hard colony colony cannot be delaminated.

which is spicule-free. The cloaca} channels channels are The narrow, cloaca} spicule-free. which is poorly developed and the

Holotype: Holotype: siphons siphons surrounding small common apertures. cloaca} The are spicules crowded

150 150

of of the tadpole with only 2 papillae. adhesive

shallow shallow waters of the Indian, western, and central Oceans. Pacific

principally principally characterized by a short dorsal lobe at the oral siphon and the shape

grass, grass,

Rock Rock Islands, 7°20.19'N; m; Indonesia, 134°32.12'E, 1 north Manado, Sulawesi,

Material Material Examined: dance dance of symbiotic unicellular algae and calcareous spicules. Collected on sea

D, D,

Figure 9. Spicules. A, B, B, A, Spicules. 9. Figure

IOµm

Didemnum guttatum guttatum Didemnum

. .

sp. E, E, sp.

Trididemnum polyorchis polyorchis Trididemnum

Didemnum linguiferum linguiferum Didemnum

Monniot Monniot

& &

Monniot

. sp n.

: :

New New Ascidians

n. sp n.

. .

C, C,

. .

Didemnum biglutinum biglutinum Didemnum

F, F,

Didemnum nigrum nigrum Didemnum

n. sp n.

Monniot, 1994. 1994. Monniot,

. .

Scale bars: bars: Scale

1 51 51

Romanov, Romanov, 1989,

T. T. vostoki

Faulkner's Faulkner's group, is the second

J. J.

F. F. Monniot, 1995

Pl. Pl. 2C

Monniot Monniot & Monniot, 1987

collected collected by

Fig. Fig. 9C, Pl. 2D

Micronesica Micronesica 29(2), 1996

A2 A2 Did C 295-Palau, Koror State, Rock Islands,

F. F. Monniot, 300, 1995: fig. pl. 1, I A and III D; New Cal

MNHN: MNHN:

Didemnum Didemnum biglutinum

Trididemnum, Trididemnum,

Monniot Monniot & Monniot, 27, Polynesia; 1987a: F. Monniot, 1995:

Didemnum Didemnum apuroto

The The zooids are small and difficult to extract from the tunic. The oral siphon

The The colonies are encrusting thin sheets, inflated underwater, of a greenish

Geographic Geographic Distribution: This species described from Polynesia and found

The The colonies are white, encrusting on ramified bryozoans and algae (Pl. 2C).

Remarks: Remarks: This

The The spicules are rather large compared to the size of the zooids. They have

is very very is short with 6 round lobes. The cloacal aperture forms an horizontal slit, as

ible. ible. The pigment is mostly in irregular specks in the upper part of the tunic.

sheet sheet without spicules against the substratum. The whole tunic is tough, but flex

There There a is layer with crowded spicules between the zooid layer and the thin fibrous

light light grey color, with black spots (Pl. 2D). In formalin they are dark brown. The

without without pigment or cells spicules. The zooids are in the upper part of the colonies.

Material Material Examined:

common common cloaca} apertures are irregularly raised in chimneys of translucent tunic

Ngerkuul Ngerkuul Pass, 7°19.22'N; 134°29.74'E, m. 10

Didemnum Didemnum biglutinum

edonia. edonia.

to to distinguish from other white didemnids.

in in New Caledonia and Micronesia is probably more widespread but it is difficult

thin thin tunic.

the the colony surface. The zooids and larvae are similar to those from Polynesia and

Chuuk Chuuk lagoon, 7°22.30'N; 151°50.64'E, 9 m.

Material Material Examined: MNHN: A2 Did C 306-Federated States of Micronesia, New New Caledonia, they are very small. The spicules are crowded throughout the

300, 300, New Caledonia.

They They are soft, very thin. When fixed the zooids appear as small protuberances at

Didemnum Didemnum apuroto

species of of species the genus having more than one testis lobe.

and and colony shape, and numerous coils of the sperm duct.

from from Peter the Great Bay, in the Japan Sea, has larger zooids, different spicules

an an average diameter of µm 40 and a maximum of µm. 50 They size are star-shaped

with with numerous short conical rays (Figs. 9A, B).

152 152 loose loose of coils the sperm duct (Fig. The 8B). ovary was not developed in the studied

colony. colony. No larvae were found.

loop, loop, is made of 2 or sometimes, 3 very protruding follicles, encircled by 3 to 4

under under the thorax. The gut is a wide loop (Fig. 8C). The testis, as large as the gut mata mata (Fig. 8A). The retactor muscle is long. The abdomen lies at a right angle

Guinea, Eastern Fields Atoll (Coral Sea), 140 km SW of Port Moresby, Moresby, Port of SW km 140 Sea), (Coral Atoll Fields Eastern Guinea,

sad I Island,

Holotype: MNHN: A2 Did C 229-Indonesia, north Sulawesi, west of Talisei Talisei of west Sulawesi, north 229-Indonesia, C Did A2 MNHN: Holotype:

Ocean, but is difficult to collect. collect. to difficult is but Ocean,

in the oozooid. The spicules are made of very numerous needles in balls. balls. in needles numerous very of made are spicules The oozooid. the in

species probably has a wide distribution in the tropical central and western pacific pacific western and central tropical the in wide a distribution has species probably

intestine forms a twisted loop twisted a forms intestine

zooids have a large thorax with a wide cloaca! aperture without a languet. The The languet. a without aperture cloaca! wide a with thorax large a have zooids

between 4 pairs of long ectodermal ampullae. The stigmata are developed early early developed are stigmata The ampullae. ectodermal long of pairs 4 between

l0°02.48'S; 145°36.28'E., 23 23 m. 145°36.28'E., l0°02.48'S; large with the trunk about 750 µm long. The 3 adhesive papillae are urn-shaped urn-shaped are papillae adhesive 3 The long. 750 µm about trunk the with large

fig. 8, 8, fig.

Didemnumfragi/is Didemnumfragi/is

7°06

Material Examined: Examined: Material

Northern Japan has only 3 pairs of larval ectodermal ampullae. ampullae. ectodermal larval of 3 pairs only has Japan Northern

Monniot & Monniot, 1987a has a different color, a large oral siphon, and 6 pairs 6 pairs and siphon, oral large a color, different a 1987a has & Monniot, Monniot

2 adhesive papillae and four pairs of ectodermal ampulllae. ampulllae. ectodermal of pairs four and papillae adhesive 2

of ectodermal ampullae in its larva. larva. its in ampullae ectodermal of

ular rays gathered in balls (Pl. 1 1 (Pl. balls in gathered rays ular

papillae with thin and long stalks. They have 4 pairs of digitiform ectodermal ectodermal digitiform of pairs 4 have They stalks. long and thin with papillae

otolith are present. The tail is short, in half a circle around the trunk. trunk. the around circle a half in short, is tail The present. are otolith

buds; its branchial sac is well developed with 3 rows of stigmata. The ocellus and and ocellus The stigmata. of 3 rows with is well sac developed branchial its buds;

with 7 to 8 coils of the sperm duct. The ovary is normally placed between the the between placed normally is ovary The duct. sperm the of coils 8 to 7 with

papillae on each side, the dorsal pair larger than the others. The larva has no no has larva The others. the than larger pair dorsal the side, each on papillae

anterior, followed by a post-stomach, mid-intestine and rectum well defined. The The well defined. rectum and mid-intestine post-stomach, a by followed anterior,

testis and the stomach. The larvae are small, 550 µm. They only have 2 adhesive adhesive 2 have only They 550 µm. small, are larvae The stomach. the and testis

The abdomen is folded under the thorax. The gut loop is wide, the stomach is is stomach the wide, is loop gut The thorax. the under folded is abdomen The

ning close to the posterior end of the endostyle. There are 6 stigmata in a row. row. a in stigmata 6 are There endostyle. the of end posterior the to close ning

spherical testis follicle clearly protrudes from the middle of the gut loop, encircled encircled loop, gut the of middle the from follicle testis spherical protrudes clearly

third transversal branchial sinus. The retractor muscle is moderately long, begin long, is moderately muscle retractor The sinus. branchial transversal third

a result of the contraction. The thoracic organs are round and placed above the the above placed and round are organs thoracic The contraction. the of result a

This species is bright white, encrusting, soft but with crowded spicules. The The spicules. crowded with but soft encrusting, white, bright species is This

Geographic Distribution: Malaysia, New Caledonia, Polynesia, Palau. The The Palau. Polynesia, Caledonia, New Malaysia, Distribution: Geographic

Geographic Distribution: New Caledonia, Palau. Palau. Caledonia, New Distribution: Geographic

h pclsaesal 3µi mxmm diameter maximum µmin 33 small, are spicules The

Remarks: This species differs from other tropical tropical other species from differs This Remarks:

8'N; 134°15.6'E, 15-25 m. m. 15-25 134°15.6'E, 8'N;

FH, FH,

51.87'N; 125°04.08' E., I m. MNHN: A2 Did C 279-Papua New New 279-Papua C Did A2 MNHN: m. I E., 125°04.08' 51.87'N;

pl. I, I, pl.

H, H,

Sluiter, 1909: 56, Malaysia; Monniot Monniot Malaysia; 1909: 56, Sluiter,

Polynesia; F. Monniot, 1995: 310, New Caledonia. Caledonia. 1995: 310, New Monniot, F. Polynesia;

MNHN: MNHN:

Didemnumfragilis Didemnumfragilis

Didemnum guttatum guttatum Didemnum

Monniot Monniot

Figs. 9D and I I 2E Pl. 0, and 9D Figs.

. .

A2 Did C 326-Palau, Koror State, Big Big Drop-Off, State, Koror 326-Palau, C Did A2

The larvae have a characteristic appearance; rather rather appearance; characteristic a have larvae The

C). C).

& &

Monniot: New Ascidians Ascidians New Monniot:

Didemnum risirense risirense Didemnum

(Sluiter, 1909) 1909) (Sluiter,

new species species new

Didemnum Didemnum

& &

. .

Nishikawa, 1990 from from 1990 Nishikawa,

hyhv ay acic many have They

Monniot, 1987a: 30, 30, 1987a: Monniot,

Didemnum pitipiri pitipiri Didemnum

by its larva with with larva its by 153 153

i3 i3

'.] '.]

U1 U1

a a

is is covered with 7 to 8

It It

A, A, thorax; B, abdomen; C, D, larvae.

: :

n. n. sp.

Micronesica 29(2), 1996 1996 29(2), Micronesica

Didemnum Didemnum guttatum

Figure Figure 10.

The The zooids have color a greenish in formalin. The oral siphon particularly is

Description: Description: The are colonies encrusting, up to 4 mm thick and cen several

anterior anterior position, small and (Fig. especially is the IOB) intestine does not exhibit

undivided undivided testis overlaps the mid-intestine (Fig. IOB).

well separated separated well The segments. gonads are in the gut loop, but, when mature, the

above above the 4th stigmata row, near the endostyle (Fig. The retractor IOA). muscle

is is not folded under the thorax. The gut loop is wide. The stomach, in a very

in the first of the the in The the of first 4 rows. thoracic organs are protruding, circular, and placed

its dorsal margin is produced into a short languet (Fig. 1 OA). There There stigmata are 6 dorsal OA). produced its margin is into a short languet (Fig. 1

extends extends from the posterior extremity of the esophago-rectal neck. The abdomen

long, long, ending in 6 short lobes The (Fig. cloaca] aperture IOA). a is transverse slit,

spicules are are spicules crowded throughout the test. The zooids are numerous, the cloaca}

mulated mulated in clumps, are present in the part deepest of the colony.

channels channels are narrow and limited to the thorax Abundant level. dark accu cells,

timeters across, across, timeters with an irregular margin. marbled The is surface and rough. Uni

white white and grey or brown (Pl. 2E). The openings common are cloaca} round. The

cellular green cellular algae present were in patches at the colony surface. The color was 154 154

There are 3 adhesive papillae circled by about 30 elongated ectodermal ampullae ampullae ectodermal 30 elongated about by circled papillae 3 adhesive are There

D). The larvae reach 800 µm in length. The tail is is tail The length. in µm 800 reach larvae The D).

the intestinal loop, its top covered by 5 to 7 turns of the sperm duct (Figs. llA, llA, (Figs. duct sperm the of turns 7 to 5 by covered top its loop, intestinal the

intestine found in the genus (Fig. 11 11 (Fig. genus the in found intestine

at the begining of the gut loop. It is followed by the common segments of the the of segments common the by followed is It loop. gut the of begining the at

a white mass which does not stain with hemalum. The undivided testis testis lies besides undivided The hemalum. with stain which not white does mass a

sac. The retractor muscle is along the short esophago-rectal waist. The abdomen abdomen The waist. esophago-rectal short the along is muscle retractor The sac.

lies horizontally. The esophagus is very short, so the spherical stomach is is located stomach spherical the so short, is very esophagus The lies horizontally.

located vertically on the rim of the cloaca! aperture at the level of the third row row third the of level the at aperture cloaca! the of rim the on vertically located

of stigmata (Figs. 1 lA, B). There are 6 stigmata in the 1 2 the in first of branchial the rows 6 (Figs. are B). stigmata There of stigmata lA,

tentacles are numerous are tentacles

of the opening (Figs. 1 lA, B). On each side the lateral organs are large lamellae lamellae large are organs lateral the side each On 1 B). (Figs. lA, opening the of

under it. The oral siphon is tubular with 6 short lobes (Figs. l lA, B). The oral oral The B). l lA, (Figs. lobes short 6 with tubular is siphon oral The it. under

branchial sac. A small languet, simple or bifurcated, is located at the upper rim rim upper the at located is bifurcated, or simple languet, small A sac. branchial

level. The spicules are dense throughout the colony. colony. the throughout dense spicules are level. The

cloaca} cavities are wide and the colony is easily tom in two sheets at the thoracic thoracic the sheets at two in is easily tom colony cloaca} the wide and cavities are

the branchial sac, and some pigment is also found above the testis. The common common The testis. the above found also is pigment some and sac, branchial the

with white, with star-shaped common cloaca} apertures. The pigment is is restricted pigment The cloaca} common apertures. star-shaped with white, with

and 1 to 2 mm thick. In formalin the surface of the colony is reddish, marbled marbled reddish, is colony the of surface the formalin In thick. mm 2 to 1 and

to the surface layer in the tunic, but red cells are abundant in the thorax, especially especially thorax, the in red cells but abundant are surface layer tunic, the in the to

town. town.

Holotype: Holotype:

covered with symbiotic algae, which seemed absent from the colonies found at 23 23 at found colonies the from which seemed absent algae, symbiotic with covered

m in New Guinea. Guinea. New in m

ture, and the pigmentation. Collected at only 1 m depth in Sulawesi, Sulawesi, was partly it in 1 depth only m at Collected pigmentation. the and ture,

vae are not gemmiparous. gemmiparous. not are vae

arising from a compact center. Their diameter reaches 70 70 µm. reaches diameter Their center. compact a from arising crescent under the tail attachment. There is also one patch on each side, side, posterior each on patch one is also There attachment. tail the under crescent

to the sensory vesicle. The 3 adhesive papillae are short and wide, circled by a a by circled wide, and short are papillae adhesive 3 vesicle. The sensory the to

patches justifying the species name (Figs. lOC, D). These pigments are green in in green are pigments These D). lOC, (Figs. name species the justifying patches

The tail makes makes tail The ring of numerous, rather irregular ectodermal ampullae, (Figs. IOC, D). The lar IOC, The D). (Figs. ampullae, ectodermal irregular rather numerous, of ring

coils of the sperm duct. The ovary lies between the testis and the stomach. The The stomach. the and testis the between lies ovary The duct. sperm the of coils formalin. There are 3 odd patches: one dorsal, one round ventral, and one in a a in one and ventral, round one dorsal, one patches: odd 3 are There formalin.

found at the colony surface. They are large, some with a trunk l mm in length. length. in mm l trunk a with some large, are They surface. colony the at found

larvae are incubated in the test below the zooids, but when mature, they can be be can they mature, when but zooids, the below test the in incubated are larvae

The zooids have a narrow thorax, that is larger than the abdomen folded folded abdomen the than larger is that thorax, narrow a have zooids The

Description: The colonies are thin, flat, encrusting, about 10 cm in extent extent in cm 10 about encrusting, flat, thin, are colonies The Description:

The main characters of the species are the wrinkled surface, the larval struc larval the surface, wrinkled the species are the of characters main The

The spicules (Fig spicules The

MNHN: MNHN:

¼ ¼

of the body circle. The tadpoles have characteristic pigment pigment characteristic have tadpoles The circle. body the of

A2 Did C 307-Indonesia, north Sulawesi, near Manado Manado near Sulawesi, north 307-Indonesia, C Did A2

. .

Didemnum Didemnum linguiferum

The cloacal aperture is large, exposing a large part of the the of part large a exposing is large, aperture cloacal The

9D) are regular, stellate with a few conical sharp rays rays sharp conical few a with stellate regular, are 9D)

Monniot Monniot

& &

Figs. 9E, 11 11 9E, Figs.

Monniot: New Ascidians Ascidians New Monniot:

C). C).

The intestinal loop is wide and contains contains and wide is loop intestinal The

new species species new

¼ ¼

the length of the trunk. trunk. the of length the 155 155

C, C,

D. D. misakiense

mm mm

Monniot, Monniot, has 1983

0.5 0.5

6 6

E E

0 0 "' "'

F. F.

with with a cloacal languet

of of another zooid;

(Sluiter, (Sluiter, have 1895) spicules

thorax thorax

ligulum ligulum

B, B,

D. D.

Didemnum Didemnum

zooid; zooid;

, ,

a a row.

29(2), 29(2), 1996

D. D. asteropum

in in

sp.: sp.:

n. n.

icronesica icronesica

Monniot, Monniot, has 1969 more stigmata a in row, strong

larva. larva.

. .

E, E,

linguiferum linguiferum

gut; gut;

Sluiter, Sluiter, and 1909

of of the

Didemnum Didemnum

details details

1. 1.

I I

D, D,

Didemnum Didemnum obscurum

D. D. chartaceum

Figure Figure

Remarks: Remarks: This species differs from other

The The spicules (Fig. 9E) are stellate, of a large up size to µm, 80 with numerous

scribed. scribed.

rap heal heal rap muscles, fewer epidermal ampullae in the larva and smaller spicules.

(Oka (Oka & Willey, 1892) has a different colony structure and the larva is not de

more more stigmata in a row, different and spicules a gemmiparous larva.

and and undivided testis in the following manner. From examination of the type spec

with with shorter rays and more stigmata

imens, imens,

pyramidal pyramidal rays.

of of stigmata can be seen.

(Fig. 1 lE). lE). (Fig. 1 The larva is not gemmiparous. Below the ocellus and otolith, 3 rows 156 156 Monniot & Monniot: New Ascidians 157

Didemnum mo/le (Herdman, 1886) Diplosomoides mo/le Herdman, 1886: 310, Indonesia. Didemnum mo/le: Kott, 1980: 2 and 1982: 98 and synonymy, lndo-Pacific; Millar, 1988: 829, Maldives Islands and Madagascar; Lafargue & Vasseur, 1989: 64, Ma yotte; F. Monniot, 1995: 316, fig. 10, New Caledonia. Material Examined: MNHN: A2 Did C 226-Indonesia, north Sulawesi, near Manado, l 0 23.50'N; 124°44.60'E, 10 m. Specimens from Seychelles and Vietnam are also present in the MNHN collections. The colonies always have the same urn shape with a terminal cloacal aperture and secrete abundant mucus. The color is highly variable corresponding to the respective proportions of the spicules, brown pigment cells and symbiotic algae. Larvae are present in the colonies from Sulawesi collected in May. Colonies also contained parasitic copepods and a pontoniine shrimp. Geographic distribution: The species is common from the Indian Ocean to the western Pacific, down to 50 m depth and more. It is generally found in well lit habitats.

Didemnum moseleyi (Herdman, 1886) Leptoclinum moseleyi Herdman 1886: 272, Philippines. Didemnum moseleyi: Kott, 1981: 169 and synonymy; Kott & Goodbody, 1982: 520, Hong Kong; Vasseur, 1985: 149, Polynesia; Monniot & Monniot, 1987a: 37, Polynesia; Lafargue & Vasseur, 1989: 64, Mayotte; Nishikawa, 1990: 100, Japan. Material Examined: MNHN: A2 Did C 308-Federated States of Micronesia, Chuuk Lagoon, NW Uman Reef, 7°18.50'N; 151°51.50'E, 25 m. The pink-red colonies are encrusting, hanging on vertical and overhanging surfaces, appearing to "drip" downward. They are several centimeters across and 1.5 mm thick. The pigment turns light brown in formalin and the colony is tan and white with dark spots made by the oral apertures. The consistency is hard, the colony is brittle, and breaks easily. There are two superimposed layers of tunic densely filled with spicules and separated by wide cloaca! channels around elon gated thoraces. The zooids have a wide funnel-like oral siphon with 6 triangular lobes. The cloacal opening is wide, without a languet. The lateral organs are small and lo cated at the postero-ventral corner of the thorax. The branchial sac is wider at the upper side with 6 stigmata in the first half row. The abdomen lies perpendicular to the thorax. The intestinal loop is narrow and curved. The testis is covered with 6 to 7 coils of the sperm duct. The larvae are small, with a length of 400 µm, and some are pigmented at the posterior part of the trunk. The spicules are stellate with numerous pointed lobes. In addition, much larger spicules with few rays are scattered throughout the colony. Geographic Distribution: The species is common in the Indo-Pacific.

concentrated concentrated in

C, C, abdomen;

; ;

is is

B, B, detail of the gut

mm mm

; ;

5 5

new new species

thorax

A, A,

: :

sp.

Figs. Figs. 9F, 12

n. n.

Micronesica 29(2), 1996 1996 29(2), Micronesica

Didemnum Didemnum nigrum

A2 A2 Did C 311-Philippines, Cebu Province, Olongo Island,

Didemnum Didemnum nigrum

larva. larva.

, ,

MNHN: MNHN:

Figure Figure 12.

Description: Description: The colony has a deep opaque black color in formalin. It is

small small round pigment spots.

the the substratum, the colony has a continuous thin layer of similar spicules, to the

surface surface layer. The black color of the tunic is diffuse but also

races. races. The abdomens of the zooids in lie the upper part of a thick vesicular and

from from the thick basal test channels by common wide surrounding cloaca} the tho

black tunic, totally devoid of spicules. The The larvae black Against present are tunic, this layer. totally in devoid spicules. of

siphons siphons and cloacal languets of the zooids. This hard surface test is separated

black superficial spicule-free layer layer above black spicule-free a superficial thin spicular layer containing the oral

folded on itself. The maximum length is 12 cm cm and mm. The the maximum maximum 3 on is folded thickness length itself. 12

their their margins without The colony irregular is spicules. in outline, parts with some

The The internal structure is peculiar. When sectioned, there is a very thin, soft and

158 158 tough, tough, rubbery; the surface is smooth. The common openings are cloaca} lobed,

off off Santa Rosa village, m. 30 10°15.00'N; 124°01.07'E, Holotype: Holotype:

described from Polynesia. The colony is thicker here, but is obviously in a resting resting a in is obviously but here, is thicker colony Polynesia. The from described

(Fig. 14A). 14A). (Fig.

papillae with very thin stalks, and 4 pairs of stout ectodermal ampullae. The 3 3 The ampullae. ectodermal stout of pairs 4 and stalks, thin very with papillae

in abundant mesenchymatous tissue. A single testis follicle is developed in only a a single follicle A testis only tissue. in is developed mesenchymatous abundant in

rows of stigmata are already differentiated. This larva is not gemmiparous. gemmiparous. is not larva This differentiated. already are stigmata of rows cloaca) aperture is narrow, without a languet. The lateral thoracic organs are are organs thoracic lateral The languet. a without narrow, is aperture cloaca)

µm in length and the tail makes makes tail the and length in µm

times. One larva was found in the deepest layer of the colony. The trunk is 500 500 is trunk The colony. the of layer deepest the in found was larva One times.

few zooids, in the primary gut loop. The sperm duct is empty and coiled several several coiled and empty is duct sperm The loop. gut primary the in few zooids,

ageal peduncle. The abdomen is almost spherical. The twisted gut loop is is loop twisted included gut spherical. The is almost abdomen ageal The peduncle.

the fourth row. The retractor muscle, of a medium size, extends from the esoph the from size, extends medium a of muscle, retractor The row. fourth the

Didemnum parau parau Didemnum

tains 6 stigmata in the first two rows, 5 stigmata in the third row, 4 stigmata in in stigmata 4 row, third the in stigmata 5 rows, two first the in stigmata 6 tains

small, located above the third transverse branchial sinus. The branchial sac con sac branchial The sinus. branchial transverse third the above located small,

resting state in the observed colony. The oral siphon has 6 pointed lobes. The The lobes. pointed 6 has siphon oral The colony. observed the in state resting

ible. It is pulled off in one sheet as the common cloaca) cavity is narrow. is narrow. cavity cloaca) common the as sheet one in off is pulled It ible.

and inflated under water. The test is entirely filled with spicules but remains flex remains filled spicules is entirely but test with The water. under inflated and

papillae on stout peduncles, circled by about 20 ectodermal vesicles. The larva is is vesicles. larva The 20 ectodermal about by circled peduncles, stout on papillae Islands, Makada Island, 4°06.28'S; 152°25.45'E, IO IO 152°25.45'E, 4°06.28'S; m. Island, Makada Islands,

Material Examined: Examined: Material

cated along the esophago-rectal waist. The abdomen (Figs. 12B, C) lies at a right right 12B, a lies (Figs. C) at abdomen The waist. esophago-rectal the along cated

muscular bundles follow the rapheal line (Fig. 12A). The retractor muscle is lo is muscle retractor 12A). The (Fig. line rapheal the follow bundles muscular

stigmata. The branchial sac is black or very dark purple. There are IO IO stigmata are There purple. dark very or black is sac branchial The stigmata. not gemmiparous. gemmiparous. not

lobes. The lateral thoracic organs are circular, protruding above the 3rd row of of row 3rd the above protruding circular, are organs thoracic lateral The lobes.

cloaca} apertures (Fig. 12A), with a narrow base and muscular fibers in the large large the fibers in muscular and base narrow a 12A), with (Fig. cloaca} apertures dark and opaque. The trunk measures 750 µm. There are 3 narrow adhesive adhesive narrow 3 are There µm. 750 measures trunk The opaque. and dark

(Fig. 12D) are incubated in the vesicular deepest part of the colony. They are very very are They colony. of the vesicular the deepest in part 12D) incubated are (Fig.

with 6 sharp lobes. The cloaca} aperture is wide, uncovering a large part of the the of part large a uncovering wide, is cloaca} aperture The lobes. sharp 6 with large and flat, covered by 6 to 7 turns of the sperm duct (Fig. 12C). The larvae larvae 12C). The (Fig. duct sperm the of turns 7 to 6 by covered flat, and large

lighter crescent is seen inside of the intestinal loop on the right side. The testis is is testis The side. right the on loop intestinal the of seen is inside crescent lighter

angle under the thorax. The body wall, digestive tract and gonads are black. A A black. are gonads and tract digestive wall, body The thorax. the under angle

ameter, and have many short rays. rays. short many have and ameter, in the first row, fewer in the others and the whole sac is triangular. Two strong strong Two triangular. is sac whole the and others the in fewer row, first the in

branchial sac. T-shaped languets are present in all zooids, larger near the common common the in near all larger present are zooids, languets sac. T-shaped branchial

The spicules are small 30 µm, globular, with crowded radial needle-like rays rays needle-like radial crowded with globular, 30 µm, small spicules are The

Remarks: This description fits very well the typical description of of description typical well the very fits description This Remarks:

The colonies (Pl. 2F) are encrusting, snowy white, rather irregular in outline outline in irregular rather snowy white, encrusting, are 2F) (Pl. colonies The

The zooids have the smaller thoraces than abdomens but seemed to be in a a in be to seemed but abdomens than thoraces smaller the have zooids The

h siue Fg F, itiue i n ae ae peia,4 m n di in µm 45 spherical, are layer one in distributed 9F), (Fig. spicules The

The zooids are difficult to extract from the tunic. The oral siphon is is tubular siphon oral The tunic. the from extract to difficult are zooids The

Monniot Monniot

Didemnum parau parau Didemnum

MNHN: MNHN:

Monniot & Monniot: New Ascidians Ascidians New Monniot: & Monniot

& &

Monniot, 1987a: 39, fig. 12 A-D, pl. II G G II pl. 12 fig. A-D, 1987a: 39, Monniot,

A2 Did C 309-Papua New Guinea, Duke of of York Duke Guinea, New 309-Papua C Did A2

Fig. 14A, Pl. 2F 2F 14A, Pl. Fig.

¾ ¾

Monniot Monniot

of a turn around it. There are 3 adhesive adhesive 3 are There it. around turn a of

& &

Monniot, 1987 1987 Monniot,

D. parau parau D. 159 159

. .

is is unknown. It is a

Monniot, Monniot, 1987a: 40,

& &

A2 A2 Did C 264--Palau, Ngerong,

new new species

Indonesia, Indonesia, north Bangka Sulawesi,

D. D. per/ucidum

F. F. Monniot, 1983

MNHN: MNHN:

m. m.

Fig. Fig. Pl. 13, 2G

1 1

Micronesica Micronesica 29(2), 1996

Figs. Figs. Pl. 14B, 15, 2H

A2 A2 Did C 228

F. F. Monniot, pl. 29, fig. IID, 12, 1983a: Guadeloupe; Mon

Didemnum poecilomorpha poecilomorpha Didemnum

MNHN: MNHN:

Didemnum Didemnum perlucidum

A2 A2 Did 230-Indonesia, C north ofTalisei side west Sulawesi,

Kott, Kott, 3-5, fig. 1982: 101 Palau, Philippines.

Laboute, Laboute, 486, 1985: Tahiti; Monniot

& &

Monniot,1995: Monniot,1995: A,B, fig.14 321, pl. IVE, New Caledonia.

. .

MNHN: MNHN:

51.52'N; 51.52'N; 125°03.84'£, 40 m.

44.77'N; 44.77'N; 125°09.26'£,

Description: Description: The colonies form large encrusting sheets, of 3 mm maximum

Geographic Geographic Distribution: The origin of

The The spicules have a regular stellate shape with few conical rays.

The The zooids have a long thorax, but the oral siphon is wide and short with 6

The colony The colony 2G) (Pl. large, encrusting, is thin but with The irregular swellings.

Geographic Geographic Distribution: Known only from Polynesia and Papua New

thickness, thickness, with a brown and white mottled surface, a design reminiscent of a thin

7°06.90'N; 7°06.90'N; 134°22.78'£, m. 12

Island, Island, l

Holotype: Holotype:

Didemnum Didemnum viride:

Senegal) as well as as as well Senegal) the Indian Ocean (Maldive Islands, Zanzibar), and the Pacific

(New (New Caledonia, Polynesia) and now Indonesia.

fouling fouling that species colonized many harbors in the Atlantic (Brazil, West Indies,

by by 4 long divergent ectodermal on vesicles each side. The 3 rows of stigmata are organs organs are horizontal, below the of level the fourth row of stigmata (Fig. 13A)

testis testis follicle is circled by 7 to 8 coils of the sperm duct (Fig. The 13B). larvae

differentiated differentiated in early stages of development of oozoids (Fig. 13C).

the the colony. The larvae are incubated in the thin basal test layer.

siphons. siphons. The zooids are inserted in groups in test pilars connecting both sides of measure measure about µm 500 in length. The separated well 3 adhesive papillae are lined

rather rather solid in spite of its brittle appearance due to the extension of the cloaca} sharp sharp lobes (Fig. The 13A). cloacal aperture is large exposing most of the bran muscle is thin thin muscle is and long (Fig. The gut 13A). loop forms a folded loop. The unique

The The stigmata are elongated, 6 to 8 were counted in the first row. The retractor

Island, Island, l

The The not anus covered is by the posterior part the of body The wall. lateral thoracic

channels. channels. The superficial layer of the test extends only to the level of the oral chial chial sac (Fig. The 13A). body wall covers only the oral area and the endostyle.

Material Material Examined:

niot, niot, Monniot cloacal cloacal apertures are wide and round. The zooids are crowded. The colony is

160 160

Polynesia; Polynesia; F

Didemnum Didemnum perlucidum

Guinea. Guinea.

similar. similar. The spicules are very characteristic. state, state, the zooids are not feeding. The single larva is somewhat larger but very

except just above the sensory vesicle, and in the front part above the adhesive adhesive the above part front the in vesicle, and sensory the above just except

she identified as as identified she

average diameter is 55 55 is µm diameter average

constriction (Fig. l 5C). There is no rastrum with symbiotic algae, and no buds. buds. no and algae, symbiotic with rastrum l is no There 5C). (Fig. constriction

papillae (Fig. 15G). With its 3 adhesive papillae and 6 pairs of digitiform ampul digitiform of pairs 6 and papillae adhesive 3 its 15G). With (Fig. papillae

680 µm in length, is coated with symbiotic unicellular algae all around the trunk, trunk, the around all algae unicellular symbiotic with is coated length, in 680 µm

short pointed rays. Their shape is very uniform throughout the colony and the the and colony the throughout uniform very is shape Their rays. pointed short phago-rectal waist is long and the retractor muscle becomes isolated at its pos its at isolated becomes muscle retractor the and long is waist phago-rectal

ois(i. E.Tebacil a a tgaa nte2frtrw.Te eso The rows. first 2 the in stigmata 6 has sac branchial l The 5E). (Fig. zooids

lae, the anterior part of the larva is separated from the oozooid body by a strong strong a by body oozooid the from is separated larva the of part anterior the lae, cloacal aperture is narrow (Figs. 15A, D, E). The thoracic organs are protruding, protruding, are organs thoracic The 15A, E). D, (Figs. is narrow aperture cloacal

ical above the mid intestine and covered by 6 to 9 coils of the sperm duct (Figs. (Figs. duct sperm the of coils 9 to 6 by covered and intestine mid the above ical

(Figs. 15B, F). A dark pigmented crescent lies inside the loop. The testis is is testis spher The lies loop. crescent inside the pigmented dark 15B, A (Figs. F).

terior part. The digestive loop is wide, not bent, with a small anterior stomach stomach anterior small a with bent, not wide, is loop digestive The part. terior

greenish and faded. The surface design is very similar to that of of that to similar is very design surface The faded. and greenish

veinous network, giving the specific name (Pl. 2H). In formalin the color turned turned color the formalin In 2H). (Pl. specific the name giving network, veinous

located above the third transverse sinus, close to the endostyle in fully fully in developed endostyle the close to sinus, transverse third the above located

300 µm for the abdomen. The oral aperture is tubular with 6 pointed lobes. The The lobes. pointed 6 with tubular is aperture oral The abdomen. the for 300 µm

level, in a dense spicular tunic. tunic. spicular dense level, a in

onies are tough as a result of the narrow cloacal canals, restricted to the thoracic thoracic the to restricted canals, cloacal narrow the of result a as tough onies are

ak re nlsosaesatrd ntedeetpr o h oois h col The colonies. the of part deepest the in scattered are inclusions green Dark

tatum tatum

15B, F). The larvae are very characteristic. They are almost spherical. The trunk, trunk, The spherical. almost are They very characteristic. are larvae 15B, The F).

The spicules (Fig. 14B) are crowded throughout the test. test. the The spicules 14B) They have numerous (Fig. throughout are crowded

The zooids are small, with a maximum length of 500 µm for the thorax and and thorax the for 500 µm of length maximum a with small, are zooids The

Remarks: Kott (1982) (1982) Philippines which and specimens Kott examined Palau from Remarks:

n. sp. Unicellular algae are embedded in the superficial layer of the test. test. the of layer superficial the in embedded are algae Unicellular sp. n.

Figure 13. 13. Figure

larva

. .

Didemm,m perlucidum perlucidum Didemm,m

Didemnum viride viride Didemnum

Monniot Monniot

. .

& &

(Herdman, 1906), however her specimens are are specimens her 1906), however (Herdman,

Monniot: New New Ascidians Monniot:

. ono, 1983 Monniot, F.

: :

A, thorax; B, abdomen; C, C, abdomen; B, thorax; A,

ienm gut Didemnum 161 161 162 Micronesica 29(2), 1996

Figure 14. Spicules. A, Didemnum parau Monniot & Monniot, 1987. B, Didemnum poecilomorpha n. sp. C, Didemnum rubeum n. sp. D, Didemnum stercoratum n. sp. E, Didemnum sp. A. F, Didemnum sp. B. Scale bars: lOµm.

Bootless Inlet, north of Motupore Island, 9°30.86'S; 147°16.63'E, 10 10 147°16.63'E, m. 9°30.86'S; Island, Motupore of north Inlet, Bootless

Material Examined: Examined: Material

Synonymy: see F. Monniot, 1995: 1995: Caledonia 324, New Monniot, see F. Synonymy:

quite different from Kott's specimens. specimens. Kott's from different quite

Museum, the type specimen of of specimen type the Museum,

similar to to similar

similar to the specimens of of specimens the to similar

Figure 15. 15. Figure

Didemnum poeci/omorpha. poeci/omorpha. Didemnum

in another colony; G, larva larva G, colony; another in

in the same colony (symbionts not figured); D, E, F, thoraces and abdomen abdomen and thoraces F, E, D, figured); not (symbionts colony same the in

Didemnum poecilomorpha poecilomorpha Didemnum

0,25 0,25

Didemnum psammathodes psammathodes Didemnum

MNHN: MNHN:

mm mm

Monniot Monniot

A2 Did C 267-Papua New Guinea, Port Moresby, Moresby, Port Guinea, New C 267-Papua A2 Did

. .

viride viride

D. D.

& &

Monniot: New New Ascidians Monniot:

viride viride

We have examined, on loan from the British British the from loan on examined, have We

that we examined from Polynesia, but are are but Polynesia, from examined we that

n. sp.: A, B, thorax and abdomen; C, larva larva C, abdomen; and thorax B, A, sp.: n.

rm Ceylon from

(Sluiter, 1895) 1895) (Sluiter,

. .

Its characters are very very are characters Its

0.25 0.25

mm mm 163 163

mm mm

0.5 0.5

. .

30.95'E, 30.95'E, m. IO

~ ~

( (

•· •·

larva

, ,

; ;

thorax

, ,

; ;

new new species

zooid

, ,

.: .:

. .

Micronesica Micronesica 29(2), 1996

um um Figs. Figs. Pl. 14C, 3A 16,

Didemnum Didemnum rubeum

A A

mnum mnum e rub

Did

. .

A2 A2 Did C 297 and 298- Palau, Rock Islands, Ngerkuul Pass,

Figure Figure

Description: Description: The living colonies are bright red or pinkish red, encrusting

MNHN: MNHN:

The The superficial layer of the tunic containing the unicellular symbiotic algae

Geographic Geographic Distribution: This fouling species is widely distributed in all

The The colonies are encrusting, grey, with pellets incorporated fecal in the tunic

151°40.66'E, IO m. m. 151°40.66'E, IO

and and the zooids turn themselves green in formalin. The zooids are large and per-

rocks rocks in large sheets reaching cm 20 across and 2 to 5 mm in thickness (Pl. 3A).

densely packed packed densely in especially the spicules, basal layer against the substratum. The 7°19.22'N; 7°19.22'N; 134°29.74'£, and Airai Channel, 7°21.03'N; 134

common common cloacal apertures. The colonies are white inside, resulting from the

Under Under water they are on inflated, with regular the surface with swellings terminal

cavity cavity extending at the of level the thoraces.

colonies colonies contract when touched. This is due to the very large common cloaca!

Holotype: Holotype: MNHN Did 296--Federated C :A2 of Micronesia, States Chuuk Atoll,

164 164

Anaw Anaw Channel (channel through barrier reef on North of Side atoll), 7°34.09'N;

warm warm of seas all oceans.

from from different oceans and the larvae are the same. and and few spicules. The zooids have the same morphology as in other specimens

uce r peet Fg 1B.Te spaorca wit sln ad a a re a has and long is waist esophago-rectal 17B). The (Fig. present are muscles

tractor muscle longer than the thorax. The abdomens lie horizontally under the the under horizontally lie abdomens The thorax. the than longer muscle tractor

long stigmata in the first two rows, 5 in the third, 4 in the fourth row. Rapheal Rapheal row. fourth the in 4 third, the in 5 rows, two first the in stigmata long

at the level of the third transverse sinus (Figs. 17 17 (Figs. sinus transverse third the of level the at

(Figs. 17 17 (Figs.

veloped around groups of thoraces. thoraces. of groups around veloped

aperture is large, without languet; it does not uncover the entire branchial sac sac branchial entire the uncover not does it languet; without large, is aperture

angle. The oral siphon is wide, long, with 6 triangular lobes (Fig. 17 17 lobes (Fig. is 6 with siphon wide, cloaca! long, triangular A). The oral The angle.

abundant fecal pellets (hence the species name). The cloaca! canals are well de well are canals cloaca! The name). species the (hence pellets fecal abundant

less numerous in the abdominal layer with a few in the basal test that contains contains that test basal the in few a with layer abdominal the in numerous less

in life life in

spicules are crowded in the upper layer of the colony containing the thoraces and and thoraces the containing colony the of layer upper the in crowded are spicules

across, around mangrove roots or on sea weeds. The color is cream, tan or yellow yellow or tan is cream, color The sea weeds. on or roots mangrove around across,

are located on each side of the furrows and sometimes on the elevated areas. The The areas. elevated the on sometimes and furrows the of side each on located are

into many polygonal areas by deep furrows as in a pavement. The oral apertures apertures oral The pavement. a in as furrows deep by areas polygonal many into

Bay, 10°18.71 'N; 124°00.72'E, 1 1 m. 10°18.71 124°00.72'E, Bay, 'N;

Holotype: Holotype:

the structure of the colony with common cloacal openings at the top of surface surface of top the at openings cloacal common with colony the of structure the

presence of reddish symbiotic algae and the deeply pigmented larva, as well as by by well as as larva, pigmented deeply the and algae symbiotic reddish of presence

swellings. swellings.

of crowded flat ended rays of equal length. The largest diameter is about 30 µm. µm. 30 about is diameter largest The length. equal of rays ended flat crowded of

is possible that the ectodermal ampullae divide later. later. divide ampullae ectodermal the that possible is

side, with an irregular outline. These larvae are not completely developed and it it and developed completely not are larvae These outline. irregular an with side,

The abdomen is not folded under the thorax (Fig. 16A). The digestive loop has has loop digestive 16A). The (Fig. thorax the under folded not is abdomen The

They are encircled by 6 coils of the sperm duct. The ovary is located on the side side the on located is ovary The duct. sperm the of coils 6 by encircled are They

to observe. They have 3 adhesive papillae and 4 ectodermal ampullae on each each on ampullae ectodermal 4 and papillae adhesive 3 have They observe. to

below the zooid level. They are deeply pigmented and their structure is difficult difficult is structure their and pigmented deeply are They level. zooid the below

of the testis. The larvae (Fig. 16C) are incubated in the dense layer of the tunic tunic the of layer dense the in incubated are 16C) (Fig. larvae The testis. the of

The lateral thoracic organs are near the rim of the cloaca! aperture between the the between aperture cloaca! the of rim the near are organs thoracic lateral The

stout (Fig. 16A). The branchial sac has 8 stigmata in the first row (Figs. 16A, 16A, B). (Figs. row first the in 8 stigmata has sac branchial 16A). The (Fig. stout are very close to each other (Fig. 16A), this is difficult to see without staining. staining. without see to difficult is this 16A), (Fig. other each to close very are

no secondary curve. The gonads are located on the gut loop. The two testis follicles testis follicles two The loop. gut the on located are gonads The curve. secondary no

of the upper part of the siphon in zooids encircling the common cloaca! apertures. apertures. cloaca! common the encircling in zooids siphon the of part upper the of

aperture is wide, without a languet (Figs. 16A, B), except for a small prolongment prolongment small a 16A, B), for except (Figs. languet a is without wide, aperture 3rd and 4th rows of stigmata (Figs. 16A, B). The retractor muscle is long and and long is muscle retractor The B). 16A, (Figs. stigmata of rows 4th and 3rd

pendicular to the colony surface. The oral siphon is short with 6 lobes. The cloaca! cloaca! The 6 lobes. with is short siphon oral The surface. colony the to pendicular

The zooids have a long thorax when relaxed and the abdomen is at a right right a at is abdomen the and relaxed when thorax long a have zooids The

Description: The colonies form thin crusts 2 mm thick and several several centimeters and 2 thick mm crusts thin form colonies The Description:

The spicules (Fig. 14C) of variable sizes appear spherical as they are made made are they as spherical appear sizes variable of 14C) (Fig. spicules The

This species differs from all other other all from differs species This

Dark pigment cells are scattered everywhere among the spicules in the tunic. tunic. the in spicules the among everywhere scattered cells are pigment Dark

(Pl. (Pl.

A, B). B). A,

3B), white in formalin. They are soft and flexible. The surface is is divided surface flexible. and The soft are They formalin. 3B), in white

MNHN: MNHN:

The lateral thoracic organs are protruding small round lamellae lamellae round small protruding are organs thoracic lateral The

A2 Did C 313-Philippines, Cebu, Mactan Island, Magellan Magellan Island, Mactan Cebu, 313-Philippines, C Did A2

Didemnum stercoratum stercoratum Didemnum

Monniot Monniot

Figs. 14D, 17, Pl. 3B 3B 17, Pl. 14D, Figs.

& &

Monniot: New Ascidians Ascidians New Monniot:

Didemnum Didemnum

new species species new

A,B). A,B).

species with two testes by the the by testes two with species

The branchial sac has 6 6 has sac branchial The 165 165

. .

larvae

mm mm

0.3 0.3

Didemnum Didemnum perlucidum

1996 1996

thoraces; thoraces; abdomen; C, D, E,

, ,

n. n. sp.:

Micronesica Micronesica 29(2)

8 8

,. ,.

il il

0 0

V, V,

stercoraftlm stercoraftlm

Didemnum Didemnum

igure igure 17.

Remarks: Remarks: Macroscopically, this resembles of species

The The are spicules stellate, made of numerous fused needles. Their diameter

the the gut, and the presence of abundant in pellets fecal the basal tunic.

characters characters including the spicule shape, the lateral thoracic organs, the shape of

F. Monniot, 1983, but is more opaque and colored. It differs by several anatomical anatomical several more by F. opaque but Monniot, and colored. is It differs 1983,

not not gemmiparous.

of of stigmata. The gut also partially is differentiated and horizontal. The larvae are

can can reach 40 µm (Fig. 14D).

Ocellus Ocellus and otolith are present above the branchial sac that already has 3 rows

pairs pairs of ectodermal ampullae, and separated from the oozooid by a constriction.

intestinal intestinal loop and covered with 5 to 7 coils of the sperm duct (Fig. 7C). The l

inal inal layer of the colony. They are small, 0.4 mm with 3 adhesive papillae and 4

ovary ovary against is the The testis. larvae (Figs. E) are 17D, incubated in the abdom

stomach, stomach, and the intestine loop is folded (Fig. 7C). The l single testis is in the

common common cloaca) cavity. The intestinal segments are not defined, well except the 166 166

are visible as white spots. The colonies are easily removed from the substrate the from removed easily are colonies The spots. white as visible are

rather small groups of spicules which give a spotted aspect. The oral apertures apertures oral The aspect. spotted a give which spicules of groups small rather

not have dense spicules, as are found throughout the remainder of the tunic, but but tunic, the of remainder the throughout found are as spicules, dense have not

formalin, but is inflated underwater. Under the microscope, the surface layer does does layer surface the microscope, the Under underwater. inflated is but formalin,

rud h cmo coc) prue (l 3C) (Pl. apertures cloaca) common the around

tan Island, underwater cave, l0°15.62'N; 123°59.11 'E, 10 10 m 123°59.11 'E, l0°15.62'N; cave, underwater Island, tan

aeil Examined Material

this is the case, it will be a new species. species. new a be will it case, the is this

out whether the thoracic structure described here is the normal adult one and if if and one adult normal the is here described structure thoracic the whether out

old thoraces and no abdominal buds. More material would be necessary to find find to necessary be would material More buds. abdominal no and thoraces old

el eaae ahsv ppla ad pis f coeml ampullae ectodermal of pairs 4 and papillae adhesive separated well that do not seem to feed. If the thoraces look like buds, there were no remaining remaining no were there buds, like look thoraces the If feed. to seem not do that

developed larva was observed, located in the basal part of the colony. It has 3, 3, has It colony. the of part basal the in located observed, was larva developed

µm (Fig. 14E), are spheres made of rod-like rays. rays. rod-like of made spheres are 14E), (Fig. µm

coils of the sperm duct. The ovary lies between the testis and the stomach. On the the On stomach. the and testis the between lies ovary The duct. sperm the of coils

gemmiparous. The thorax is well developed with 3 rows of stigmata. stigmata. of rows 3 with developed well is thorax The gemmiparous.

parts from the anterior part of the esophago-rectal waist. The gut forms a twisted, twisted, a forms gut The waist. esophago-rectal the of part anterior the from parts

produced into a short tube. The branchial sac has 10 oval stigmata in the first first the in stigmata oval 10 has sac branchial The tube. short a into produced

right side of the abdomen, in the gut loop, is a dark crescent. Only one well well one Only crescent. dark a is loop, gut the in abdomen, the of side right

empty and the thoraces seem to be aged buds. The oral siphon is short with only only with short is siphon oral The buds. aged be to seem thoraces the and empty

narrow, short loop. There is an undivided testis at the side of the gut circled by 7 7 by circled gut the of side the at testis undivided an is There loop. short narrow,

very hard. The common cloaca! channels are narrow, limited to the thoracic level. level. thoracic the to limited narrow, are channels cloaca! common The hard. very o o ah ie h ltrl hrcc ras eent seen not were organs thoracic lateral The side. each on row

could not be seen. The colony is brittle, densely filled with small spicules, but not not but spicules, small with filled densely brittle, is colony The seen. be not could

scalloped fringing lobes. The cloaca! siphon is circular, either sessile or sometimes sometimes or sessile either is circular, siphon cloaca! The lobes. fringing scalloped

Chuuk State, Kuop Atoll, 7°00. lO'N; 151 °56.20'E, 30 m 30 151 7°00. °56.20'E, lO'N; Atoll, Kuop State, Chuuk

black coral. The surface is smooth and shiny, oral or common cloacal apertures apertures cloacal common or oral shiny, and smooth is surface The coral. black

Material Examined: Examined: Material

species. species.

tributed throughout the colony. Its spicules are rare and different from the present present the from different and rare are spicules Its colony. the throughout tributed

This ascidian is encrusting, bright orange at the surface with white rings rings white with surface the at orange bright encrusting, is ascidian This

The spicules, small and crowded everywhere, with an average diameter of 40 40 of diameter average an with everywhere, crowded and small spicules, The

It is surprising to find in a rather large colony such numerous mature zooids zooids mature numerous such colony large rather a in find to surprising is It

The zooids are small, they are mature with testis and ovary, but the guts are are guts the but ovary, and testis with mature are they small, are zooids The

The colony is pink, several centimeters in length, and encrusting branches of of branches encrusting and length, in centimeters several pink, is colony The

Didemnum psammathodes psammathodes Didemnum

: :

MNHN: MNHN:

Monniot Monniot

2Dd 0-hlpie, eu suh at Mac east south Cebu, 303-Philippines, C Did A2

A2 Did C 305-Federated States of Micronesia, Micronesia, of States 305-Federated C Did A2

lo a audn fcl elt, u te ae dis are they but pellets, fecal abundant has also

Didemnum Didemnum

Fig. 14F, Pl. 3C 3C Pl. 14F, Fig.

& &

Monniot: New Ascidians Ascidians New Monniot:

Fig. 14E 14E Fig.

sp. A A sp.

. .

B B

. .

The colony is 1.5 mm thick in in thick mm 1.5 is colony The

. .

The retractor muscle muscle retractor The

. .

It is not not is It

. .

The The 167 167

new species species new

cm cm patches. The colorless but more

IO IO

Papua Papua New Guinea, Eastern Fields Atoll

Fig. Fig. Pl. 3D 18,

Micronesica Micronesica 29(2), 1996

Dip/osoma Dip/osoma multitestis

Description: Description: The translucent colonies appear blue underwater with white

The zooids are large, at least 2 mm mm for the at thorax are large, 2 The zooids least and l for abdomen, mm the

Remarks: Remarks: No described species allies all characters grouped here, such as

The spicules reach reach The µmin 40 spicules diameter, they are pointed long with stellate few

The The zooids are large. The oral siphon is wide, cylindrical, with 6 triangular

18A), sometimes 7 or or 7 sometimes The often 8, missing. 18A), tentacles are numerous, at in 24, least

rated rated by a constriction from the tubular rectum. The anus bilabiate is (Figs. 18B,

testine testine extends, after a constriction into an enlarged rectum without caeca sepa

D), D), posteriorly prolonged in a enlarged post-stomach. The olive shaped mid-in

fourth row of stigmata. There is a large anterior anterior large a stigmata. fourth There unperforated of is row branchial area the in gut gut loop is wide with a tubular esophagus, a cordiform stomach (Figs. C, 18B,

domen domen much is smaller than the thorax which is folded under it (Fig. The 18A).

3 3 orders of The size. cloacal siphon is narrow, circled by a ring of it muscles; is without without retractor muscle (Fig. The 18A). base of the thorax is straight. The ab

first row. The endostyle is narrow. narrow. is The endostyle row. first The esophago-rectal short is waist and narrow,

sac sac (Fig. The 18A). stigmata are long, numerous, with an average of 25 in the

located located either in the middle of the thorax (Fig. or at 18A) its base, in front of the

type type colony forms a cushion of 45 mm in diameter and 20 mm in thickness. It wall wall is totally transparent. At the rim of the oral siphon are 6 denticules (Fig.

white white in formalin. This is due to very dense micropapillae. The remaining body

this this size varying with the size of the gonads. The oral siphon remains opaque

has has openings. common 3 cloaca}

opaque opaque zooids are slightly The visible. consistency of the tunic jelly-like. The is

rings rings around the oral apertures (Pl. 3D). The colonies are large inflated, with few

(Coral (Coral km Sea), southwest 140 of Port Moresby, m. 20 145°39.90'E, 10°00.66'S;

Holotype Holotype MNHN: A2 Dip A 109

common common apertures. cloaca} They form

one one colony observed, the cannot species be named.

mata, mata, and esophago-rectal a long waist. But due to of larvae absence and the only

small small groups of in spicules the superficial spicular layer, a large number of stig

rays rays (Fig. 14F).

at at a right angle to the thorax. The primary loop of the intestine contains the

serted serted anteriorly along a very long esophago-rectal peduncle. The abdomen lies

large large oocyte. No mature larvae found were in the colony studied.

gonads: gonads: the of sperm with a coils duct testis 7 large and, aside, the ovary one with

168 168

the the 3rd transverse sinus. The branchial sac is wide with 8 to stigmata 10 in the

the the branchial whole The sac. lateral thoracic organs are protruding of at the level

thoracic thoracic level. first first row and 8 stigmata in the posterior rows. The retractor muscle is long, in

sharp sharp aperture lobes. The cloaca} is wide, with no languet, but does not uncover common common cavities are cloaca} wide and the colonies are easily split in two at the

colorless, with a large (2 mm) trunk mm) (2 large a with colorless,

mro dvlp nte aa lyr f h colony the of layer basal the in develop embryos

3 anterior adhesive papillae are well separated. Three long ectodermal papillae papillae ectodermal long Three separated. well are papillae adhesive anterior 3

D). When the testis is well developed, the ovary is small with few few small is located with oocytes, is ovary testis the well the developed, When D).

between the testis and the rectum. Only one large oocyte matures at a time. The The time. a at matures oocyte large one Only rectum. the and testis the between

wall. The testis follicles, from 5 to 7, are arranged in an irregular rosette (Figs. (Figs. rosette irregular an in arranged are 7, to 5 follicles, from testis The wall.

gonads are not located inside the gut loop but in a pouch of the abdominal body body abdominal the of pouch a in but loop gut the inside located not are gonads

connected to the stomach by a long duct crossing the gut loop (Fig. l l The 8E). (Fig. loop gut the crossing duct long a by stomach the to connected

C, D). The pyloric gland tubules converge to an ampulla against the intestine, intestine, the against ampulla an to converge tubules gland pyloric The D). C,

18B, C, D). The sperm duct is straight and opens against the anus (Figs anus the against opens and straight is duct sperm The 18B, D). C,

Figure 18. 18. Figure

of the gut; F, larva F, gut; the of

Diplosoma multitestis multitestis Diplosoma

Monniot Monniot

. .

& &

n. sp.: A, thorax; thorax; A, sp.: n.

Monniot: New A New Monniot:

. .

The tail is a quarter of the body circle. circle. The body the of quarter a is tail The

B, B,

. .

The tadpoles (Fig. l 8F) are are l 8F) (Fig. tadpoles The

C, D, abdomens; E, detail detail E, abdomens; D, C,

ans ans

. .

18B, 18B, C,

16 9 9

Herdman, Herdman,

(Sluiter, (Sluiter, 1909)

corals. corals. The color is

64; 64; F. Monniot, 1994:

: :

multifi.dum multifi.dum

Diplosoma Diplosoma viride

. .

Acropora Acropora

new new species

is is strongly adherent to the substrate

Diplosoma Diplosoma

(Sluiter, (Sluiter, 1909)

It It

Millar, Millar, 1975, which has a large cloaca}

(Hartmeyer, (Hartmeyer, 1909)

A2 A2 Dip A 103-lndonesia, north Sulawesi,

Micronesica Micronesica 29(2), 1996

The The species distributed is across all the Indian, and

A2 A2 Dip A 105-Palau, Koror State, Rock Islands,

: :

Figs. Figs. 19A, B, 20A, Pl. 3E

D. D. multifidum

Lissoclinum Lissoclinum hadium

Diplosoma Diplosoma similis

MNHN: MNHN:

Diplosoma Diplosoma virens

MNHN: MNHN:

synonymy: synonymy: Kott, 1980: 22 and synonymy; Kott, 1981: 193;

Hartmeyer, Hartmeyer, n. 1456, nov. 1909: for

had had mature colonies in July in Palau

124°58.87'E, 124°58.87'E, 8 m.

A2 A2 Lis A 107-Palau, Channel, 7°17.5'N; 134°27.8'E.

virens virens

also also differs from

. .

It It

Description: Description: The colonies are yellow and brown exteriorly, white internally

Geographic Geographic Distribution: This species is widespread in the tropical Indo Pa D

IO'N; IO'N;

Geographic Geographic Distribution

This This ascidian was found in soft thin crusts on

Remarks: Remarks: This new differs species from

in in the basal part. The surface has large swellings at the top of which are very wide

common common cloaca} apertures (Pl. 3E). The oral apertures are seen as dark brown

MNHN: MNHN:

(Coral (Coral Sea), km 140 southwest of Port Moresby, 10°01.47'S; 145°38.49'E, m. IO

10, New New 10, Caledonia Holotype: Holotype: MNHN: A2 Lis A 109-Papua New Guinea, Eastern Fields Atoll

1982: 114; Vasseur, Vasseur, 1982: 114; Lafargue 1985: 151; & Vasseur, 1989

1906. 1906.

saltwater saltwater river, 7°14.67'N; 134°24.43'E, Im. cific Ocean, Ocean, cific extending to Natal.

Material Material Examined:

Diplosoma Diplosoma virens:

Leptoclinum Leptoclinum virens

1°45. 1°45.

the the western and central Pacific Oceans.

species. species.

typically typically green with an intense blue glint.

and and brittle. It mostly inhabits shallow bottoms, less than meters IO deep. The

colonies colonies from Manado did not have larvae, but they are never frequent in this

Material Material Examined:

opening opening and a much smaller, gemmiparous larva.

by by the colony systems, the large number of stigmata and the absence of pigment

cells. cells.

species. species.

170 170

bearing bearing the papillae and another spherical posterior vesicle in which the tail is

inserted. inserted. The larva is not gemmiparous. There are no symbiotic algae in this

otolith. otolith. The branchial sac of the oozoid developed well is with 4 rows of stigmata.

are are folded backwards on each side. The sensory vesicle contains an ocellus and The The embryonic abdomen elongated is between a large anterior ball like structure,

from the tunic. The oral siphon is short with 6 lobes. The cloaca! aperture is is wide aperture cloaca! The lobes. 6 with is short siphon oral The tunic. the from

with crowded spicules. The cloaca! channels are wide and the zooids are arranged arranged are zooids the wide and are channels cloaca! The spicules. crowded with

hard basal part. The zooids are deeply pigmented dark brown. brown. dark pigmented deeply are zooids The part. basal hard

in groups in pillars of tunic. The larvae are incubated in the upper layer of the the of layer upper the in incubated are larvae The tunic. of pillars in groups in

spots. The upper part of the colony is soft and inflated, the basal layer is harder, harder, is layer basal the inflated, and soft is colony the of part upper The spots.

The zooids (Fig. 19A) are large (more than 2 mm) and very difficult to extract extract difficult to very 2 and mm) than 19A) (more large are (Fig. zooids The

Figure Figure

sp.: sp.:

19. 19.

C, zooid; D, gonads; E, thorax; F, G, larvae. larvae. G, F, thorax; E, gonads; D, zooid; C,

Lissoc/inum badium badium Lissoc/inum

Monniot Monniot

n. sp.: sp.: n.

& &

Monniot: Monniot:

A, A,

zooid; B, B, larva zooid;

New Ascidians Ascidians New

. .

ssoc/

inum nebulosum nebulosum inum

n. n.

1 71 71

n. n. sp.

sp. Scale bars: bars: sp. Scale IOµm.

. .

Lissoclinum Lissoclinum nebulosum

n. n. sp. B,

A trio/um trio/um A quadratum

D, D,

. .

Micronesica 29(2), 1996 29(2), Micronesica

Lissoclinum Lissoclinum badium

Lissoclinum Lissoclinum tunicatum

C, C,

Figure Figure 20. Spicules. A,

19A). The The 19A). gut is divided into several compartments, with a spherical stomach.

gemmiparous. gemmiparous.

5 5 to 9 adhesive papillae and the same number plus one ectodermal ampullae on

gut, gut, located vertically between 2 large (Fig. vesicles The 19B). larvae are not

each each side (Fig. The branchial 19B). sac already has 4 rows of stigmata, and the

straight. straight. The larvae are almost spherical, mm 1 for the trunk. There is a ring of

pigment cells. The esophago-rectal The esophago-rectal short, waist is without cells. pigment a retractor (Fig. muscle

The The testis against lies the gut loop with (Fig. 2 follicles The 19A). spermduct is

body body and wall the branchial tissue, as the abdomen, are invaded by large brown

organs organs are small near the of cloacal the edge aperture, to close The the endostyle.

ganglion. The The thorax ganglion. stigmata The with lateral row. first long the in thoracic 10 is 72 72

(Fig. (Fig. A 19A). very small languet is inserted at its upper rim, close to the neural 1

They consist of numerous needles; the length of the rays is not always equal. equal. always is not rays the of length the needles; numerous of consist They

from which emerges the tail. The larva is not gemmiparous. gemmiparous. is not larva The tail. the emerges which from

ularly spaced. The 4 rows of stigmata are already developed in the tadpoles and and tadpoles the in developed already are stigmata of rows 4 The spaced. ularly

number of these papillae varies in different larvae, from 18 to 20, and are irreg are and 20, 18 to from larvae, different in varies papillae these of number

the gut is elongated between the anterior yolky vesicle and a clear posterior vesicle vesicle yolky vesicle a clear posterior anterior and between the is gut the elongated

The tadpoles are large, up to 1.3 mm for the trunk. The 3 long adhesive papillae papillae adhesive 3 long The trunk. the for 1.3 mm to up large, are tadpoles The

rounded at their base by a ring of ectodermal short ampullae (Figs. 19F, G). The The 19F, G). (Figs. ampullae short ectodermal of ring a by base their at rounded

The eggs migrate posteriorly, to achieve their development isolated in the tunic, tunic, the in isolated development their achieve to posteriorly, eggs migrate The

r wdl eaae etnig rm lre ok vsce hy r ec sur each are They vesicle. yolky large a from extending separated widely are

the rectum. The ovary develops against the sperm duct, anteriorly to the testis. testis. the to anteriorly duct, sperm the against develops ovary The rectum. the

below the abdomen. The young embryos are rapidly enclosed enclosed rapidly are in embryos young The a spicular capsule. below abdomen. the

mature, the single testis is prominently protruding on the side of the gut loop loop gut the of side the on protruding prominently is testis single the mature,

with 12 stigmata in the first row. There is no muscular thoracic appendage. The The appendage. thoracic muscular no is There row. first the in 12 stigmata with

(Figs. l 9C, D). The sperm duct arises from the middle of the testis and follows follows and testis the of middle the from arises duct sperm The l D). 9C, (Figs.

The branchial sac extends down the entire length of the thorax (Figs. 19C, E) E) 19C, (Figs. thorax the of length entire the down extends sac branchial The

abdomen is folded under the thorax. It is flat in the absence of gonads. When When gonads. of absence the in flat is It thorax. the under folded is abdomen

organs form transverse hollows in front of the second transverse branchial sinus. sinus. branchial transverse second the of front in hollows transverse form organs

of the branchial sac. 24 oral tentacles are inserted on a ring. The lateral thoracic thoracic lateral The ring. a on inserted are tentacles oral 24 sac. branchial the of

fibrous and difficult to tear off. off. tear to difficult and fibrous

(Fig. 19E), but no sphincter. The cloaca! aperture is large and exposes exposes is part a large large and cloaca! The aperture sphincter. 19E), no but (Fig.

which lies perpendicular to it. The oral siphon has 6, very small pointed lobes lobes pointed small very 6, has siphon oral The it. to perpendicular lies which The zooids and larvae are orange in formalin, this color progressively fades. The The fades. progressively color this formalin, in orange are larvae and zooids The

common cloaca! openings are large at the junction of the wide cloaca! channels. channels. cloaca! wide the of junction the at large are openings cloaca! common

superficial layer of the colonies is jelly-like, the deepest parts of the tunic are are tunic the of parts deepest the jelly-like, is colonies the of layer superficial

marbled with white, giving a cloudy aspect (Pl. 3F) (suggested in the species species the in (suggested 3F) (Pl. aspect cloudy a giving white, with marbled

accumulation, either immediately below the colony surface or deeper in the tunic. tunic. the in deeper or surface colony below the immediately either accumulation,

name). The white patches, mostly around the zooids, are the result of a spicule spicule a of result the are zooids, the around mostly patches, white The name).

Spicules are in thick masses or only in capsules around the zooids and larvae. The The larvae. and zooids the in only around capsules masses Spicules or thick in are

ters across and about 4 mm thick. The color varies from grey to brown, irregularly irregularly brown, varies grey from to color 4 The thick. mm about and across ters

other specimen. specimen. other

goon, Polle Island, 7°21.54'N; 151°35.63'£, 10-30 m. m. 10-30 151°35.63'£, 7°21.54'N; Island, Polle goon,

Holotype: Holotype:

present species by the absence of a small cloaca! languet, the shape and the color color the and shape the languet, small cloaca! a of absence species the by present

merous adhesive papillae, and 2 testis follicles in the abdomen. It differs from the the from differs It testis follicles abdomen. 2 the in and papillae, adhesive merous

of the colonies. colonies. the of

the largest are 35 µm. This species has a raspberry smell, noted by the collectors. collectors. the by smell, noted raspberry a species has This 35 µm. are largest the

The spicules (Fig. 20B) are 55 µm in maximum diameter, all sizes mixed sizes all diameter, maximum in µm 55 are 20B) (Fig. spicules The

In the more relaxed zooids, the thorax is 2 times longer than the abdomen, abdomen, the than longer times 2 is thorax the zooids, relaxed more the In

Description: The colonies are soft, fleshy, encrusting sheets several centime several sheets fleshy, encrusting soft, are colonies The Description:

Remarks: Remarks:

The spicules are small balls of numerous radially arranged needles (Fig. 20), 20), needles (Fig. arranged radially numerous of balls small spicules are The

MNHN: MNHN:

Lissoclinumjaponicum Lissoclinumjaponicum

2LsA 1-eeae Sae fMcoei, hu La Chuuk Micronesia, of States 118-Federated A Lis A2

Lissoc/inum Lissoc/inum nehulosum

Monniot Monniot

Figs. 19C-G, 20B, Pl. 3F 3F Pl. 20B, 19C-G, Figs.

& &

Monniot: New Ascidians Ascidians New Monniot:

Tokioka, 1958 has similar larvae with nu with larvae similar 1958 has Tokioka,

new species species new

MNHN: MNHN:

2LsA 119 A Lis A2

173 173 . .

L. L.

Lis

larva. larva.

C, C,

differs differs from the white

It It

(Sluiter, 1909) also also has (Sluiter, spicular 1909)

Monniot, Monniot, Polynesia; 50, 1987a:

B, B, gonads;

& &

new new species

zooid; zooid;

A, A,

(Gottschaldt, (Gottschaldt, 1898)

Papua Papua New Guinea, Eastern Fields Atoll

sp.: sp.:

n. n.

A2 A2 Lis A 104, 108, 105-Palau: Ngerong,

Micronesica Micronesica 29(2), 1996

Figs. Figs. 20C, Pl. 21, 3G

1980: 18 and and 18 synonymy. 1980:

tunicawm tunicawm

Lissoclinum Lissoclinum triangulum

MNHN: MNHN:

Lissoc/inum Lissoc/inum tunicatum

Kott, Kott,

Lissoclinum Lissoclinum patella

Gottschaldt, Gottschaldt, 653. 1898:

Kott, Kott, Monniot 113; 1982:

Lissoclinum Lissoclinum

21. 21.

in in having an undivided testis, no atrial languet and no retractor muscle,

Monniot Monniot & Monniot, which has 1987, very peculiar spicules.

Figure Figure

Geographic Geographic Distribution: This species is very common in tropical areas of

The The type of specimen

Remarks: Remarks: This without new species, symbiotic algae, differs from other

(Coral (Coral Sea), km 140 southwest of Port Moresby, l0°00.66'S; m. 145°39.90'E, 72

Holotype: Holotype: MNHN: A2 Lis A 121

the the Indian and Pacific oceans. It has previously been recorded from the Palau

F. F. Monniot,1992: New 572, Caledonia.

Islands, Islands, Arch Big Lake, 7°15.8l'E; 134°24.9l'E, m. 1

Islands Islands by Kott 1982). (

7°06.90'N; 7°06.90'N; 134°22.78'E, m; 10 Ngercheu, 7°05.39'N; 134°16.24'E, 0.5 m; Rock

Material Material Examined:

Lissoclinum Lissoclinum patella:

Lissoclinum Lissoclinum patellum:

Didemnoides Didemnoides patella

cells that that cells do not exist in the studied species here.

174 174

tadpoles tadpoles have fewer ectodermal ampullae and a body wall with large vesicular

taratara taratara

of of the spicules. Nevertheless, the shape of the gut segments is different, and the

capsules capsules around the zooids and larvae, and a more angular but similar structure

a a larva that not is gemmiparous and a different arrangment of the larval papillae.

The The structure of the colony is also characteristic. soclinum soclinum Monniot & Monniot: New Ascidians 175

Description: The colonies are jelly-like, brown, surrounding octocorallian axes. The white spots visible in transparency are due to spicule capsules around the zooids (Pl. 3G). The cloacal canals are wide, the common cloacal apertures are not visible on preserved material but in living colonies are seen to be large bordered with a transparent chimney of tunic. The zooids are very contracted. They are well separated from each other in the colony, each enclosed in a layer of tunic containing small spicules. The oral siphon is tubular with 6 very short lobes. The cloacal aperture is bound by a muscular ring; it is sometimes large when not contracted but appears narrow after fixation (Fig. 21A). The lateral thoracic organs are small and difficult to see, making a horizontal slit above the second transverse branchial sinus. There is no retractor muscle. The number of stigmata in a row cannot be counted but they are numerous. The esophageal peduncle is short (Fig. 21A) and the abdomen is folded under the thorax. The stomach is heart-shaped and the different segments of the intestine are well individualized. The undivided testis is on the gut loop (Fig. 21B); spherical it may reach a very large size, as large as the entire abdomen. The sperm duct is straight. The ovary lies against the sperm duct in the gut loop (Fig. 21B). The larvae (Fig. 21C) are incubated in the tunic, near the parent zooids. They are enclosed in a spicular capsule. They measure about 1 mm. They have 3 well separated adhesive vesicles, with a crown of short ectodermal ampullae at the base of their stalk. An average of 20 ectodermal ampullae were counted in the tadpoles. The larvae are gemmiparous (Fig. 21C). The spicules (Fig. 20C) have rays of irregular length, they are about 20 µm in diameter. Remarks: This species has many similarities to other Lissoc/inum species with jelly-like colonies, few spicules and a gemmiparous larva. However, L. abdominale F. Monniot, 1982, L. calycis F. Monniot, 1992, L. mereti Monniot & Monniot, 1987, L. punctatum F. Monniot, 1992, L. verrilli (Van Name, 1902) have different spicules, no brown pigment and 4 pairs of larval ectodermal ampullae. L. pacifi cense Kott, 1981 also has different spicules and 6 pairs of larval ectodermal am pullae.

Lissoclinum voe/tzkowi ? (Michaelsen 1920) Didemnum voeltzkowi Michaelsen 1920: 54 Lissoc/inum voeltzkowi: Kott, 1980: 13; 1981: 190; 1982: 112; F. Monniot, 1992: 579. Material Examined: MNHN: A2 LIS A 102, and 103-Palau: Ngerong, sea grass, 7°06.90'N; 134°22.78'E, 0.5 m; S.W. Airai Channel, Ngermutidech, 7°18.74'N; 134°3l.12'E, intertidal. The colonies are soft, encrusting corals or weeds on sea grass beds. When living the color is grey and green. The color is irregular, a result of the relative number of spicules and brown pigment cells, and the abundance of the symbiotic unicellular algae. The surface can be smooth or wrinkled, with some papillae

L. L.

Monniot, Monniot,

& &

(Sluiter, (Sluiter, 1905)

voeltzkowi, voeltzkowi,

L. L.

bistratum bistratum

L. L.

(Sluiter, (Sluiter, 1909)

The Palau specimens have fewer rays rays and The fewer have Palau specimens

Pl. Pl. 3H

Micronesica Micronesica 29(2), 1996

bistratum. bistratum.

L. L.

Leptoc/inides Leptoc/inides dubius

Sluiter, pl. Sluiter, 69, 3, pl. 4, fig. Indonesia. 7, 1909: fig. 10,

F. F. and Monniot, Monniot synonymy; 681 1989:

nor nor to those of

26.57'N; 26.57'N; 124°44.48'£, m. 23

The The characters of the zooids with agree the previous descriptions of this spe

The colonies are encrusting but fleshy, 5 to 8 cm across across cm to and are thick. Their encrusting cm The colonies 5 8 but fleshy, 1

Geographic Geographic Distribution: Madagascar, Australia, Philippines, Cal New Fiji,

Remarks: Remarks: Without ovaries and larvae, the identification of these colonies The The spicules of the Palau specimens are not exactly similar to those of

The The branchial sac has 8 stigmata in each half row. The abdomen is much

The The cloacal cavities are large, surrounding groups of zooids and extending

1991: 377, New New 377, Caledonia. 1991:

in in a rosette) and only by circled one complete of coil the sperm duct. Large sized

larvae larvae in lie the deepest part of the colony.

lular lular symbiotic algae. No symbiotic algae were found in the interior. The large

cies; two two tubular grouped in bunch a cies; (not siphons follicles with the arranged testis

color color pink is (Pl. 3H) but they may be partially covered with bright unicel green

nado, nado, l The The are spicules only numerous in the basal part of the colony .

Material Material examined: MNHN: A2 Lep 31-Indonesia, of Ma north west Sulawesi, common common cloacal openings expose the brown-red internal color of the colonies.

Leptoc/inides Leptoc/inides dubius:

Polysyncraton Polysyncraton dubium

voeltzkowi, voeltzkowi,

they they are thicker here. However, variability some has been from observed different

edonia, edonia, Palau.

species based based species on this difference. single

is is a allied and species, closely it may be possible that additional occur in species geographical geographical areas, so it is doubtful the Palau could specimens represent a new

the the western Pacific.

scribed scribed from Australia, New Caledonia and Fiji. But

remains remains doubtful. The color and colony shape correspond to

observed, observed, and the zooids totally were immature. No larvae were present.

follicle and and parallel follicle to the rectum. The not ovaries were developed in the colonies

below below the intestine. The sperm duct originates posteriorly, runs over the testis

smaller smaller than the thorax. The gut loop is narrow and the single testis is follicle

176 176

level of of level the second transverse There sinus. no is retractor muscle.

wide wide with no languet. The lateral thoracic organs are circular protruding at the

has has no lobes, its rim plain is or raised slighly in two lips. The cloacal aperture is

sizes. The The zooids open apertures. at sizes. colony the surface by slit-like The oral siphon

tunic, tunic, crowded with spicules. Spicules are numerous everywhere, with variable

below below the abdominal layer. Against the substrate a is dense, rather thick layer of

located located mostly at the thick edge of the colony. Generally the consistency is firm

mm mm in thickness. with with a slippery surface. The colonies reach centimeters several across and 2 to 5 the western Pacific; Australia, Korea, Philippines, Malaysia, Indonesia. The spe The Indonesia. Malaysia, Philippines, Korea, Pacific;western Australia, the cies was collected down to 80 m depth.80 m to cies collecteddownwas

Holotype: ° .4N 125°21.84'E, m; 25 2°l 7.24'N; MNHN: omn laa aetrs asd n o selnso te ooy ufc. The surface. colony the of swellings low on raised apertures cloacal common

green, blue-black or grey, mottled orange, with darker grey in patcheswithsome darker orange, grey,orgreen, places blue-black mottled (Pl. 4A). The internal color was grey was color internal The 4A). (Pl. hcns vre fo 2 o m Te pclsae etitd o h ts layer test the to restricted are spicules The mm. 7 to 2 from varies thickness °75N 42.', 10 m. 34°27.8'E, 7°17.5'N; h etr ts i atclry og ad iru, n i pt fter ag size large their of spite in and fibrous, and tough particularly is test entire The (1

surrounding the oral siphons. They are mixed with black ramified pigment cells. pigment ramified black with mixed are They siphons. oral the surrounding

3°15', odr hne, °07N 142.', ih Hue Channel, House Light 134°22.l'E, 7°10.7'N; Channel, Wonder 134°31.55'E,

5 themm),very zooids are difficultdamage. remove the colonytowithout from

egahc itiuin Te pce a iegorpi itiuin in distribution geographic wide a species has The Distribution: Geographic

ecito: hs pce a cletd n rcy otm n ws dark was and bottom rocky a on collected was species This Description:

Fig

o.s

2 e 4 ad 8Plu Mtedu Brir ef 7°16.51 'N; Reef, Barrier Mutremdiu, 48-Palau, and 47 Lep A2

re 22 re

MNHN:

l mm

arva.

Leptocli

2 e 3-noei, ot Slws, un Island, Ruang Sulawesi, north 32-Indonesia, Lep A2

etciie oscitans Leptoc/inides

ides oscita

Monniot & Monniot: New Ascidians New Monniot: & Monniot

Figs. 22, 24A, Pl. 4A Pl. 24A,22, Figs.

. sp. .

The smooth surface is pierced by rounded by is pierced surface smooth The

zooid;

new speciesnew

tho

ax; C, ax;

, ab ,

dom

ens; E, ens;

177 is divided

Four rows

Leptoclinides

Leptoclinides sulawesii

There is no retractor muscle.

B).

(Sluiter, 1909)

22A,

(Sluiter, A, larva 1909): of the type colony; B,

Micronesica 29(2), 1996 29(2), Micronesica

Figs. 23, Pl. 24B,4B

Hastings, 92, 1931: Great Barrier Reef and New Zea

Sluiter, 60-Indonesia. 1909:

Leptoclinides reticulatus

Leptoc/inides reticu/atus

larva of another colony

Sluiter, whose 1909 type has tubular cloacal siphons, rounded instead of

Figure 23.

The oral and cloacal apertures have smooth without edges The cloacal lobes.

Remarks: raysThe are more spicule numerous than in

The (Fig. spicules 24A) are large, reaching µmin90 diameter.

178

aperture not is tubular, evencontractedin animals, but (Fig. and simpleis sessile

same level on each side, but above the second branchial sinus on the left and above the third sinus on the right (Figs 850 µm. The 3 short stout adhesive papillae are diverging, and alternate with 4 of stigmata are already well developed in the tadpole, behind the ocellus and 22A) similar to similarname.22A) The yawn,a thoracicthe henceare species grouped muscles thorax (Fig. 22A) with a narrow gut loop. The testis (Figs. 22C, in 6 longitudinal bundles on each The side. thoracic organs are not located at the into 3 to 5 follicles encircled by 4 to 6 coils of the sperm duct. The body wall contains black fusiform pigment cells. The trunk of the larvae measures 800 to

There are at least stigmata 14 in the first row. The abdomen is smaller than the pairs ectodermal of papillae with an enlarged triangular tip (Fig. 22E) n.sp., but with the same chisel-shaped tips. The differs from species rufum otolith pigmentramified cells, and a different larva.

Leptoclinides reticulatus: land; Tokioka, 89, Palau 1967: and Gilbert Islands. Material Examined: MNHN: A2 Lep 34-Papua New Guinea, Duke of York Islands, Mioko Island, 4°13.85'S;m 23 152°28.16'E, Didemnum reticulatum white. The superficial layer of the colony is transparent and soft. Under this pel this Under soft. and is transparent colony the of superficial layer The white.

edii pget el.Ti lyrcno esl e eaae fo te bottom the from separated be easily cannot layer This cells. pigment dendritic part of the colony. The cloacal channelsencirclingcloacal of The the colony. clustersof zooids below extend part

hc wt a irglr uln. h upr ufc i lc, ih odn rings golden with black, is surface upper The outline. irregular an with thick rud h oa oeig ad ht rns n h rm f h cmo cloacal common the of rim the on rings white and openings oral the around apertures at the top of low swellings (Pl. 4B). The underside of the colonies is colonies the of underside The 4B). swellings (Pl. low of top the at apertures

licle, the tunic is tough, and difficult to tear. It is spiculesdenselyfilled and with It tear. difficult to and tough, is tunic licle, the

the zooid abdomens but are not very wide. very not are but abdomens zooid the

tubular, and without lobes. The cloacal aperture is located in front of the fourth the of front in is located aperture cloacal The lobes. without and tubular, row of stigmata, either sessile or on a short tube. The thorax is wide, with a wide is wide, a with thorax The tube. short a sessile on either or stigmata, of row

The colonies are encrusting on rocks, several centimeters across, l to 2 cm 2 to l across, centimeters several rocks, on encrusting are colonies The

The zooids are difficult to remove from the tunic. The oral siphon is long, is siphon oral The tunic. the from remove to difficult are zooids The

iue 4 Siue. A, Spicules. 24. Figure

Sutr 1909). C, (Sluiter,

cl as 1 bars: Scale Oµm.

Monniot

etciie sulawesii Leptoclinides

etciie oscitans Leptoclinides

Monniot: New Ascidians New Monniot:

n. sp. n.

. sp. n.

etciie uniorbis Leptoclinides

etciie reticulatus Leptoclinides

n. sp. n.

9 n. sp.

Leptoc/inides sulawesii

n. sp., which has darkhas n. sp., which pigment

new species

species, the species, zooids are extremely dif

mm in length). muscular Six bundles ex

L. oscitans

Micronesica 29(2), 1996

Leptoc/inides

Figs. 24C, 25A-C, Pl. 4C

Leptoclinides sulawesii

A2 Lep 30-lndonesia, north Sulawesi, west of Manado,

MNHN:

A2 Lep 8 and 29-Palau, Wonder Channel, 7°10.7'N; 134°22.l'E, 12

Remarks: This species differs from differs Remarks: This species

The spicules (Fig. 24B) are stellate, with pointed rays. This species has similar are pointed with stellate, rays. speciesThis24B) (Fig. The spicules Geographic Distribution: Indonesia, Palau, Gilbert Islands, Papua New

Description: Large encrusting sheets that reach more than 20 cm in length

The thorax is elongated (at least

23.52'N; 124°32.64'£, m. 38

180

otolith stigmata presentof rows are that branchial above the 4 with alreadyare sac

straight base. The stigmata could not be counted. The lateral thoracic organs are round above the middle of the fourth row of stigmata. There is no retractor surface design. The basal layer of the colony was much thicker in our sample, slightly twisted. The twisted. stomach and slightly small is folliclesThere spherical. testis to are4 3

below the zooids. They are rather small (450 µm) with 3 adhesive papillae, 2 the apertures. The larvae and are spicules different. lacks black pigment has cells, a wide gut loop, more and testis follicles different developed. spicules.

characters as those of the type specimens which has been examined. It corresponds It examined. has been which characters thosetype the specimensas of Guinea, New Zealand. closely to the Tokioka's the to but from Palau,closely had are smaller similara which specimens muscle. The abdomen is folded under the thorax. The gut loop is narrow and possibly due to the presence of larvae at this stage of development. encircled by 5 coils of the sperm duct. The larvae (Figs. 23A, B) are incubated cells and cells no tube cloaca} because of the peculiar golden and white rings around ectodermal vesicles on each side, one dorsal and one ventral. The ocellus and MNHN:

and 5 mm in thickness. In life, they are rusty orange (Pl. 4C) and translucent internally. The pigment quickly fades in alcohol, and becomes dark red in for

of tunic devoid of spicules. The colonies are not shiny but the superficial layer

Holotype: the deepest fibrous layer in contact with the substrate. The larvae are incubated in the basal test. As in many m, and Mutremdiu, Barrier Reef, 7°16.5l'N; m. 134°31.55'E, 12 tend longitudinally Both and cylindrical onsiphons are eachwithout side. fringing lobes (Fig.lobes 25A). The oral siphon has a strong sphincter. The cloaca]tube arises malin. Numerousapertures protrude cloaca} at colonythe a bylined surface, ring of the from third the level row of stigmata. The thoracic organs are located above the third branchial sinus (Fig. 25A). An unperforated area separates the pre does not contain and spicules a is deep orange color. There are no black pigment cells. The cells. spicules are mostly restricted to the zooid level, and become sparse in

ficult to ficult extract from the common test where they are crowded.

pharyngeal groove from the first row of stigmata. About long 14 stigmata were

is no bud. bud. no is

ectodermal ampullae (Fig. 25C). The 4 rows of stigmata are clearly visible, there there visible, clearly are stigmata of rows 4 The 25C). (Fig. ampullae ectodermal

does not form caeca, it is not clearly separated from the mid-intestine. The gonads gonads The mid-intestine. the from separated clearly is it not caeca, form not does

a large stomach (Fig. 25B), an annular post-stomach. The rectum is enlarged but but is enlarged rectum The post-stomach. annular an 25B), (Fig. stomach large a

25A). There is no retractor muscle. The gut loop is wide with a short esophagus, esophagus, short a with wide is loop gut The muscle. retractor no is There 25A).

counted in the first row. The esophago-rectal peduncle is short and narrow (Fig. (Fig. narrow and short is peduncle esophago-rectal The row. first the in counted tips. Their diameter is about 45 µm, rarely 50 50 µm. rarely µm, 45 about is diameter Their tips.

the sperm duct (Fig. 25A). The larval size is variable with an average length of of length average an with variable is size larval The 25A). (Fig. duct sperm the

separate zooids. Three to 5 testis follicles are in a circle surrounded by 7 coils of of coils 7 by surrounded circle a in follicles are testis 5 to Three zooids. separate

are normally located on the side of the gut loop. Testis and ovary were found in in found were ovary and Testis loop. gut the of side the on located normally are

850 µm for the trunk. The 3 adhesive papillae are sided by only 3 pairs of wide wide of pairs 3 only by sided are papillae adhesive 3 The trunk. the for 850 µm

The spicules few spicules with 24C) stellate, The have chisel-shaped are rays (Fig. The rays.

Figure 25. 25. Figure

clinides uniorbis uniorbis clinides

Leptoclinides sulawesii sulawesii Leptoclinides

n. sp.: D, thorax; E, F, abdomens. abdomens. F, E, thorax; D, sp.: n.

Monniot Monniot

& &

Monniot

n. sp.: A, zooid; B, abdomen; C, larva. larva. C, abdomen; B, zooid; A, sp.: n.

: :

New New Ascidians

Lepto 181 181

L. L.

(Sluiter, (Sluiter,

Leptoc/in

(Sluiter, (Sluiter, 1909)

Polysyncraton Polysyncraton doboense,

species species from Fiji under the

Leptoc/inides Leptoc/inides rufum

new new species

Leptoc/inides Leptoc/inides cuspidatum

Polysyncraton Polysyncraton

Federated Federated States of Micronesia, Chuuk Lagoon,

Sluiter, Sluiter, which 1913, is completely different from

Micronesica Micronesica 29(2), 1996

Figs. Figs. 24D, 25D-F, Pl. 4D

Leptoclinides Leptoclinides uniorbis

A2 A2 Lep 35

(Sluiter, (Sluiter, described 1913) under the name

MNHN: MNHN:

does does not correspond to any New Caledonian (F. Monniot species 1989)

Polysyncraton Polysyncraton doboense

n. n. sp. by the cloacal aperture, the the and spicules, pigment cells the larva.

The The zooids have long tubular oral siphons ending denticules in (Fig. 6 25D).

Description: Description: The colonies are across cm large orange and sheets, several to 2

Kott Kott (1981) redescribed a true

For For comparison, the type and paratype of the Indonesian species

Remarks: Remarks: This differs species from

duct duct only one coil describes and runs along the rectum (Figs. 25E, F). The ovary

made made of 6 to arranged follicles 8 in a compact rosette (Figs. 25E, F). The sperm

is is short. The loop digestive is wide with differentiated well regions. The testis is

small, and and small, located at the base of the siphon. cloaca! There no is retractor muscle.

to to count the stigmata 18 in the first row. The lateral thoracic organs are circular,

smooth. When fixed in formalin the red pigment dissolves and the fixative become become formalin and in the fixative the dissolves pigment smooth. red When fixed musculature is weak and and weak musculature reduced very the easy thus is (Fig. 25D), cloacal cavity is it

the the abdomens. The colonies are easy to collect as the underside is wrinkled but

The The abdomen is not always folded under the thorax. The esophago-rectal waist

The cloaca} siphon is a tube at the level of the fourth fourth the of stigmata. of The row thoracic tube a at level the siphon is cloaca} The

layer is pierced with large meandering meandering large with penetrate pierced also which canals the is tunic layer below The The thorax is wide and square with a distinct angle at the end of the endostyle.

thin thin sheets extends the zooid layer, with but fewer, still numerous This spicules.

substrate, substrate, the test forms a crust with very crowded Between spicules. these two deeply-colored. deeply-colored.

open open at the top of swellings (Pl. 4D). The zooids are irregularly spaced. The

3 mm mm thick, with 3 an irregular outline. The round large common apertures cloaca!

colonies colonies are made of several layers. On the upper surface layer and against the

Lematol Lematol Bay Polle and between Pata Islands, 7°21.45'N; m. 10 151°35,53'E,

Sluiter's Sluiter's sample. Holotype: Holotype:

name name

longitudinal longitudinal thoracic on There muscles each are side. stigmata in the 10 first row.

1909) has has many 1909) stigmata fewer and dark pigment remaining cells in fixative.

small but but small not differentiated. completely

ides ides doboense numerous numerous in the fibrous very test; they are large and stellate. The zooids have 8

grooves grooves in a polygonal design. The layer superficial of the colony contains the

There There are and 3 generally testis follicles of coils 6 the sperm duct. The larvae are

182 182

zooids, zooids, separated from the basal layer by the cloaca! canals. The are spicules not

citans citans

colonies colonies are encrusting, about 10 cm across. The cloaca! canals form surface

deposited deposited in the Senckenberg Museum at Frankfurt have been examined. The

or or to other Indo-Pacific type that species we have studied. It differs from

which which has 4 pairs of larval ectodermal ampullae. sulawesii sulawesii Monniot & Monniot: New Ascidians 183 develops against the testis with one large oocyte maturing at a time. The larvae were not seen. The spicules have a maximum diameter of 40 µm with numerous rays (Fig. 24D) with tips in points or cut. Remarks: This species differs from Leptoc/inides dubius (Sluiter, 1909) by the very regular distribution of the testis follicles in a rosette, and the gut being a simple flat loop. It differs from Leptoc/inides multilobatus Kott, 1954 which has 20 testis lobes, a very large stomach and spicules at the colony surface only.

Atriolum quadratum new species Figs. 20D, 26, Pl. 4E Holotype: MNHN: A2 Atr 9- Federated States of Micronesia, Chuuk Atoll, bar rier reef, channel (Salat Pass) on the south east side of the atoll, 8 m.

Figur e 26. A trio/um quadratum n; sp.: A, zooid ; 8, detail of the gut; C, D, abdo mens; E, growth of the brood pouch. 184 Micronesica 29(2), 1996

Description: The colonies form large encrusting sheets, 2 to 3 mm thick, with surface swellings each with a terminal circular common cloacal opening (Pl. 4E). The color is grey or pale violet-blue. The colonies are inflated under water. The general consistency is flexible but tough. The spicules are crowded in the super ficial layer of the colony and more sparse inside. The zooids are crowded. The abdomens are folded under the thoraces. The oral siphon is tubular with 6 shallow lobes. The oral tentacles, at the base of the siphon, are very numerous, in 4 orders of size. There is a large imperforate area between the prepharyngeal band and the branchial sac (Fig. 26A). There are 12 stigmata in each of the 4 rows. An imperforate area lies along each side of the dorsal line. The 3 rapheal languets are slightly displaced to the left side. The cloacal siphon is tubular, very long, at the base of the thorax. It has an internal velum at its tip (Fig. 26A). The small cup-like thoracic organs are placed on the begining of the cloacal siphon, above the esophageal opening. The thoracic mus culature is weak, longitudinal. A brood pouch, small and empty in the figure (Fig. 26E), is located at the base of the cloacal siphon on the right side of the thorax. The esophagorectal waist is short and begins far from the base of the endostyle (Fig. 26A). The shape of the thorax (siphons excluded) is rectangular, the abdo men is located at its postero-dorsal angle. There is no retractor muscle. The ab domen is much smaller than the thorax (Fig. 26A). The esophagus is tubular, the stomach spherical without plications. There is a cylindrical post-stomach, sepa rated from the intestine by a constriction. The mid-intestine is tubular, a little wider than the posterior intestine when empty. A distinct bend is located between the mid-intestine and the posterior intestine (Figs. 26B, C). The testis is made of 3 to 5 follicles, located above the mid-intestine, surrounded by 4 coils of the sperm duct (Fig. 26D). The ovary, with one large oocyte, takes the place of the testis when the latter has disappeared (Fig. 26C). No embryos were found in the brood pouch. The majority of the zooids of the large colony studied were in a male phase, although a few had an ovary and the start of a brood pouch. The spicules are large, the largest reach 100 µm, with few long pointed rays (Fig. 20D). Remarks: This species differs from Atriolum robustum Kott, 1983 in having several testis follicles. It differs from Atriolum marsupialis F. Monniot, 1989 in its different spicules, thorax and cloacal siphon. The shape of the gut is also different. The esophago-rectal waist is shorter here, and displaced dorsally. Even without larvae, A. quadratum is obviously different from the other two species of A trio/um. The species name comes from the square shape of the thorax.

Family: Polycitoridae Po/ycitor circes Michaelse~, 1930 Polycitor circes Michaelsen, 1930: 495, fig.2; Kott, 1990a: 169 and synonymy. Material Examined: MNHN: A3 Pol A 17-Indonesia, north Sulawesi, mouth of Teluk Korakora inlet, l 0 43.97'N; 124°58.03'E, 33 m.

vascular processes. The stomach is large, asymmetrical; the intestine forms a loop loop a forms intestine the is asymmetrical; large, stomach processes. The vascular

The oral tentacles are very numerous in several circles. In addition the anterior anterior the addition In circles. several in numerous very are tentacles oral The

are very short. The abdomen is very elongated, and ends in one or several several ends in or is one ramified and very elongated, abdomen The very are short.

and its ascending limb is bent against the stomach (Figs. 27 A, B). The rectum rectum 27 The B). (Figs. A, stomach the against bent is limb ascending its and

internal part of the oral siphon is covered with minute papillae. The branchial sac sac branchial The papillae. is minute with covered siphon oral of the part internal

fibers regularly spaced. This strong musculature extends onto the long abdomen. abdomen. long the onto extends musculature strong This spaced. fibers regularly

has long stigmata, about 25 on each side in the first row. The rapheal languets languets rapheal The row. first the in side each on 25 about stigmata, long has

The body wall is thick with very numerous longitudinal and transverse muscle muscle transverse and longitudinal numerous very with thick is wall body The

and greenish oocytes; the larvae are colorless. Both siphons have 6 pointed lobes. lobes. 6 pointed have siphons Both colorless. are larvae the oocytes; greenish and

cloacal openings in the center. The tissues are colorless except the orange stomach stomach tissues colorless The orange are except the center. in the openings cloacal

surface to the substrate. They are arranged with the oral openings in circles circles with in openings oral the with arranged are They substrate. the to surface

embedded sand. The color varies from dark green to tan or marroon. There is a a is There marroon. or tan to green dark from varies color The sand. embedded

the colony surface and between the zooids. zooids. the between and surface colony the

(Pl. 4F). Many green or brown spheres of varied sizes are found in the tunic at at tunic the in found sizes are varied of spheres brown or green Many 4F). (Pl.

star or honeycomb pattern at the surface due to the zooid arrangement in circles circles in arrangement zooid the to due surface the at pattern honeycomb or star

ippines, Cebu Province, Olongo Island, off Santa Rosa village, l0°14.99'N; l0°14.99'N; village, Rosa Santa off Island, Olongo Province, Cebu ippines,

ions l cm thick, and up to 9 cm across with a rubbery but smooth surface, with with surface, smooth but rubbery a with across cm 9 to up and thick, cm l ions

Papua New Guinea, New Ireland, Kalili Harbor, 3°26.37'N; 151°56.0l'E;-Phil• 3°26.37'N; Harbor, Kalili Ireland, New Guinea, New Papua

diu, Barrier Reef, 7°16.51 'N; 134°31.55'E; West reef 7°l9.4'N; 134°l3.9'E, l m; l 134°l3.9'E, 7°16.51 Reef, Barrier 134°31.55'E; diu, reef 7°l9.4'N; West 'N;

;Wne Canl Rc Ilns72.0N l40.5E itria; Mutrem intertidal; l34°02.15'E, Islands,7°22.30'N; Rock Channel, Wonder m;

°33';115.5E . n ol ince °l5'; 15l 151°52.25'E, 7°2l.54'N; Polle Pinnacle, 7°23.30'N; 0.5 and m

7°16.80'N; 134°25.92'E, 0.5 m; Venture marine lake, 7°19.21 'N; 134°29.33'E, 1 1 7°19.21 134°29.33'E, lake, 'N; marine Venture m; 0.5 134°25.92'E, 7°16.80'N;

Federated States of Micronesia, Chuuk lagoon, north side of Dublon Island, Island, Dublon of side north lagoon, Chuuk Micronesia, of States Federated

doab, Ngersuul, 7°24.51 'N; 134°35.89'E, 0.5 m; Ngeruktabl marine lake, lake, marine Ngeruktabl m; 0.5 134°35.89'E, 7°24.51 'N; Ngersuul, doab,

Material Examined: Examined: Material

Eudistoma amp/um: amp/um: Eudistoma

Polycitor amp/us amp/us Polycitor

Australia and the Philippines. Philippines. the and Australia

l24°0l.07'E, 9 9 m l24°0l.07'E,

licles and a central ovary. The larvae are large large are larvae The ovary. central a licles and

common base. Zooids open on the upper surface in circular systems. The test is is test The systems. circular in surface upper the on open Zooids base. common

longitudinal plications. The gonads lie in the gut loop with numerous testis fol testis numerous with loop gut the in lie gonads The plications. longitudinal

bryos are incubated all along the oviduct. oviduct. the along all incubated are bryos

anterior adhesive papillae triradially arranged, and no ectodermal vesicles. vesicles. Em ectodermal no and arranged, triradially papillae adhesive anterior

translucent and firm. The oral and cloacal siphons have 6 lobes. The branchial branchial The lobes. 6 have siphons cloacal and oral The firm. and translucent

sac has at least 15 rows of stigmata. The stomach wall has numerous narrow narrow numerous has wall stomach The stigmata. of rows 15 least at has sac

0,ad 0-aa: ia Canl 7l.4N l43.8E l ; at Babel east m; l l34°3l.28'E, 7°l9.24'N; Channel, Airai 105-Palau: 104, and

The zooids (Fig. 27 A) may reach l cm in length, they extend from the colony colony the from they extend length, 27 in cm (Fig. l zooids The reach A) may

The colonies from all localities are similar. They are circular encrusting cush encrusting circular are similar. all They localities are colonies from The

Geographic Distribution: This species is recorded from the western Pacific, Pacific, western the from recorded species is This Distribution: Geographic

Several lobes reaching 6 cm in height and 3 cm in diameter arise from a a from arise diameter in cm 3 and height in cm 6 reaching lobes Several

. .

Sluiter, 1909: 1909: 21. Sluiter,

Kott, 1990a: 194 and synonymy. synonymy. 194 1990a: and Kott,

MNHN: MNHN:

Eudistoma amp/um amp/um Eudistoma

Monniot & Monniot: New Ascidians Ascidians New Monniot: & Monniot

A3 Eud 77, 78, 84, 86, 96, 97, 98, 100, 102, 103, 103, 102, 100, 98, 97, 96, 86, 84, 78, 77, Eud A3

Fig. 27, Pl. 4F 4F 27, Pl. Fig.

(Sluiter, 1909) 1909) (Sluiter,

, ,

the trunk 2.5 mm long, with 3 3 with long, mm 2.5 trunk the

35.63'E, l m; l 35.63'E, 185 185

; ;

larva

, ,

; ;

C C

The The central. ovary is

abdomens

C). C).

, ,

3 3

"' "'

, ,

; ;

ooid

, z ,

29(2), 29(2), 1996

1909): 1909):

, ,

(Sluiter

B B

Micronesica Micronesica

papillae. papillae.

amp/11m amp/11m

larval larval

Eudistoma Eudistoma

. .

E, E, detail of

0 0

Figure Figure 27

Geographic Geographic Distribution: The species was already known from Palau, the

Banda Banda Sea, Kei Islands, Truk, Ponape and Majuro Islands and Queensland in

Australia. Australia.

(Fig. (Fig. 27D). There are papillae, adhesive irregularly 3 spaced. The young tadpoles

large large concentric rows of irregular papillae on each anteriorly side, (Fig. 27E).

have have only between odd the adhesive papillae, swellings the mature larvae have 3

facing the the facing stomach also extends anteriorly and posteriorly onto the The intestine.

One One or two larvae are incubated in the cloacal cavity. The trunk is µm 900 long

are are numerous, a in bunch the gut inside loop (Figs. 27B,

pyloric pyloric duct enters the stomach junction at the of the intestine. The follicles testis

starts starts behind the posterior loop by an annular enlargment. The pyloric gland 186 186

the tunic but yellow-green spherical plant cells are present. The consistency is is consistency The present. are cells plant spherical yellow-green but tunic the

of the colony has a short stalk arising from a common base. There is no sand in in sand no is There base. common a from arising stalk short a has colony the of

form balls 6 to 8 cm in diameter around living branches of of living branches around diameter in cm 8 to 6 balls form

rier reef), 7°16.16'N; 134°31.74'E, 134°31.74'E, 18 m. reef), 7°16.16'N; rier

Holotype: Holotype:

Description: The colonies are made of several several to of which heads coalescent made colonies are The join Description:

Figure 28. 28. Figure

MNHN: MNHN:

, ,

larva

Eudistomafasciculum Eudistomafasciculum

. .

Eudistoma gilboviride gilboviride Eudistoma

A3 Eud 83-Palau, Koror State, Mutremdiu lagoon (on bar (on lagoon Mutremdiu State, Koror 83-Palau, Eud A3

Eudistoma fasciculum fasciculum Eudistoma

Monniot Monniot

0.5 0.5

Fig. 28A-C, Pl. 4G 4G Pl. 28A-C, Fig.

mm mm

& &

. .

Monniot

Sutr 1909) (Sluiter,

.: .:

A, zooid; B, posterior part of the abdomen; abdomen; the of part B, posterior zooid; A,

: :

New Ascidians Ascidians New

new species species new

: :

D, E, zooids. zooids. E, D,

Acropora. Acropora.

Each lobe lobe Each 187 187

(Sluiter, (Sluiter,

has has different

Eudistoma Eudistoma laysani

E. E. o/ivaceum

. .

(Sluiter, (Sluiter, 1909)

(Van (Van Name, The structure 1902). the of colony is

Micronesica Micronesica 29(2), 1996

Figs. Figs. 28D, E, Pl. 4H

has has smaller zooids, smaller larvae, weak abdominal

Kott,1990a: Kott,1990a: 206, Queensland, Australia.

Eudistoma Eudistoma gilboviride

Sluiter, 1909: 6, 6, Indonesia Sluiter, 1909:

E. E. laysani

Eudistoma Eudistoma o/ivaceum

The The zooids (Figs. 28 D, E) have 6 lobes at each cylindrical siphon. The tho

The The are colonies stalked, with an average of 2 cm in height, with the zooids

Remarks: This species is a similar in appearance appearance in to similar a is species Remarks: This

The The zooids are up long, to mm 8 in length contracted for the less They ones.

the the abdomen. The branchial sac has 3 rows of long stigmata. The abdomen is

racic racic musculature is made of many thin fibers, extending in thin ribbons along

stomach stomach and dark pigment cells around the most mature larvae, the eggs are

the the colony where they open; they are not grouped in systems.

hard hard with dark pigment The cells. zooids are not pigmented except the orange

yellow. The The yellow. zooids are parallel to each other, perpendicular to the flat surface of

opening opening on the flat top of the lobes. The general color is dark green, but large

1900) and and 1900)

Rasch Rasch 3 m. Passage, 5°09.27'S; 145°49.82'E,

pale pale irregular yellow patches are on seen the upper surface (Pl. 4H). The tunic is

Material Material Examined: MNHN: A3 Eud 88 and 101-Palau, Koror State, Light

house house Channel, 7°17.03'N; 134°27.82'E; m.-Papua 12 New Guinea, Madang,

Eudistoma Eudistoma gilboviride:

Polycitor Polycitor gilboviridis

the the same. The zooids in our have species a more developed musculature and a

pigment pigment spots which remain and in fixatives, a smaller and different larva.

musculature musculature and lacks thoracic dark pigment spots.

thicker thicker body wall.

into into pointed lobes (Fig. The 28C). larval trunk mm 0.8 is in length.

to to the The left side. 3 adhesive papillae have thin and long stalks. They are sep

arated arated by 4 pairs of ectodermal ampullae, irregular in shape, their tip produced

of the tadpole is not exactly encircling the the the anterior of encircling not tadpole papillae is exactly displaced a but little is

Among Among 4 or embryos 5 in a zooid, only one maturity reaches in a time. The tail

oocytes oocytes (Fig. 28B). The larvae (Fig. 28C) are incubated in the cloacal cavity.

the the upper part of the stomach has 2 humps (Fig. The 28A). gut loop and wide is

an an irregular outline; the typhlosole is raised in a ridge and on the opposite side

from from the posterior extremity of the zooids. The orange stomach posterior is with

contains very numerous testis follicles (Fig. 28A). The ovary is central with several several central with The ovary is (Fig. 28A). follicles contains numerous testis very

188 188

rows rows oflong stigmata the entire branchial occupy without sac unperforated areas.

have have posterior abdominal vascular processes. Both siphons are short A). (Fig. 28

consists consists of numerous longitudinal and transverse fibers regularly crossed. The 3

disappear disappear in The fixatives. 6 cloacal lobes are round. The thoracic musculature

The abdomen is contracted contracted The strong distance muscular abdomen 2 ending at bundles, some is by

The The oral siphon has 6 triangular lobes. At its base are 4 brown spots, which

slightly protruding protruding slightly at the upper surface of the lobes. No observed. were systems

The The opaque tunic more becomes in specimens living transparent after fixation. tough. tough. The color is brown-orange, with green (Pl. 4G). The siphonal lobes are

wall and the larvae. All attempts to clear this pigment remained unsuccessful. unsuccessful. remained pigment this clear to attempts All larvae. the and wall

unperforate space is anterior to the rows of stigmata stigmata of . rows the to anterior is space unperforate

shape of the gut, different ectodermal ampullae in the larvae (Fig. 29D). A large large A 29D). (Fig. larvae the in ampullae ectodermal different gut, the of shape

obvious. obvious.

dermal ampullae seen as small vesicles extending in more than the anterior half half anterior the than more in vesicles extending small as seen ampullae dermal

cavity, rarely two. They are highly pigmented, not well developed and their struc their well and developed not pigmented, highly are They two. rarely cavity,

fueul egh ltee ltrly Fg 2C.Tee r eynmru ecto numerous very are There 29C). (Fig. laterally flattened length, unequal of

The tail forms only half a circle. The 3 adhesive papillae, in the median line, are are line, median the in papillae, adhesive 3 The circle. a half only forms tail The

of the trunk, and along the ventral side (Fig. 29C). 29C). (Fig. side ventral the along and trunk, the of

ture is particularly difficult to study. They strongly contract when dehydrated. dehydrated. when contract strongly They study. to difficult particularly is ture

mid-intestine. The ovary is central. One larva is incubated in the thoracic cloacal cloacal thoracic the in is incubated larva One is central. ovary The mid-intestine.

nads are located in the gut loop, behind the stomach (Figs. 29A, B). The small small The B). 29A, (Figs. stomach the behind loop, gut the in located are nads

culature consists of numerous longitudinal fibers in a continuous coating. The The coating. continuous a in fibers longitudinal numerous of consists culature

hrx egh h aymtia soah svlmnu ad otro. h go The posterior. and voluminous is stomach asymmetrical The length. thorax

more than 30 long stigmata. The less contracted abdomens reach 2.5 times the the times 2.5 reach abdomens contracted less The stigmata. long 30 than more

testis follicles are so numerous that they sometimes extend in a pouch below the the below pouch a in extend sometimes they that numerous so follicles testis are

branchial sac has no anterior unperforated area (Fig. 29A) and the first row has has row first the and 29A) (Fig. area unperforated anterior no has sac branchial

1954b whose type specimen has been examined, however some differences are are differences some however examined, been has specimen type whose 1954b

qa eghad ae6rudd oe rud h prue. h hrcc mus thoracic The apertures. the around lobes rounded 6 have and length equal

ranged. ranged.

is tough but gelatinous internally. The zooids are well spaced and vertically ar vertically and spaced well are zooids The internally. gelatinous but tough is

no systems are visible visible (Pl. are systems no

Holotype: Holotype:

marbled with yellow is very characteristic. characteristic. is yellow very with marbled

mentions colonies with zooids in small systems. The green color of the colony colony the of color green The systems. small in zooids with colonies mentions

nesia, Queensland, and now in Palau and New Guinea. Guinea. New and Palau in now and Queensland, nesia,

correspond well to the previous descriptions with the exception that Kott (1990a) (1990a) Kott that exception the with descriptions previous the well to correspond

650 to 770 770 µm. 650 to

wide loop. The testis follicles are not very numerous and do not fill the posterior fill posterior the not do and numerous very not follicles testis are The loop. wide

124°32.62'£, 43 m. m. 43 124°32.62'£, stalks and are bordered with 4 pairs of ectodermal papillae. Their trunk measures measures trunk Their papillae. of ectodermal 4 with pairs bordered are and stalks

being incubated in the thoracic cloacal cavity. The 3 adhesive papillae have short short have papillae 3 adhesive The cavity. cloacal thoracic the in incubated being

gut loop. The ovary is located alongside the testis follicles. testis follicles. the larva is one alongside is only There located ovary The loop. gut

long, enlarged in the posterior part where the post-stomach and intestine form a a form intestine and post-stomach the where part posterior the in enlarged long,

. .

The The

Remarks: Externally this species resembles resembles species this Externally Remarks:

Almost all zooids of the colony are contracted. Both siphons are tubular, of of tubular, are siphons Both contracted. are colony the of zooids all Almost

Description: Completely encrusted with sand, the colony is a brown color, color, brown a is colony the sand, with encrusted Completely Description:

Geographic Distribution: Distribution: Geographic

Remarks: These characters and especially the color and shape of the colonies colonies of the shape and especially color the and characters These Remarks:

E. snakabri snakabri E.

pigment in in pigment

MNHN: MNHN:

3Ed7-noei, ot Slws,Mnd, l Sulawesi, Manado, north 76-lndonesia, Eud A3

has protruding test papillae at the colony surface, a different different a surface, colony the at papillae test protruding has

E. incrustatum incrustatum E.

Eudistoma incrustatum incrustatum Eudistoma

SA). SA).

Monniot Monniot

Figs. 29A-C, Pl. Pl. 29A-C, Figs.

Eudistoma gilboviride gilboviride Eudistoma

Both test and zooids are dark brown. The colony colony The brown. dark are zooids and test Both

& &

Monniot: New Ascidians Ascidians New Monniot:

is darker, much more abundant in the body body the in abundant more much darker, is

new species species new

SA SA

Eudistoma snakabri snakabri Eudistoma

has been recorded in Indo in recorded been has

Tokioka, Tokioka,

23.56'N; 23.56'N; 189 189

snakabri snakabri

mm mm

.5 .5

Eudistoma Eudistoma

. .

larva

, ,

; C ;

1996 1996

zooid

(Sluiter, (Sluiter, 1900)

B, B,

A, A,

.: .:

n. n.

A3 A3 Eud 89-Papua New Guinea, Madang,

. .

Micronesica Micronesica 29(2),

larva

, ,

incrustatum incrustatum

Eudistoma Eudistoma laysani?

MNHN: MNHN:

1954: 1954:

, ,

Eudistoma Eudistoma

. .

Tokioka

Figure Figure 29

The colonies are are The colonies translucent white heads with short stalks, crowded together,

5°08.44'S; 5°08.44'S; 145°48.IS'E, 2 m on mangrove roots.

Material Material Examined: uncertain. uncertain.

not not mature fully and without larvae, identification the remains species somewhat

forming forming balls on mangrove roots. The diameter reaches 2 cm. As the zooids are 190 190

ovary is central. Generally, two thin and long conical stolonic vessels inde- vessels arise stolonic conical long and thin two Generally, central. is ovary

round testis follicles form a dense mass posterior to the stomach (Fig. 30A). The The 30A). (Fig. stomach the to posterior mass dense a follicles testis form round

gut loop, close to the beginning of the ascending limb of the intestine. Numerous Numerous intestine. the of limb ascending the of beginning the close to loop, gut

side of the stomach. A large clear pyloric vesicle is sometimes located inside the the inside located vesicle sometimes is pyloric clear large A stomach. the of side

limb of the intestine is enlarged in two caeca two in is enlarged intestine the of limb

tubules have some swellings near the short common duct entering the posterior posterior the entering duct common short the swellings near some have tubules

The zooids seem to have two stomach pouches. The first part of the ascending ascending the of part first The pouches. stomach two have to seem zooids The

the tubules extending down the rectum to the level of the stomach (Fig. 30F). The The 30F). (Fig. level stomach the of the to rectum the down extending tubules the

is inflated in a ball, sometimes more voluminous than the stomach (Figs. 30E, F) 30E, (Figs. stomach the than voluminous more sometimes ball, a is in inflated

groove (Fig. 30F). After a short and narrow cylindrical segment, the post-stomach post-stomach cylindrical segment, the narrow and short a 30F). After (Fig. groove

stomach is asymmetrical with 2 dorsal swellings separated by a longitudinal longitudinal a by separated swellings dorsal 2 with asymmetrical is stomach

is long is

in the first row. The dorsal languets are displaced on the left side. The abdomen abdomen The side. left the on displaced are languets dorsal The row. first the in

aa (Fig mata

muscles which form strong ribbons along the abdomen. The body wall wall body The is abdomen. thick. the along The muscles ribbons strong which form

separate them from the branchial sac. The branchial sac has 3 rows of long stig long of rows 3 has sac branchial The sac. branchial the from them separate

lobes.The body wall has numerous and regularly spaced thoracic longitudinal longitudinal thoracic spaced regularly and numerous has wall body lobes.The

oral tentacles, in 3 orders of size, are inserted on a line. A large unperforated area area unperforated large line. a A on size, of inserted are 3 orders in tentacles, oral

oral one with 6 sharp rounded lobes, the cloaca! siphon with, sometimes, 8 8 sometimes, with, siphon cloaca! the lobes, rounded sharp 6 with one oral

are calcium oxalate. Several kinds of pigment cells cells circular of Several kinds pigment dark coexist test: calcium the in oxalate. are

sions, hardly soluble in acetic acid, polarising the light, which may mean that they they that mean which may light, the acetic acid, soluble polarising in sions, hardly

ones and others more clear and fusiform. fusiform. and clear more others and ones

depressions contain a ring of oral openings and a central concavity with cloaca! cloaca! with concavity central a and openings oral of ring a contain depressions

zooids systems are obvious at the upper surface of the colonies, where circular circular where colonies, the of surface upper the at obvious are systems zooids

apertures. The tunic is vacuolar and contains numerous crystalline surface inclu surface crystalline numerous contains and vacuolar is tunic The apertures.

crusted particles on the surface and inside the test. In formalin, the color turns turns color the formalin, In test. the inside and surface the on particles crusted

dark brown. The consistency is rubbery and the colonies are hard to tear. The The tear. to hard are colonies the and rubbery is consistency The brown. dark

thickness. They are brown or dark purple (Pl. 5C, D), with sometimes few en few sometimes with D), 5C, (Pl. purple dark or brown are They thickness.

7°19.4'N; 134°29.6'E; marine lake north of Ngel Channel entrance, 7°18.3'N; 7°18.3'N; entrance, Channel Ngel of north lake marine 134°29.6'E; 7°19.4'N;

o Sae Rc Ilns Tpua, °59'; 134°27 7°15.94'N; Topkukau, Islands, Rock State, ror

0mPlu es aeda, grul 72.1'; 3°58',itria; Ko 7°24.51 intertidal; 134°35.89'E, Ngersuul, 'N; Babeldoab, east m-Palau, 60

uaei aao l Manado, Sulawesi,

Material Examined: MNHN: A3 Eud 73,74,75,81,85 and 107-lndonesia, north north 107-lndonesia, 73,74,75,81,85 and Eud A3 MNHN: Examined: Material

Eudistoma reginum reginum Eudistoma

etr toia Pcfc aai Plu Piipns Nw aeoi ad Aus and Caledonia New Philippines, Palau, Hawaii, Pacific: tropical western

134°29.l'E, 9 m m 9 .. 134°29.l'E,

tralia. tralia.

All zooids are shortened by contraction by shortened are zooids All

The colonies form low cushions, measuring up to 20 cm across and 1 cm in in 1 cm and across cm 20 to up measuring cushions, low form colonies The

Geographic Distribution: Distribution: Geographic

. .

h u opi eycaatrsi (Figs characteristic is very loop gut The

. .

30A), the first one dorsally curved dorsally one first the 30A),

Kott, 1990: 1990: Queensland,Australia. 228, Kott,

23.52'N; 124°32.64'E, 18 m, and 1°45.IO'N; 124°58.87'E, 124°58.87'E, 1°45.IO'N; and 18 m, 124°32.64'E, 23.52'N;

Eudistoma reginum reginum Eudistoma

Monniot & Monniot: New Ascidians Ascidians New Monniot: & Monniot

Eudistoma /aysani /aysani Eudistoma

Fig. 30, Pl. 5C, D D 5C, Pl. 30, Fig.

. .

The pyloric gland is is well gland pyloric developed, The

Kott, 1990 1990 Kott,

. .

There are about 20 to 25 stigmata stigmata 25 to 20 about are There

Both siphons are cylindrical, the the cylindrical, are siphons Both

. .

30A, B, E, F). The short brown brown short The F). E, B, 30A,

has a wide distribution in the the in distribution wide a has

01 'E, 1 m; shallow Bay, Bay, shallow 1 m; 01 'E,

191 191 . .

e e

larva

, ,

d. d.

mm mm

D). D).

; ;

0.5 0.5

" " a a

"' "'

Pal

m m

pyloric pyloric glan

fro

e e

s s

ooid

l of of l th

, , z

1996 1996

r~ r~

, ,

: : I I

\ \

I I

1 1

.I .I

j j

sometimes sometimes branched, some alternate with

esi; esi;

1990

) )

, ,

) )

, ,

) )

\ \

Sulaw

ronesica 29(2) ronesica

Kott

m m

1111111 1111111

d d fro

egi

a a

E, E, zo

; ;

isto

ala

30. 30.

from from P

gure gure

A A

2 2

side side of the tail attachment, two lateral are swellings also pigmented. The ecto

3 stout adhesive papillae (Figs. 30C, D) deeply pigmented in dark dark pigmented in D) deeply brown. papillae 30C, stout (Figs. On adhesive 3 each

the the papillae, adhesive others are lined on each (Figs. side 30C,

dermal dermal anterior papillae are variable

matures matures in cavity. The the thoracic tadpole cloaca} large. is It characterized is by

pendently pendently from the enlarged posterior part of the abdomen. Only one embryo 19

ot Ffn Island Fefan north

Holotype: Holotype:

the samples collected at Palau and Indonesia. Indonesia. and Palau at collected samples the

deposited in the Queensland Museum, were examined. They are very similar to to similar very are They examined. were Museum, Queensland the in deposited

egahc Distribution Geographic

Remarks: specimens from Heron Island (Australia), identified by Kott and and Kott by identified (Australia), Island Heron from specimens Remarks:

Figure 31. 31. Figure

A A

MNHN

larva

. .

uitm rubigino Eudistoma

: :

, ,

A3 Eud 112-Federated States of Micronesia, Chuuk lagoon, lagoon, Chuuk of Micronesia, States 112-Federated Eud A3

Eudistoma rubiginosum rubiginosum Eudistoma

30'N; 151 151 30'N;

Monniot Monniot

3 3

9 9

: :

uesad Idnsa Palau Indonesia, Queensland,

Fig. 31, Pl. 5B 5B 31, Pl. Fig.

um um

& &

Monniot

. .

sp.

64'E, 24 24 m 64'E,

l l

: :

m m

New Ascidians Ascidians New

, ,

z ,

new species species new

ooid

. .

s; s;

, ,

i o te gut the of ail

. .

; ;

, , 193 193

5 5

cm cm in

5 5

1975 1975

SE SE

Millar, Millar,

Fig. Fig. Pl. 32,

Micronesica Micronesica 29(2), 1996

The The gonads are located inside the gut loop, below the

mm mm in length when partially relaxed. Both siphons are

Eudistoma Eudistoma toealensis

5 5

C). C).

Millar, Millar, 222, Kei Islands, 1975: Toeal.

The pyloric gland consists of few large tubules tubules the stom large facing gland The of few pyloric consists

C). C).

23.13'N; 23.13'N; m;-Palau, 1 124°34.22'E, Peliliu Island, dock area, 6°59.l0'N;

Numerous Numerous lobes in a bunch arise from a common base. They have a cylin

The The secondary curve of the intestine seems indicative of the and species not

The peculiarity of this species is the the abundant is The peculiarity with of mixed orange this species pellets fecal

Description: Description: The are colonies flat cushions, long or circular of about

The The zooids reach

134°13.30'E, intertidal intertidal 134°13.30'E, and Ngercheu, 7°05.39'N; intertidal. 134°16.24'E,

parallel. parallel.

and inside the colony, but superficially harder harder the and colony, but inside on superficially the stalks and common the base.

is covered covered is with epibionts and sand. The test is gelatinous and soft on the heads

nado, nado, l

Material Material Examined: MNHN: 82-Indonesia, Eud A3 79, 72, north Ma Sulawesi, The The zooids are except in the heads irregularly crowded where they are everywhere

about about cm in 1 length and cm in 1 diameter. The basal mass on which they raise

an an orange color to the colonies (Pl. SE). The heads are unequal, and measure

without without encrusting material on the lobes. The orange-red pigmented zooids give

drical drical stalk and an enlarged head containing the thoraces. The test is translucent

Eudistoma Eudistoma toea/ensis

the the result of a contracted state.

rectum rectum (Fig. 31 (species name). name). (species Another peculiarity is the pyloric duct entering the stomach half

inclusions inclusions filling the tunic which give the uncommon rusty color to the colony

located located posteriorly. No other organs could be seen.

the the tail the is same length. There are 3 adhesive papillae in the median and line

stomach. stomach. They consist of a cluster of and spherical testis follicles a central ovary.

ach. ach. The common pyloric duct crosses the gut loop and enters the stomach wall

way way along instead of posteriorly.

a a short ampulla leading to an enlarged mid-intestine curved before reaching the

rectum. rectum. The stomach far is posterior, followed by a narrow cylindrical segment,

the the first row, which is curved dorsally. The abdomen has a long esophagus and

pairs pairs of bifurcated ectodermal ampullae (Fig. 31D). The ocellus and otolith are

194 194

Only Only one larva incubated is at a time in the thoracic The cavity. trunk and long is

at at mid length (Fig. 31

side. side. About stigmata 30 on the right side and 25 on the left side were counted in

forate forate areas but with long stigmata. The rapheal languets are displaced to the left

numerous numerous The fibers. muscle branchial sac long is (Figs. without B), 31A, unper

tubular, tubular, the the cloaca} longest; each has 6 lobes. The body wall is opaque with

in in the center. and pellets The with tough, tunic fecal small orange is filled vesicles.

systems systems of about zooid 8 oral apertures in a ring and grouped apertures cloaca}

There There are also small inorganic particles.

which which turned uniformly rusty throughout in formalin. The smooth surface shows maximum maximum dimension and cm thick. 1 The color in was ochre-yellow life (Pl. 9B)

limb of the intestine makes a short curve at its its curve level. at short limb inside of located intestine a the are makes gonads The

has no special characters (Figs. 32A, B). The stomach is large and the ascending ascending the and large is stomach 32A, B). The (Figs. special characters no has

counted on the right side. They are less numerous on the left side. The abdomen abdomen side. The left the on less are numerous side. They right the on counted

the gut loop. Two to 4 embryos are incubated in the cloaca} cavity at different different cloaca} at the cavity in incubated are embryos 4 to Two loop. gut the

the longitudinal ones. The branchial sac is narrow but up to 20 stigmata were were stigmata 20 to up but narrow is sac branchial The ones. longitudinal the

thoracic muscles make a regular network, the transverse fibers are as strong as as strong as are fibers transverse the network, regular a make muscles thoracic

lobes (Fig. 32A). The lobes are slightly protruding at the colony surface. The The surface. colony the at protruding slightly are lobes The 32A). (Fig. lobes

The zooids are large, up to 1 cm in length. Both siphons are short with 6 6 with short are siphons Both length. in cm 1 to up large, are zooids The

Figure 32. 32. Figure

region; C,D, larvae. larvae. C,D, region;

Eudistoma toealensis toealensis Eudistoma

N N

3 3

Monniot Monniot

& &

Millar, 1975:A, zooid; B, detail of the stomacal stomacal the of detail B, 1975:A, zooid; Millar,

Monniot: New Ascidians Ascidians New Monniot:

C C

0.5 0.5

mm mm 195 195

species. species.

Eudistoma Eudistoma laysani

Eudistoma Eudistoma

to to 3 cm in height. In formalin

deposited deposited in the Copenhagen

1 1

Papua Papua New Guinea, Duke of York

sp. sp.

D). D).

Eudistoma Eudistoma

large ectodermal ectodermal large ampullae, each and deeply irreg

Micronesica Micronesica 29(2), 1996

A3 A3 Eud 109

5 5

Eudistoma Eudistoma toealensis

cm cm in diameter and

or or

5 5

4 4

MNHN: MNHN:

The The gonads are simultaneously present in the gut loop, testis with few

. .

adhesive papillae which are not not are spaced, and papillae regularly which developed, not equally adhesive

Remarks: Remarks: The peculiar color and shape, colony rectum the and long small the

The The zooids have a strong musculature and are deeply retracted inside the

Geographic Geographic Distribution: This species seems limited to very shallow sub

The The colonies are opaque, dark purple with black spots which correspond to

Remarks: Remarks: These differ specimens from the related closely

The The type specimen of

5 5

the the cavity cloacal of some zooids.

Mature Mature tadpoles were not present. An undeveloped embryo being incubated is in

the zooids the number number are of follicles testis not reminiscent of any described

level the the level rectum is bent, this does not seem to be due to the contracted state of

lobes lobes and a central ovary. A protruding egg is often present above the ovary.

anterior anterior to the loop and remains isodiametrical along its length. At the stomach

is is narrow and cylindrical as the mid-intestine, both separated by a constriction.

in the less contracted zooids, it seems to have 2 lateral swellings. The post-stomach The post-stomach lateral swellings. 2 to have contracted zooids, seems less it the in

the the pigment diffuses and the takes fixative a red color. No sand present is in the is long. The The stomach asymmetrical, long. short, is is located at¾ the abdominal of length

equal in size. The number of stigmata in a row could could not counted. number The The abdomen row a be stigmata in of size. in equal

the the zooid apertures. The surface smooth. is The tunic is tough and rubbery. The The The posterior intestine begins without caeca in the descending limb of the gut,

colony. The The colony. body opaque, is wall dark purple. Both siphons have and 6 lobes are

colony colony but only some rare pellets. fecal

Material Material Examined:

colonies colonies are stalked, to 1

Islands, Islands, Mioko Island, 4°13.85'S; 152°28.l6'E, 23 m.

strates strates in Indonesia and Palau.

3 to to 3

influence of of the substrate influence during the growth of the colony. The larvae of the type

for for was specimens the also Sulawesi found in the type.

and and irregular ectodermal ampullae are along each side. The variability observed

vary vary and in size in shape, even in a same lobe of the colony. They are long with

oblique oblique position as described by Millar. This structure may be the result of an

a a thick base containing crowded zooids and fecal pellets, and raised lobes in an

196 196

(Sluiter, 1900) by their larger zooids and a different structure and size of the larvae. larvae. the of and structure their different zooids a larger by and size 1900) (Sluiter,

Zoologisk Zoologisk Museum has been examined. The colony is very similar to ours, with

and and their number are variable (Figs. 32C,

ularly ularly the divided, ventral one extending longitudinally. The shape of the papillae

D), the trunk reaching a length from 1.1 to 1.3 mm. The 4 conical adhesive papillae papillae adhesive conical 4 mm. The to trunk D), the from length reaching a 1.3 1.1

the the posterior part of the abdomen. The mature tadpoles are elongated (Figs. 32C,

are are intercalated between development development stages. Stolonical often vessels, branched, project from the side of

f uln Island Dublon of

Exostoma ianthinum: ianthinum: Exostoma Me)Iln, etre,72.9N 151°49 7°25.29'N; west reef, Island, (Moen)

Polycitor ianthinus ianthinus Polycitor 7°10

tertidal; Mutremdiu, 7°16.Sl'N; 134°31.SS'E, intertidal; Wonder Channel, Channel, Wonder intertidal; 134°31.SS'E, 7°16.Sl'N; Mutremdiu, tertidal;

ai,71.7N 143.7£ 1 ;Rc Ilns 7 Islands, Rock 12 m; 134°30.37'£, 7°18.47'N; Basin, Moresby, Bootless Inlet, south Motupore Island, 9°31.81 9°31.81 Island, 147°17.0S 'S; Motupore south Inlet, Bootless Moresby,

Material Examined: Examined: Material

identify the specimen to the species species level. the to specimen the identify

However, the contraction and the presence of a single colony do not allow to to allow not do colony single a of presence the and contraction the However,

. .

'; 134°22 7'N;

Fig

re 33 re

. .

Exosroma Exosroma

1 'E, 1 m;-Federated States of Micronesia, Chuuk Atoll, Weno Weno Atoll, Chuuk Micronesia, of States 1 1 m;-Federated 'E,

, ,

lie, 1909 Sluiter,

7°22

MNHN

ot 1990a Kott,

Exostoma ianthinum ianthinum Exostoma

5N 115.7£ 4 m; 45 151°54.77'£, 95'N;

ia11

Monniot Monniot

hi11

: :

A3 Exo 1 to 7-Palau, Koror State, Turtle Island Island Turtle State, Koror 7-Palau, 1 to Exo A3

m m

20 20

Fig. 33, Pl. SF SF 33, Pl. Fig.

& &

: :

234 and synonymy synonymy 234 and

Monniot

uite

, ,

1909): 1909): A,

: :

New New Ascidians

(Sluiter, 1909) 1909) (Sluiter,

02'E, 4 m; Halimeda sand ridge east east ridge sand Halimeda m; 4 02'E,

, zooids; zooids; ,

Papua New Guinea, Port Port Guinea, New Papua

18.48'N; 134°28.12'E, in 134°28.12'E, 18.48'N;

B B

C, go C,

. .

, ,

3 m. m. 3 197 197

(Sluiter (Sluiter 1895)

Pl. Pl. 5H

Micronesica Micronesica 29(2), 1996

A3 A3 Hyp Bl and 2-Papua New Guinea, Madang,

Kott, Kott, and 106 1990a: synonymy.

Hypodistoma Hypodistoma deerratum

MNHN: MNHN:

Sluiter, Sluiter, 167. 1895:

Both Both siphons are long and tubular. They have 6 short lobes. counted We 14

The The stomach has a smooth wall. The digestive loop contains the gonads: a

The The colonies are massive lobes, to 15 20 cm in extent, with a large volcano

Geographic Geographic Distribution: The was species already known from the tropical

Remarks: Remarks: The characterized is species by its wide and cav cloaca] extensive

The The tunic is tough, opaque and dark brown in formalin. The zooids are

The The colonies are large irregular masses up to cm 25 across, and cm 5 thick.

thorax. thorax. When mature, this larva has a developed well thorax with differentiated

144°57.?'E, 18 m. m. 144°57.?'E, 18

to 20 stigmata to stigmata 20 in the first of the of rows 3 the branchial A sac. unperforated large appendage. appendage. One larva incubated is in a brood pouch appended to the base of the

short, short, displaced to the left side.

area area lies between the tentacles and the branchial sac. The rapheal languets are

central central ovary and The small few follicles. testis abdomen ends in a long vascular

tunic. tunic. Large cloaca] channels extend between these two parts. The zooids are

restricted restricted to the upper layer of the colony, the center consists of a firm core of

translucent translucent smooth undulations hollow encircling rounded parts. The zooids are

distributed distributed in groups.

Material Material Examined:

like like aperture 2 cm in diameter (Pl. 5H). The surface has grey, tan, or brown

Hypodistoma Hypodistoma deerrata:

Distoma Distoma deerrata

Tab Tab m; 'S; Boisa Island, 12 Island 4°00.0?'S; 145°50.37'E, on barrier reef, 5°10.21

stomach, stomach, inside the gut loop. No larvae were found in these colonies.

the the stomach. behind the Male gonads a developed in cluster follicles of testis were

western Pacific (Palau, (Palau, western Pacific Philippines, Indonesia and New Guinea).

33C). The The 33C). pyloric gland consists of tubules around the part of the rectum facing

ities, ities, and the zooid structure.

tudinal tudinal located crest is on of the each typhlosole. side The narrow mid-intestine is

before before entering the posterior intestine that much is wider but without caeca (Fig.

long. long. The stomach is followed by a posterior stomach. In some zooids a longi

rated rated area also extends at the base of the branchial sac. The esophagus is very

an an unperforated area. There are about 20 stigmata in the first row. An unperfo

the thorax thorax The the It branchial elongated lobes. B). has below (Figs. 6 also 33A, is sac

198 198

irregularly irregularly spaced, oriented directions. The all in oral siphon has rounded 6 lobes.

The The siphon cloaca) long is and cylindrical, issued from the postero-dorsal part of

is is wrinkled, rubbery. Sometimes thin epibionts are attached in patches to the

few very large openings of the cloaca} cavities (Pl. SF). The surf ace ace of large very the openings colony (Pl. cavities few SF). of The surf the cloaca}

swollen and and swollen the oral apertures of the zooids are irregularly distributed. There are

colony colony surface. The The color varies from grey to brown, mottled. The colony surface is irregularly

h soah clrd noag i ie soiesae ntemdl fte ab the of middle the in olive-shaped is life, in orange in colored stomach, The

men is elongated, linked to the thorax by a narrow and long esophago-rectal waist. waist. long esophago-rectal and by a narrow men is thorax linked the to elongated,

domen. The post-stomach is dilated in a ring before the constriction leading to to leading constriction the before ring a in dilated is post-stomach The domen.

weak, mostly longitudinal. There are 2 unperforated areas anterior and posterior posterior and anterior areas 2 unperforated are There longitudinal. mostly weak,

due to the zooids being enclosed in capsules of spicules. spicules. of capsules in enclosed being zooids the to due to the branchial sac. The stigmata are elongated and very numerous. The abdo The numerous. very and elongated are stigmata The sac. branchial the to

State, Ushi Island passage, 7°40.40'N; 151°49.50'£, 30 30 m. 151°49.50'£, 7°40.40'N; passage, Island Ushi State,

Reef, 7°16.51 'N; 134°26.71 'E, 20 m;-Federated States of Micronesia, Chuuk Chuuk Micronesia, of States 134°26.71 m;-Federated 7°16.51 20 Reef, 'N; 'E,

Material Examined: Examined: Material

despite Kott's (1990a) estimate that there is only one species species of one is only there that (1990a) estimate Kott's despite

theless the spicule shape is very characteristic. characteristic. is very shape spicule the theless

than the present records suggests. suggests. records believe We is probable, present the its in presence Australia than

ant edtrie, n te pcfcietfcto rmis neti. Never uncertain. remains specific identification the and determined, be cannot

recorded in New Caledonia (F. Monniot, 1988). It very likely has a wider range range wider a likely has very 1988). It Monniot, (F. Caledonia New in recorded

the colony and the zooids. As the larvae are missing, the place of incubation incubation of place the missing, are larvae the As zooids. the and colony the Cystodytes punctatus punctatus Cystodytes

in New Caledonia for the color, the spicule shape and arrangement, the shape of of shape the arrangement, and shape spicule the color, the for Caledonia New in

spicules with indentated margins. Both siphons have 6 lobes. The 6 testis follicles follicles testis 6 The lobes. 6 have siphons Both margins. indentated spicules with

ace ecig4 mars.Tezodaetrs r ragd ncrua sys circular in arranged are apertures zooid The across. cm 40 reaching patches

are in a circle. Larvae are not present. present. not are circle. Larvae a in are

tems. The dark pigment cells are more abundant in the deepest part of the tunic tunic the of part deepest the in abundant more cells are pigment dark The tems.

than in the upper layer, which contains some patches of poorly formed spicules. spicules. formed poorly of patches some contains which layer, upper the in than

Atoll, oceanside reef channel west of Pis Island, 7°38.97'N; 151 151 7°38.97'N; m. 53 Island, °41.22'E, Pis of west channel reef oceanside Atoll,

Material Examined: MNHN: A3 Cys 19-Federated States of Micronesia, Chuuk Chuuk States of Micronesia, 19-Federated Cys A3 MNHN: Examined: Material

Fig. 8, pl.I B-C, New Caledonia. Caledonia. New B-C, pl.I 8, Fig.

filiform papillae arise from this part. part. this from arise papillae filiform

lands and Torres Strait. Strait. Torres and lands

ot Asrla o uesad ad o sas nw fo tePiipns Is Philippines the from known also is now and Queensland, to Australia north

vesicles are large, very flat, and attached to an abdominal extension. Very thin thin Very extension. abdominal an to attached and flat, very large, vesicles are

siphons, a large unperforated area in front of 3 rows of stigmata. The adhesive adhesive The stigmata. of rows 3 of front in area unperforated large a siphons, Cystodytes aucklandicus aucklandicus Cystodytes

The colonies are of firm consistency, lobated and vitreous with white spots spots white with vitreous and lobated consistency, firm of are colonies The

The zooids have short siphons with 6 lobes. The thoracic musculature is is musculature thoracic The lobes. 6 with siphons short have zooids The

Geographic Distribution: The species described from New Zealand has been been has Zealand New species from The described Distribution: Geographic

The zooids have the abdomen enclosed in a capsule made of disk-shaped disk-shaped of made capsule a in enclosed abdomen the have zooids The

This specimen corresponds well to the samples of of samples the to well corresponds specimen This

The colonies (Pl. 5G) are encrusting, brown, rubbery, about 5 mm thick, in in thick, mm 5 about rubbery, brown, encrusting, are 5G) (Pl. colonies The

Geographic Distribution: This species was previously known from western western from known previously species was This Distribution: Geographic

Cystodytes punctatus? punctatus? Cystodytes

MNHN: MNHN:

F. Monniot, 1988: 1988: 11, 222, fig. pl.2A Monniot, F.

Cystodytes aucklandicus? aucklandicus? Cystodytes

Nott, 1892: 323, New Zealand; F. Monniot, 1988: 1988: 215 Monniot, F. Zealand; 1892: New 323, Nott,

Monniot Monniot

A3 Cys 80, and 81-Palau, Mutremdiu, Barrier Barrier Mutremdiu, 81-Palau, and 80, Cys A3

& &

Monniot: New Ascidians Ascidians New Monniot:

Pl. 5G 5G Pl.

F. Monniot, 1988 1988 Monniot, F.

Nott, 1892 1892 Nott,

C. C.

aucklandicus aucklandicus

B. B.

New Caledonia. Caledonia. New

Cystodytes Cystodytes

collected collected

there. there. 199 199

by by the larval structure. However,

F. F. Monniot, 1988

Cystodytes Cystodytes

Micronesica Micronesica 29(2), 1996

A3 Cys 82, and and A3 Cys 82, 83-Palau, Rock Islands, Ngerkuul

All zooid zooid characters All but correspond species this closely,

Cystodytes Cystodytes solitus

MNHN: MNHN:

F. F. Monniot, fig. 223 12A-C. 1988:

punctatus. punctatus.

C. C.

Geographic Geographic Distribution: The described species from New Caledonia, and

The zooids appear very differently according according to appear differently their relaxation. state zooids very The of Both

The The form colonies encrusting cm across sheets several and 1 cm thick, lav

The The disk-shaped have spicules a thin rim.

This This sample looks exactly like the colonies collected in New Caledonia and

Pacific Ocean. Ocean. Pacific

now now known from Palau, certainly has a larger distribution the in tropical western

numerous numerous dark lacking pigment in the zooids. cells, There papillae are adhesive 3

measures 1.2 mm. mm. measures 1.2

bundle extends along the abdomen. Only one large larva is incubated in the cloaca} cloaca} the incubated in larva abdomen. is along the large one extends Only bundle

encircled by encircled a rod made from 4 pairs of ectodermal ampullae. The larval trunk

the the center of the rosette straight. is The ovary anterior is to A the muscular testis.

arranged arranged in a at circle the bottom of the gut loop. The sperm duct starting from

cavity, but there is no no brood but is cavity, there pouch. contains The the larvae epithelium of external

relaxed relaxed animals. The esophagus narrow, is the stomach is just located below the

enlarged enlarged part at the end of the descending limb of the gut. are The follicles testis

middle middle of the abdomen length. The gut loop is wide. The rectum with begins an

the the thorax and the abdomen. On the the left above body side, depressed deeply is

part part of the esophagus, on the right side of the body. The abdomen is long in

the the esophagus aperture. The body wall forms a horizontal fold above the upper

racic musculature racic musculature weak with is thin There a fibers. is strong constriction between

rapheal rapheal languets are rather short and displaced slightly to the left The side. tho

large unperforated large unperforated area in the upper lies part of the branchial The of first the sac.

than than the oral one. There are about 20 to 24 oral tentacles in 3 orders of A size.

4 4 rows of stigmata contain 22 to 28 stigmata each and are bent dorsally. The

individual individual cloacal openings. Dark pigment are cells scattered in the entire thick

siphons siphons are tubular, with 6 round lobes, but the siphon cloaca} is much longer

smooth. smooth. The oral siphons are arranged in circles whose centers have groups of Cystodytes Cystodytes solitus

spicules spicules arranged around the abdomens of the zooids. The colony surface is

ness ness of the tunic.

ender ender or purple color. in The tough tunic is and translucent calcareous with white

Material Material examined:

m. m.

Pass, Pass, m 7 7°19.22'N; and 134°29.74'E, Kaibaku Cave, 7°19.47'N; 9 134°29.42'E,

is mainly is mainly distinguished from other

identification. identification.

the absence of larvae larvae not does of here the absence us to absolutely certain be allow the specific of

described as described

gut gut loop. The ovary is not developed and no larvae were found.

the the intestine. The testis is made of long arranged follicles in a rosette inside the 200 200

is bright blue in life in blue bright is

is 8.5 cm in length and 2.5 cm in diameter, tapering at its free extremity. The color color free its The extremity. at tapering 2.5 is cm 8.5 diameter, in and cm length in

6A). The largest has a stalk 7 cm long and the head containing numerous zooids zooids numerous containing head the and long 7 cm stalk a has largest The 6A).

Nephtheis ma/ayensis ma/ayensis Nephtheis

Nephtheisfaciformis Nephtheisfaciformis

nesia. nesia.

region

Nephtheis centripetens centripetens Nephtheis

Material Examined: Examined: Material

Nephtheis Nephtheis

145 Colella thomsoni thomsoni Colella

Oxycorynia fascicularis fascicularis Oxycorynia

50.37'£, 10 10 m. 50.37'£,

The colonies are groups of stalked lobes arising from a common base (Pl. (Pl. base common a from arising lobes stalked of groups are colonies The

Fig

Papua New Guinea, Madang, Tab Island barrier reef, 5°10.21 5°10.21 'S; reef, barrier Island Tab Madang, Guinea, New Papua

(?) (?)

e 34 e

gon

Gould, 1852: 1852: Gould, 52, fig. Philippines. 621, pl.

. .

s; s;

Nephtheisfascic

Herdman, 1886: 94, pl.10-13. Philippines. Philippines. 1886: 94, pl.10-13. Herdman,

C, C,

Nephtheisfascicularis? Nephtheisfascicularis?

, ,

MNHN: MNHN:

Sluiter, 1909 Sluiter,

dark blue or dark green in alcohol. The tunic is soft, trans- soft, is tunic The alcohol. in green dark or blue dark

Sluiter, 1909 Sluiter,

rva rva

Sluiter, 1909 Sluiter,

von Drasche, 1882: 177, Caroline Islands. Islands. 1882: 177, Caroline Drasche, von

Monniot Monniot

!I !I

;a ;a

N N

A3 Nep 1 and 2-Indonesia, Banta Island, Komada Komada Island, Banta 1 A3 2-Indonesia, Nep and

laris? laris?

Fig. 34, PI. 6A 6A PI. 34, Fig.

: : & &

: :

(von (von

39, pl.3, fig 39, pl.3,

: :

Monniot: New Ascidians New Ascidians Monniot:

36, pl.3, fig 36, pl.3,

38, pl.2, fig.17, pl.3, fig.12, pl.8, fig.4, Indo fig.4, fig.12, pl.8, fig.17, pl.3, pl.2, 38,

rasc

(von Drasche, 1882) 1882) Drasche, (von

e 1882) e

3-4, pl.7, fig.2, fig.2, Indonesia. pl.7, 3-4,

8-10, Indonesia. Indonesia. 8-10,

: :

A, A,

zooi

; ;

boe wi abdomen ,

1 1

mm mm

th th 201 201

fas

N. N.

fascicu/aris, fascicu/aris,

as as described

the the sample la

N. N.

In In

The The thoracic mus

fascicularis. fascicularis.

is is long, totally obscured by

N. N.

It It

fascicularis. fascicularis.

N. N.

Nephtheis fascicu/aris fascicu/aris Nephtheis

Sluiter, Sluiter, 1909 for all characters: mus

Sluiter, Sluiter, a 1909 problem arises.

Micronesica Micronesica 29(2), 1996

centripetens centripetens

Sluiter, Sluiter, 1909 the colonies are branched, with well

N. N.

Sluiter, 1909 is definitely a a synonym Sluiter, definitely is of 1909

(Herdman, (Herdman, has 1886) zooids similar to those of

Nephtheis Nephtheis faciformis

Sluiter, Sluiter, 1909 and

Nephtheis Nephtheis malayensis

With With

Nephtheis Nephtheis centripetens

In In

Nephtheis Nephtheis thomsoni

Remarks: Remarks: These specimens correspond to

The The zooids are deeply pigmented by dark cells in a dense network of blood

short, short, curved, isodiametric, with a smooth wall. The stomach is relatively short,

single lobe, lobe, single preserved well , has no constriction between the head and the stalk.

belled belled type, or lectotype from the Siboga station 310, the largest colony with a

The The zooids are relatively small but have 20 rows of stigmata. The esophagus is

with with a square enlargment in the middle as in

comes comes from a station different from that of the type).

stigmata stigmata varies from to 13 in both 15 cases, and the esophagus has a smooth wall

clearly clearly separated from the stalk by a constriction (the sample labelled paratype

culature culature is the same, as are the gonads and the larvae. The head of the colony is

identified, identified, with insufficient characters, but may be

with with a discolored type and deeply pigmented paratype. The number of rows of

cicularis cicularis developed developed stalks but very and small young cannot The zooids. species precisely be

culature, culature, branchial sac, digestive tract and gonads.

For For the other synonymous species, the type and paratype specimens have been

studied: studied:

by by Kott The (1990a). type specimen of von Drasche has (1882) not been located.

from from New Caledonia by F. Monniot or (1988) from Australia and the Phillipines

a a triangle and a branchial sac with 6 to 8 rows of stigmata (Fig. 34C).

the the rectum with the oviduct. The latter crosses the esophagus entrance and opens

The The embryos their begin development the cloacal the in oviduct in and free become

at at the bottom of the thorax. The heart vertical in is the gut loop and voluminous.

cavity cavity when the tail is developed. In aged larvae, there are 3 adhesive papillae in

are distributed distributed are on gut the whole loop. central. The ovary is The sperm duct follows

is is not divided in regions and the anus has a thin smooth rim. The testis follicles

pigment pigment cells. The gut loop is narrow, even at its posterior end. The esophagus

middle middle (Fig. The 34B). stomach is olive shaped and smooth-walled. The intestine

above above the esophageal opening. The cloacal cavity is large and contains tadpoles.

has has longitudinal irregular plications, with an enlarged quadrate segment it its

with with to 14 25 rows of numerous long stigmata. It forms a postero-ventral pouch

of of them anastomosed dorsally and ventrally (Fig. The 34A). branchial wide sac is

The The abdomen is much smaller than the thorax.

34A). The The 34A). oral siphon has a short velum anterior to the 8 tentacles. The thoracic

sinuses sinuses in the thoracic body wall, and by blue and dark opaque cells in all ab

dominal dominal organs. Both siphons have their margin undulated by contraction (Fig.

musculature musculature mostly is transverse with numerous bundles, regularly spaced, some

side side external, is the abdomen internal.

202 202

colony colony are about the same size and the same development stage. Their ventral parent parent on the head, a little harder and opaque on the of ids the same stalk. zoo All

tip of the stolons, as in in as stolons, the of tip

Trason (1963) showed that the budding does not originate from chambers at the the at chambers from originate not does budding the that showed (1963) Trason

u rte b srblzto o h zod n rgnrto o n o svrl stro- several or one of regeneration and zooid the of strobilization by rather but

budding and the larva of the Californian species species Californian the of larva the and budding

has been largely modified by Trason (1963) who studied in detail the mode of of mode the detail in studied who (1963) Trason by modified largely been has

xeso o h bsl omn ui. oee, hs to eea ae en syn been have genera two these However, tunic. common basal the of extension

onymized (Brunetti 1987, Kott 1990a). The definition of the genus genus the of definition The 1990a). Kott 1987, (Brunetti onymized

considered this genus to be intermediate between between intermediate be to genus this considered

where they remain sterile or, more rarely, give rise to new buds" (p. 53). Garstang Garstang 53). (p. buds" new to rise give rarely, more or, sterile remain they where

Stereoclavella Stereoclavella

some of which even extend and branch in the esophageal region of the zooids, zooids, the of region esophageal the in branch and extend even which of some

presence of a basal mass of tunic "traversed in all directions by stolonial tubes, tubes, stolonial by directions all in "traversed tunic of mass basal a of presence

1949. 1949.

which we disagree. The genus genus The disagree. we which

P. aurilucens aurilucens P.

the presence of numerous larvae incubated inside the oviduct. oviduct. the inside incubated larvae numerous of presence the

characters of her definition of of definition her of characters

unknown

oas oeig h hl gt op bhn te stomach the behind loop, gut whole the covering gonads

fig. 13, 14, S Australia; Kott, 1990a: 73, fig. 24, pl. 6a-f, Australia and Philippines. Philippines. and Australia 6a-f, pl. 24, fig. 73, 1990a: Kott, 14, S 13, Australia; fig.

Material Examined: Examined: Material

Chuuk Atoll. Atoll. Chuuk

Pycnoclavelladiminuta: Pycnoclavelladiminuta:

1988 is a distinct species characterized by its smaller size, a less wide thorax, and and thorax, wide less a size, smaller its by characterized species distinct a 1988 is

collection does not extend the known range of this species. species. this of range known the extend not does collection

? Archidistoma rubripunctum rubripunctum ? Archidistoma

well as western, northern and eastern Australia and New Caledonia. The present present The Caledonia. New and Australia eastern and northern western, as well

morphology of the zooid may be different with different sizes of the colonies. The The colonies. the of sizes different with different be may zooid the of morphology

study of large populations would be necessary to establish if there is only one one only is there if establish to necessary be would populations large of study

species or more or species

long time in alcohol or formalin and collected in different parts of the world. The The world. the of parts different in collected and formalin or alcohol in time long Clave/ina diminuta diminuta Clave/ina

with a smooth wall and an enlarged pouch, in its middle. middle. its in pouch, enlarged an and wall smooth a with

and general pigmentation is difficult to establish with specimens preserved for a a for preserved specimens with establish to difficult is pigmentation general and

station 49A has zooids with only 13 to 15 rows of stigmata, a long esophagus esophagus long a stigmata, of rows 15 to 13 only with zooids has 49A station

well separated from the intestine. The sample labelled paratype from the Siboga Siboga the from paratype labelled sample The intestine. the from separated well

. .

aurilucens aurilucens

Monniot, 1988 may be a synonym. On the contrary, contrary, the On synonym. a be may 1988 Monniot,

Kott (1990a) placed this species in the genus genus the in species this placed (1990a) Kott

The colony The

Geographic Distribution: Indonesia, the Philippines and Caroline Islands, as as Islands, Caroline and Philippines the Indonesia, Distribution: Geographic

The variability in the number of rows of stigmata, length of the abdomen abdomen the of length stigmata, of rows of number the in variability The

P. auri/ucens auri/ucens P.

. .

This colony corresponds to the species defined by Kott (1990a). Some Some (1990a). Kott by defined species the to corresponds colony This

lacking epidermal papillae and and papillae epidermal lacking

rm h Biih hne, a atr him after has Channel, British the from

Herdman, 1891 b, a distinction corresponding to a more or less less large or a more to corresponding 1891 distinction a b, Herdman,

. .

, ,

in bad condition does not contain larvae, but has well well developed has but larvae, contain not does condition bad in

and and

Kott, 1957: 89, W Australia. Australia. W 89, 1957: Kott,

MNHN: MNHN:

Archidistoma diminuta diminuta Archidistoma

Clave/ina /epadiformis /epadiformis Clave/ina

ilr 1963 Millar,

. .

stanleyi stanleyi

Monniot Monniot

F. Monniot, 1988: 200, fig. 2 C,D, New Caledonia. Caledonia. New C,D, 2 fig. 200, 1988: Monniot, F.

A. diminuta diminuta A.

Pycnoc/ave/la Pycnoc/ave/la

A3 Arc A9-Federated States of Micronesia, Micronesia, of States A9-Federated Arc A3

& &

differ morphologically in the larval structure, structure, larval the in morphologically differ

Monniot: New Ascidians Ascidians New Monniot:

: :

715 fig. 21, NW Australia; Kott, 1972: 170, 170, 1972: Kott, Australia; NW 21, fig. 715

indicate that that indicate

P. stanleyi stanleyi P.

established by Garstang (1891) for for (1891) Garstang by established

after Brien Brien after

(Kott, 1957) 1957) (Kott,

Pycnoc/avella, Pycnoc/avella,

P. stanleyi stanleyi P.

Clavelina Clavelina

, ,

as major characteristic, the the characteristic, major as

having a crown of of papillae. crown a having

Archidistoma rubripunctum rubripunctum Archidistoma

& &

. .

A. richeri richeri A.

Its appearance in life is is life in appearance Its

Brien-Gavage (1927), (1927), Brien-Gavage

Savigny, 1816 and and 1816 Savigny,

Berrill & Abbott, Abbott, & Berrill

a decision with with decision a

Pycnoc/avella Pycnoc/avella

. .

Monniot, Monniot, 203 203

E. E.

Euclav

is is common

and and more is

It It

Pycnoclavella Pycnoclavella

and and for a species:

Clave/ina Clave/ina

which which have lost their

despite despite having described it

which correspond correspond division to the which

C/ave/ina C/ave/ina

which correspond correspond which to the larval 2

Pycnoclavella Pycnoclavella

without without mentioning his reasons for

from from sectioned abdomens.

Pycnoclavel/a, Pycnoclavel/a,

Archiascidia Archiascidia

Archidistoma Archidistoma

redescribing the redescribing Mediterranean con specimens

Micronesica Micronesica 29(2), 1996

(Polycitoridae) (Polycitoridae) have 3 kinds of papillae; the genus

type type of the genus, described is as producing at the

Pycnoclave/la Pycnoclave/la

the the Phlebobranchia all have the same papillae except

(1953) (1953)

In In

Peres Peres

Eudistoma Eudistoma

C/ave/ina C/ave/ina /epadiformis

P. P. auri/ucens,

has has papillae similar to those of

(Garstang (Garstang Garstang in 1895). work his (1895) describing budding in

F. F. Monniot, an other 1987, kind of papillae.

Euherdmania Euherdmania claviformis.

The The structure species studied 28 in histologically was papillae the of adhesive

The The structure stud was the larval of papillae in families all ascidian adhesive

Is Is there any to real in significance differences the budding and in the larva?

Kott (1990a) described a new family, the Pycnoclavellidae, with with Pycnoclavellidae, the family, new a described Kott (1990a)

Millar Millar used (1953, 1963)

Trason Trason (1963) also described the structure of the larva which posseses ad

Kott, Kott, The principal 1990a. characters of the family concern the structure of

C/ave/ina C/ave/ina

found found in another family. To separate the genera, we use instead the branchial primary rows; 3 rows of stigmata stigmata in of rows 3 primary rows;

development: of development: rows 2 stigmata in

single kind kind single of papilla as a character justifying a generic division and to further

base base the creation of a new family, particularly when this type of papilla is also

and and this represents one per thousand of the described ascidian Ten species. dif

ferent ferent structures were defined. Consequently it seems questionable to consider a

dumosa dumosa

Euherdmania Euherdmania

ferent ferent genera. The

the the Perophoridae. Among the Aplousobranchia, only the Didemnidae ho seem

mogenous. mogenous. The Polycitoridae and possess 4 Polyclinidae kinds of papillae in dif the the two Styelidae kinds.

scribed. scribed. A tenth condition, the absence of papillae, can be added. In the Stoli

ied ied in detail by Cloney (1977, and 1978, 1990) 9 different kinds of papillae de dobranchia, dobranchia, while papillae the Molgulidae kind and of Pyuridae one the only have

thorax, thorax, or from zooids regressed which reorganise after unfavorable conditions. or less expressed according according expressed or to or less the the species conditions of the of colonies. life

to to observe regenerations of damaged zooids in The The potential for both budding or regeneration coexists in

several years earlier. earlier. years several of of the thorax of

firmed the firmed duality of the mode of budding. a described regeneration Brien (1970)

same same time stolons and buds. It does not appear very different from the budding Tunicata Tunicata did not mention the genus

of of

the the family. The The budding of

of of Trason The (1963). other defining characters concern the proportions of the

the the larval adhesive papillae, and the mode of budding, in to reference the work different different parts of the body, or characters all of which representing exceptions in

as as the type of genus also the which included family, the monotypic genus

ella ella

using using name. this generic

terminology terminology and (1977) similar to those described by Trason in the (1957) Poly

hesive papillae papillae hesive of the type: "tubular invaginated papillae" according to Cloney's clinidae clinidae

sidered sidered to be intermediate between the Polyclinidae and the Polycitoridae.

biles. For For biles. Trason this was the (1963) main character of the con she which genus 204 204

iable according to the specimens. The tunic is covered with epibionts, including including epibionts, with covered is tunic The specimens. the to according iable

patches around the oral siphon and between the siphons; their disposition is var is disposition their siphons; the between and siphon oral the around patches

ml age ddmi clne ad yrzas Atr iain al oos disap- colors all fixation, After hydrozoans. and colonies didemnid algae, small

upwards. Both siphons are short without lobes, with a denticulated rim. The tunic tunic The rim. denticulated a with lobes, without short are siphons Both upwards.

is transparent and the body wall opaque brown-purple. There are gold yellow yellow gold are There brown-purple. opaque wall body the and transparent is

usrtm yvr sot n sot hzis Te oy shr-ie wtot sep without horn-like, is body The rhizoids. stout and short very by substratum

siphon opens perpendicularly to the body axis and the cloaca! aperture is is directed aperture cloaca! the and axis body the to perpendicularly opens siphon

aration in thorax and abdomen. The siphons are apical, widely open. The oral oral The open. widely apical, are siphons The abdomen. and thorax in aration

Material Examined: Examined: Material

Podoc/avel/a meridionalis meridionalis Podoc/avel/a

tralia. tralia.

ern Pacific and western Australia. Australia. western and Pacific ern

Clavelina meridiona/is: meridiona/is: Clavelina

and, as as and,

elongated stomach with 10 to 12 plications. Kott (1990a) describes a "large "large a describes (1990a) Kott plications. 12 to 10 with stomach elongated

artifacts"

roomy" stomach with "4 or more sometimes irregular rounded folds apparently apparently folds rounded irregular sometimes more or "4 with stomach roomy"

color disappears in fixatives. fixatives. in disappears color

by Kott (1990a), 16 by Tokioka (1952). Tokioka described and figured a rather rather a figured and described Tokioka (1952). Tokioka by 16 (1990a), Kott by

color seem characteristic of the species, as figured by Kott ( 1990a, pl. I, e, f). The The f). e, I, ( pl. 1990a, Kott by figured as species, the of characteristic seem color

pqe elwgen ace o bt sds f h oa siphon oral the of sides both on patches yellow-green opaque

tunic is transparent and the thoraces are translucent, black or purple, with two two with purple, or black translucent, are thoraces the and transparent is tunic

Ngerkuul Pass, 7°19.B'N; 134°29.87'E, 10 10 m. 134°29.87'E, 7°19.B'N; Pass, Ngerkuul

Pearl Oyster Bed, Melville Bay; Nishikawa, I 984: 116, Truk (Chuuk) Island; Kott, Kott, Island; (Chuuk) I 116, Truk 984: Nishikawa, Bay; Melville Bed, Oyster Pearl

(Pl. 6B). The groups are united at their base by a network of stolons. In life, the the life, In stolons. of network a by base their at united are groups The 6B). (Pl.

Material Examined: Examined: Material

r, n nrhr Asrla Nw aeoi, Philippines Caledonia, New Australia, northern and ern,

sac. sac.

Clave/ina ( Synclavella) arafurensis arafurensis ( Synclavella) Clave/ina

stigmata and multiplication by the anterior and posterior sides of the branchial branchial the of sides posterior and anterior the by multiplication and stigmata

1990a: 38, fig. 7, pl. 1,e-f, W. Australia, Philippines. Philippines. Australia, W. 1,e-f, pl. 7, fig. 38, 1990a:

of the first primary row only, and in in and only, row primary first the of

This species has large isolated zooids (up to 8 cm) (Pl. 6C) , attached to the the to attached , 6C) (Pl. cm) 8 to (up zooids isolated large has species This

Geographic Distribution: The species was previously recorded in all the west the all in recorded previously was species The Distribution: Geographic

In the Palau specimens we counted 14 rows of stigmata-12-16 indicated indicated stigmata-12-16 of rows 14 counted we specimens Palau the In

The colony consists of 1 to 7 zooids grouped in a cylinder of common test test common of cylinder a in grouped zooids 7 to 1 of consists colony The

egahc itiuin Peiu rcrs nld wsen suhr, east southern, western, include records Previous Distribution: Geographic

not~ not~

. .

Our specimens show a pyriform stomach with thick swellings (Fig. 35E) 35E) (Fig. swellings thick with stomach pyriform a show specimens Our

by by

Kott (1990a), a neat post-stomach. post-stomach. neat a (1990a), Kott

MNHN

Clave/ina meridiona/is meridiona/is Clave/ina

MNHN: MNHN:

ot 19aad yoyy 4,wsen n esen Aus eastern and western 48, synonymy: and 1990a Kott,

Clavelina arafurensis arafurensis Clavelina

Herdman, 1891a: 603, Australia, Port Jackson. Jackson. Port Australia, 603, 1891a: Herdman,

Monniot Monniot

: :

3Ca 67 Cla A3

Figs. 35A-D, PI. 6C 6C PI. 35A-D, Figs.

A3 Cla 68-Palau, Koror State, Rock Islands, Islands, Rock State, Koror 68-Palau, Cla A3

Fig. 35E, PI. 6B 6B PI. 35E, Fig.

& &

Tokioka, 1952: 97, fig. 5, Papua New Guinea, Guinea, New Papua 5, fig. 1952: 97, Tokioka,

Monniot: New Ascidians Ascidians New Monniot:

Clavelina Clavelina

Indonesia, north Sulawesi, Manado. Manado. Sulawesi, north Indonesia,

(Herdman, 1891) 1891) (Herdman,

Tokioka, 1952 1952 Tokioka,

division of the two larval rows of of rows larval two the of division

. .

This design and and design This 205 205

hard hard

, ,

Tokioka

cylinders, cylinders,

measured measured when

be be

B, B, posterior part of

3 3

"' "'

zooid; zooid;

, ,

C/ave/ina arafurensis arafurensis C/ave/ina

. .

larva

. .

, ,

to depend on the number of successive zooids. zooids. depend to number the on successive of

(Herdman, (Herdman, 1881):

muscles; muscles; D

Micronesica 29(2), 1996 1996 29(2), Micronesica

seems seems

meridionalis meridionalis

C, C, thoracic

51 51

3 3

V, V,

; ;

posterior posterior part of the abdomen

Clave/ina Clave/ina

E, E,

35. 35.

1952: 1952:

the the abdomen

Figure Figure

The The zooid in divided is parts 3 proportions which cannot

the thorax thorax the and abdomen, and

to to the base of the tunic. The variable length of this process may exceed that of

contracted: contracted: the thorax, the abdomen, and thick a opaque long down going process

and and opaque, which represent the tunics of eldest zooids.

pear. pear. In many specimens the basal tunic has one or several test 206 206

area ( ( area

cies. cies.

to the base of the tunic. The anus of of anus The tunic. the of base the to

maining in fixatives. These two species have also the abdomen prolongated down down prolongated abdomen the also species have two These fixatives. in maining

cluster of zooids results. In In results. zooids of cluster

part very early from the mother zooid, even before the siphons are open, and a a and open, are siphons the before even zooid, mother the from early very part

miniata miniata solitary individuals. individuals. solitary C. C.

idiona/is idiona/is

viola viola the zooids. zooids. the

are are

ave 1 larvae

and occupies the whole right side of the abdomen up to the stomach level. The The level. stomach the to up abdomen the of side right whole the occupies and

The sperm duct arises from the bottom of the gut loop, crosses the ovary seeming seeming ovary the crosses loop, gut the of bottom the from arises duct sperm The

right cloaca! cavity where numerous larvae are incubated. The heart is very large large very is heart The incubated. are larvae numerous where cavity cloaca! right

to divide it into two parts, and ends near the anus. The oviduct is dilated in the the in dilated is oviduct The anus. the near ends and parts, two into it divide to

is located on the left side of the body (Fig. 35B). The testis forms small follicles follicles small forms testis The 35B). (Fig. body the of side left the on located is

the curve of the gut loop. The anus possesses numerous round lobes. The ovary ovary The lobes. round numerous possesses anus The loop. gut the of curve the

scattered around the intestine which even overly the stomach on the right side. side. right the on stomach the overly even which intestine the around scattered

or less plicated.The post-stomach is well differentiated. The intestine begins before before begins intestine The is well differentiated. post-stomach less plicated.The or

have differentiations. differentiations. have

is separated from the thorax by a definite constriction. The esophagus does not not does esophagus The constriction. definite a by thorax the from separated is

multiply the number of rows of stigmata. The transverse vessels have internal high high internal vessels have transverse The stigmata. of of rows number the multiply

contracted parts of the branchial tissue, the stigmata measure 0.30 to 0.45 mm. mm. 0.45 to 0.30 measure stigmata the tissue, branchial the of parts contracted

part of the branchial sac, the transverse vessels appear from the endostyle to to endostyle the from appear vessels transverse the sac, branchial the of part

blades; 0 blades;

development of transverse vessels starting from the dorsal lamina. In the middle middle the In lamina. dorsal the from starting vessels transverse of development

formed from the dorsal lamina with perforation of round stigmata before the the before stigmata round of perforation with lamina dorsal the from formed

and posterior parts of the branchial sac we observed incomplete rows of stigmata stigmata of rows incomplete observed we sac branchial the of parts posterior and

straight aperture. The dorsal lamina consists of at least 27 languets. In the anterior anterior the In 27 languets. least of at consists lamina dorsal The aperture. straight

where the larvae are incubated. The muscles extend along the abdomen. A dozen dozen A abdomen. the along extend muscles The incubated. are larvae the where

two high membranes. The dorsal tubercle has the shape of a coffee-bean with a a with coffee-bean a of shape the has tubercle dorsal The membranes. high two

well developed on the left side of the body; they are very thin on the right side, side, right the on thin very are they body; the of side left the on developed well

short and stout tentacles are located behind a velum. The prepharyngeal band has has band prepharyngeal The velum. a behind located are tentacles stout and short

posterior part, these fibers do not seem to join the longitudinal fibers. They are are They fibers. longitudinal the join to seem not do fibers these part, posterior

mosed network of transverse fibers joining the dorsal longitudinal fibers. In the the In fibers. longitudinal dorsal the joining fibers transverse of network mosed

osl iescn ese.I te neir af fte hrx slctd n anasto an located is thorax the of half anterior the In seen. be can fibers dorsal

We have not been able to establish a muscular formula, the tissues being too thick thick too being tissues the formula, muscular a establish to able been not have We

and contracted. In a specimen opened along the ventral side (Fig. 35C), parallel parallel 35C), (Fig. side ventral the along opened specimen a In contracted. and

meridionalis. meridionalis.

settled close to the base of adult individuals, but no relations exist between between exist relations no but individuals, adult of base the to close settled

u seiesd nt efcl crepn t ay ecito o hs spe these of description any to correspond perfectly not do specimens Our

Remarks: Five species of of species Five Remarks:

This species seems to always have solitary zooids. Sometimes, Sometimes, zooids. solitary have always to seems species This

The abdomen (Figs. 35A, B) is cylindrical and is 3 to 4 mm in diameter. diameter. in mm 4 to 3 is and cylindrical is B) 35A, (Figs. abdomen The

C. C.

Tokioka & Nishikawa, 1976 colonies may occur. Only two species: species: two Only occur. may 1976 colonies Nishikawa, & Tokioka

C. C.

Watanabe Watanabe

meridionalis) meridionalis)

dagysa dagysa

(Herdman, 1891a) 1891a) and (Herdman,

25 mm long, have 3 adhesive papillae in a triangle (Fig. 35D). 35D). (Fig. triangle a in papillae adhesive 3 have long, mm 25

2mm. We counted more than 50 long stigmata in a half row. In the less less the In row. half a in stigmata 50 long than more counted We 2mm.

and and

C. C.

o/iva o/iva

& &

C. C.

or in different seas ( ( seas different in or

Tokioka, 1973 actively buds, but the newly formed zooids zooids formed newly the but buds, 1973 actively Tokioka,

It It

ostrearium ostrearium

has 3 large folds on the right side and the left side is side more left the and side right the on folds large 3 has

and and

Monniot Monniot

C. C.

dagysa dagysa

Clavelina Clavelina

meridionalis meridionalis

C. C.

& &

ostrearium ostrearium

ae caatrsi uiom le oo, re color, blue uniform characteristic a have

Monniot: New Ascidians Ascidians New Monniot:

C. C.

(Kott, 1957), 1957), (Kott,

viola viola

C. C.

are known to have isolated zooids. zooids. isolated have to known are

oliva). oliva).

have variable colors either in a same same a in either colors variable have

has two plain edges, it is lobated in in lobated is it edges, plain two has

(Michaelsen, 1930) 1930) exclusively have (Michaelsen,

After Kott (1990a), the species species the (1990a), Kott After

C. C.

oliva oliva

Kott, 1992b, and and 1992b, Kott,

youn_g youn_g

C. C.

zooids zooids

mer

C. C.

207 207 It It

cm cm for C.

5 5

and and 2.5 to

sensu sensu lato.

species, species, but very difficult or

Tokioka, Tokioka, 1976

In In our opinion, the variability in

. .

& &

has has transverse a musculature mostly

meridionalis meridionalis

Clavelina Clavelina

meridionalis meridionalis

seems to to seems to be closer the posterior end of the

Nishikawa Nishikawa

Fig. Fig. 36, Pl. 6D

contains contains as many issued from fibers the siphons

meridionalis meridionalis

A3 A3 Cla 65-Palau, Koror State, Ngerkuul Pass,

Micronesica Micronesica 29(2), 1996

oliva oliva

meridionalis meridionalis

MNHN: MNHN:

The The cloacal siphon is the longest one. On the upper are side

. .

Clavelina Clavelina obesa?

Nishikawa Nishikawa & Tokioka, Japan. 73, 1976:

species is not not is known, sufficiently species and presently we include the spec

the the stomach located is the in middle of the abdomen marked well with

. .

oliva oliva

The The largest zooid has a thorax 9 mm long and abdomen mm 8.5 long and a

The The colony settled on limestone consists of a cluster of zooids linked by

Geographic Geographic Distribution: distributed Widely around Australia; Indonesia.

Our Our are specimens intermediate between these two for species both the size

The The stomach of C.

The The musculature of C.

Clave/ina Clave/ina

yngeal yngeal band is a protruding single rod. The dorsal tubercle has not been clearly

encircle encircle the cloacal siphon. The tentacles 12 are short and stout. The prephar

the the body. with They the fuse musculature of the opposite along side the endostyle domen. domen. The oral siphon has a strong circular musculature, while very fibers few

(Fig. (Fig. 36A, B). There are no longitudinal either muscles on the thorax or the ab

bundles, bundles, branched and anastomosed. The do muscles not reach the dorsal of line

vascular vascular stolon 6 mm long. The musculature is mostly oblique with 13 to 16

siphons

parent; parent; after fixation it becomes a little irregular with low papillae some near the

lamina lamina and a dark line along the endostyle (Pl. 6D). The tunic perfectly is trans

base base of the cloacal siphon. The branchial sac has a dark line along the dorsal

3 3 dark spots; at the top of the endostyle, above the neural ganglion and at the

due due to gut contents

animals animals are of a pale blue with an irridescent white crescent dorsally around the

Clavelina Clavelina obesa stolons. stolons. The stolons arise from the zooids at any part of the body. In life, the cloacal cloacal siphon 6D). (Pl. The gut appears white in photographs, a color probably

7°19.13'N; 7°19.13'N; m. 134°29.87'E, 10

Material Material Examined:

the the

imens imens of this collection in the C. species

and and anatomical the characters. have anastomoses They the transverse of muscular

specimen. specimen.

folds folds in the type specimen (Kott, fig. 1990a, or 17C) no folds at all in another fibers fibers and this not was described in any species

in in C.

gut gut loop than to the thorax. The stomach has 3 marked folds. On the contrary,

to to be a good character to separate the

impossible impossible to observe in contracted specimens.

these these two there species is no agreement between the muscle formula in given the

text text by Kott and her (1990a) The drawings. seems exact disposition muscles of the

as from as from while the C. ventral side,

issuing issuing from the endostylar area. These characters seem very variable, and for

oliva. oliva.

are are distinguished by their size, 20 cm for C. 208 208

white crescent around the cloacal siphon is not mentioned. The muscle distribu- muscle The mentioned. not is siphon cloacal the around crescent white

at a distance as if they were luminescent"(Nishikawa luminescent"(Nishikawa were they if as distance a at

the zooids were shining in bluish white through the transparent test and looked looked and test transparent the through white bluish in shining were zooids the

Tokioka, 1976 with a few exceptions. The color is a little different: "When alive, alive, "When different: little a is color The exceptions. few a 1976 with Tokioka,

the stomach. stomach. the

None of the zooids have gonads. The heart is voluminous and extends beyond beyond extends and voluminous is heart The gonads. have zooids the of None

not been possible to determine exactly at which level of the thorax the anus opens. opens. anus the level thorax which the of at exactly determine to possible been not

perforated area at the posterior end of the branchial sac (Fig. 36A). 36A). esophagus sac The (Fig. branchial of the end posterior the at area perforated

ach has an irregular outline (Fig. 36C). The intestine is full of sediment and it has has it and sediment is of full intestine The 36C). (Fig. outline irregular an has ach

is cylindrical. The globular stomach has a well-marked typhlosole. The post-stom The typhlosole. a well-marked has stomach globular The is cylindrical.

os fsimt ae eaae b ih rnvre aia. hr i lre un large a is There laminae. transverse high by separated are stigmata of rows

seen. There are 14 sharp dorsal languets, of reduced length posteriorly. The 15 15 The posteriorly. length reduced of languets, dorsal sharp 14 are There seen.

Remarks: This colony has some similarity to to similarity some has colony This Remarks:

Figure 36. 36. Figure

hrx C eal fte stomach the of detail C, thorax;

Clave/ina obesa? obesa? Clave/ina

V1 V1

3 3

Monniot Monniot

Nishikawa Nishikawa

& &

Monniot: New New Ascidians Monniot:

. .

& &

Tokioka, 1976: 1976: Tokioka,

C C

"' "'

., .,

3 3

Clavelina obesa obesa Clavelina

& &

A, B, B, A,

Tokioka 1976, p. 73). A A 73). 1976, p. Tokioka

both sides of the the of sides both

Nishikawa Nishikawa

209 209 & &

species, species, by

Clavelina Clavelina

is is rather posterior at 3

obesa obesa

C. C.

has has only been previously recorded

Kott, Kott, 1990

Indonesia, Indonesia, north Sulawesi, west ofMa

is is said to have 4 longitudinal grooves on the

Micronesica Micronesica 29(2), 1996

Clavelina Clavelina obesa

Figs. Figs. 37A-C, Pl. 6E

but but we are not certain that the swellings of the test

A3 A3 Cla 61

obesa obesa

Clavelina Clavelina robusta

C. C.

obesa, obesa,

C. C.

MNHN: MNHN:

Kott, Kott, 1990a: 61, fig. 20, pl.4g and synonymy.

The The tentacles 24 orders 4 in arise from a high crest. The largest are clearly

23.74'N; 23.74'N; 'E, 124°32'41 m. IO

The The thorax is a little longer than the abdomen. The siphons are close to each

The The colonies from Sulawesi were overhanging in coral reefs mile off 1 shore.

Geographic Geographic Distribution:

The The Palau specimens differ from all other Indo-Pacific

branchial branchial sac, which is expanded into a small pouch.

reach reach the dorsal lamina. The esophagus opening is anterior to the bottom of the

is protruding is protruding and opens in a slit (Fig. 37C). The dorsal lamina is made of large

in in a row. The transverse vessels are high; the posterior one is lower and does not

the the center of a wide unperforated band. There are 20 to 24 long regular stigmata

languets languets in prolongation of the transverse vessels. The dorsal lamina is located in

yngeal yngeal groove has two high blades parallel, without dorsal V. The dorsal tubercle

behind behind the crest where the latter describes deep curves (Fig. 37C). The prephar

where where each makes an anchoring mass in the tunic (Fig. 33B), as in the genus

on on the left side of the abdomen. They form 2 bundles at the base of the abdomen

Diazona. Diazona.

are are branched. The general disposition is characteristic but the number of muscles

the the thorax, the bundles are well formed. Anteriorly and posteriorly, the muscles

siphon, siphon, the others originating from the endostylar region. In the middle part of

varies varies according to the thoracic regions and to the individuals. The muscles extend

37 A) A) is made 37 of parallel longitudinal fibers the half of which comes from the oral

Clavelina Clavelina robusta other, other, the oral one double the width of the cloacal. The thoracic musculature (Fig.

surrounding surrounding the thoraces is soft, but thick.

abdomens abdomens and vascular ampullae. The basal tunic is thick and resistant; that

arated arated from one another, joined basally in a common test which contains the

around around the siphons (Pl. 6E). The zooids, 2 cm in length, are almost entirely sep

Material Material Examined:

The The general color of the zooids was of a dark green, translucent, with yellow rings

nado, nado, l

from from the type station in the Japanese Takegasima Island facing the Kii Channel.

the the muscle distribution, the shortness of the gut and the structure of the colony.

of of the Palau specimens exactly correspond to what the Japanese authors saw.

specific character character specific of

kawa kawa & Tokioka suggested that the small papillae around the siphons may be a

& & Tokioka p. 1976, 73). They were not observed in the present specimen. Nishi

or or rows 4 of stigmata from the esophagus opening, but to it seems more be anterior

these these plications may become much less distinct and almost obscure" (Nishikawa

stomach stomach wall, distinct in young zooids, "but in some larger specimens some of

in in the Palau specimens.

tion tion and the gut shape are identical. The anus of 210 210

are scattered over the whole left side and and overlap slightly onto the right. The The right. the onto slightly overlap and side and left whole the over scattered are

has no folds. The post-stomach is present. The proximal end of the rectum is is rectum the of end proximal The present. is post-stomach The folds. no has

the bottom of the cloaca! cavity in an anus with two scalloped lobes. The massive massive The lobes. scalloped two with anus an in cloaca! cavity the of bottom the

expanded and surrounded by tubules of the pyloric gland. The The gland. pyloric the of tubules by surrounded and expanded

ovary is on the left side just posterior to the stomach. The The stomach. the to posterior side just left the on is ovary

The stomach (Figs. 37 A, B), a little posterior to the middle of the abdomen, abdomen, the of middle the to 37 posterior little (Figs. a B), A, stomach The

iue 37 Figure

area

. .

Clavelina Clavelina

C/avelina C/avelina

sp.: sp.:

robusta robusta

Monniot Monniot

, E D,

ot 1990 Kott,

, ,

both both

& &

Monniot Monniot

sides sides

: :

A, A,

or a a or

zooid; zooid;

New New

zooid

Ascidians Ascidians

B, B,

. .

abdominal abdominal

muscles; muscles;

testes testes

intestine intestine

(Figs. 37 37 B) (Figs. A,

C, C,

dorsal dorsal

ends at at ends 211 211

It It

20. 20.

= =

robusta robusta

C. C.

described described by To

Clave/ina Clave/ina

Tokioka Tokioka (1976), and partic

& &

Tokioka Tokioka & Nishikawa, These 1975.

sp. sp.

eye/us eye/us

C. C.

Clave/ina Clave/ina

Nishikawa Nishikawa (1976) is 4 E.6 to lOB.10D

& &

Micronesica Micronesica 29(2), 1996

Figs. Figs. 37D, E, Pl. 6F

specimens specimens from Palau, whose color was slightly differ

(Oka, (Oka, 1927a) and

elegans elegans

Nishikawa Nishikawa (1975, and 1976), Nishikawa

C. C.

& &

Remarks: Remarks: These specimens resemble the Japanese

Contracted Contracted the zooids reach at most 15 mm in length, with 6 to 7 mm for

The The colony a cluster is of erect zooids slightly linked to a small common base,

Geographical Geographical Distribution: The geographical distribution of

Remarks: Remarks: This sample corresponds well to the description ofTokioka (1967)

Clavelina moluccensis, Clavelina moluccensis,

ularly ularly

kioka kioka

testis testis follicles extending over the intestine and a part of the stomach. The volu

37D). 37D). They consist of a slightly protruding ovary surrounded by a thin layer of

minous minous heart is ventral. Some eggs are incubated in the right side of the cloacal

and and has no pre-stomach. Only the typholosole visible is on the globular stomach.

cavity. cavity. The larvae were not yet developed.

on on the dorsal side of the body, and some on the right side of the gut loop (Fig.

30 rows 30 rows of stigmata are separated by transverse thick and high crests. The rapheal

The The anus opens at the bottom of the cloacal cavity. The gonads are mostly located

too too contracted to accurately determine the oral tentacles and neural area. About

languets languets are blunt and short. The esophagus long is (half of the abdomen length)

does does not seem identical on both sides of the body wall. The thoracic contraction

to to the terminology of Tokioka

cles cles extend onto the abdomen (Figs. 37D, E). The anterior part of the body was on on each side, more crowded ventrally. The muscles formula established according

the the thorax and 9 mm for the abdomen. At the base of the colony, the tunic has

makes makes a distinction between branchial and dorsal muscles difficult. Thoracic mus

swellings swellings filled with vascular ampullae. There are about 20 longitudinal muscles

the the zooids were steel blue and opaque, the pigment being located in the body wall.

siphon siphon smaller, is more posterior. The gold-yellow color absent is above the neural

Each Each siphon has a golden ring, wider on the oral siphon (Pl. 6F). The cloacal

brown-purple brown-purple and opaque anteriorly.

ganglion. ganglion. After fixation the yellow color disappears and the body wall becomes

7°19.13'N; 7°19.13'N; 134°29.87'£, 3 m.

generation. generation. No extensions of the adult zooids penetrate this base. When living,

Material Material examined: MNHN: A3 Cla 66-Palau, Rock Islands, Ngerkuul Pass,

but but sometimes some individuals arise from the tunic of a zooid of the preceding

for for

Philippines Philippines were black with fluorescent green rings around the siphons.

atrial atrial apertures a very bright yellow". Kott (1990a) noted that specimens from

cludes cludes Japan, Northern and Western Australia, Palau, Philippines and Indonesia.

ent ent "body dark blue and anterior portion of zooid surrounding branchial and

of of this collection, the larvae are not fully developed and none are in the cloacal

incubation incubation of the larvae begins in the distal part of the oviduct. In the specimens

cavity. cavity. 212 212

previously described for the species, except for the color of the colony, very dark dark very colony, the of color the for except species, the for described previously

The trunk size is 1 mm, but contracted. contracted. but 1 mm, is size trunk The

most differentiated we have found has 3 triradially arranged adhesive papillae. papillae. adhesive arranged triradially 3 has found have we differentiated most

mature embryo is incubated in the posterior part of the thorax in each zooid. The The zooid. each in thorax the of part posterior the in incubated is embryo mature

h suid ooy te vr i oae a te otm f h gt op A im An loop. gut the of bottom the at located is ovary the colony, studied the

longitudinal folds probably due to contraction. The male gonads were absent in in absent were gonads male The contraction. to due probably folds longitudinal

domen is longer than the thorax. The stomach is posterior, olive-shaped, with with olive-shaped, posterior, is stomach The thorax. the than longer is domen cular bundles originate from the oral siphon on each side; some join the muscular muscular the join some side; each on siphon oral the from originate bundles cular

tgaa n ah ie Te rnvre iue pses ih ebae. h ab The membranes. high possess sinuses transverse The side. each on stigmata

tentacles are on a line. The branchial sac comprises 24 to 25 rows of about 50 50 about of rows 25 to 24 comprises sac branchial The line. a on are tentacles

All All

ribbon along the endostyle. Other muscular fibers start from the cloacal siphon. siphon. cloacal the from start fibers muscular Other endostyle. the along ribbon

esrs 5m a lat Te ihn hv a idnae mri. tog mus Strong margin. indentated an have siphons The least. at mm 15 measures

acetic acid, polarize the light and may be made of calcium oxalate. oxalate. calcium of made be may and light the polarize acid, acetic

the sides of the lobes, discoid crystals are scattered. They are not easily soluble in in soluble easily not are They scattered. are crystals discoid lobes, the of sides the

zooids. Orange pigment cells are also present at the base of the oral lobes and in in and lobes oral the of base the at present also are cells pigment Orange zooids. top of small protrusions. The bodies are perpendicular to the colony surface and and surface colony the to perpendicular are bodies The protrusions. small of top

from 4 to 5 cm in height and I to 2 cm for the diameter in the upper part. The The part. upper the in diameter the for cm 2 to I and height in cm 5 to 4 from

the branchial tissue in preserved specimens. In the superficial layer of the test, on on test, the of layer superficial the In specimens. preserved in tissue branchial the

in alcohol, but many dark purple cells persist in the test, fewer in discolored discolored in fewer test, the in persist cells purple dark many but alcohol, in

translucent test has a dark purple color, the same as for zooids. This stain fades fades stain This zooids. for as same the color, purple dark a has test translucent

ao l nado,

Material Examined: Examined: Material

Stomozoa roseola: roseola: Stomozoa

Stomozoa murrayi murrayi Stomozoa

parent and cobalt blue, and becomes purplish in the fixative. fixative. the in purplish becomes and blue, cobalt and parent

cloacal cavity. cavity. cloacal

ring at the base of the oral siphon, and not on its rim. rim. its on not and siphon, oral the of base the at ring

species (see Nishikawa & Tokioka 1976, pl. l) translucent with a narrow yellow yellow narrow a with translucent l) pl. 1976, Tokioka & Nishikawa (see species

ular. The appearance of the living living the of appearance The ular.

Clavelina roseo/a roseo/a Clavelina

incubates its larvae in the terminal part of the oviduct or at the bottom of the the of bottom the at or oviduct the of part terminal the in larvae its incubates

mens (30) than in in than (30) mens

in in

(more numerous in the Palau species: 20 instead of IO to 13 13 in IO to of instead 20 species: Palau the in numerous (more

e/egans e/egans

species have in common the shape of the colony (except that the zooids for for zooids the that (except colony the of shape the common in have species

C. C.

muscles are gathered in a longitudinal ribbon along the abdomen. The oral oral The abdomen. the along ribbon longitudinal a in gathered are muscles

Remarks: The characters of the present specimens correspond well to those those to well correspond specimens present the of characters The Remarks:

The zooids apertures, at the colony surface, are seen as 2 fringed holes at the the at holes fringed 2 as seen are surface, colony the at apertures, zooids The

The colonies consist of several conical lobes, joined at their base, measuring measuring base, their at joined lobes, conical several of consist colonies The

We do not have enough material of the present species to be able to name it. it. name to able be to species present the of material enough have not do We

eye/us). eye/us).

have common stalks) and the same general distribution of the muscles muscles the of distribution general same the and stalks) common have

23.74'N; 124°32.4l'E, 20 m. m. 20 124°32.4l'E, 23.74'N;

h nme o os fsimt i lo agr nte rsn speci present the in larger also is stigmata of rows of number The

It It

always has a striated stomach, that of of that stomach, striated a has always

Millar, 1955: 139, South Africa. Africa. South 139, 1955: Millar,

Kott, Kott,

C. C.

Kott, 1957b: 131 fig. 2-8, Red Sea. Sea. Red 2-8, 131 fig. 1957b: Kott,

MNHN: MNHN:

eye/us eye/us

Stomozoa roseo/a roseo/a Stomozoa

1990a: 157 and synonymy. synonymy. 157 and 1990a:

Monniot Monniot

(maximum 22) or in in or 22) (maximum

A3 Sto 4-Indonesia, north Sulawesi, west of Ma of west Sulawesi, north 4-Indonesia, Sto A3

Family Holozoidae Holozoidae Family

& &

Monniot: New Ascidians Ascidians New Monniot:

C. C.

eye/us eye/us

(Millar, 1955) 1955) (Millar,

is clearly different from the Palau Palau the from different clearly is

C. C.

elegans elegans

C. C.

e/egans e/egans

elegans elegans

(16 to 30). 30). to (16

C. C.

e/egans e/egans

sqie trans quite is

smr irreg more is

C. eye/us eye/us C.

and 15 15 and

213 213 C. C.

. .

N; N;

Sig

m;-Fed

5 5

. .

(Sluiter, (Sluiter, 1909)

Pl. Pl. 6G

Sigillina Sigillina

5 5

Fig. Fig. 38A, Pl. 6H

Micronesica Micronesica 29(2), 1996

A3 A3 2, Sig 4 and 5-Palau, Koror State, Ngerkuul

Sigil/ina Sigil/ina signifera

MNHN: MNHN:

Sluiter, Sluiter, 1909:

Kott, Kott, and pl.8c-d fig.34, synonymy. 100, 1990a:

51.52'N; 51.52'N; m. 18 125°03.84'E,

species was collected in Indonesia as most species of this genus inhabit inhabit genus colder of this Indonesia most in species as collected was species

Without Without gonads and larvae it is not possible to identify animals having so

The The colonies are stalked (Pl. 6H). According to the collector, the heads of The The zooids are all immature in the colony observed. Both apertures are sim

Geographic Distribution: The species was already already known Geographic Palau from was Distribution: Islands, The species

The The colonies are dark green or dark blue (Pl. 6G) with a transparent tunic

il/ina il/ina

observed. observed. zooid Some buds growing were in the colony head, to close the stalk.

raised raised in high membranes. The abdomen is narrower than the thorax. It is short

with with a narrow loop. digestive The stomach has a smooth wall. Gonads were not waters. waters.

(Fig. (Fig. they 38A), are not cut by parastigmatic The sinuses. transverse sinuses are varied varied shapes and colors. Nevertheless it is interesting to note that another

row row is slightly curved dorsally. About 50 stigmata were counted in a half row

151°50.50'E, intertidal. intertidal. 151°50.50'E,

ilar ilar with 6 lobes. The thorax (Fig. wider is 38A) than high and the first stigmata

distribution. distribution. The oral siphons are more posterior than the cloacal ones.

the the different colonies had diverse colors: white, blue, purple, orange and peach

Island, Island, l

vascular vascular appendages into extend the in stalk. observed zooid the No were systems Material Material Examined: MNHN: I-Indonesia, Sig A3 ofTalisei west north Sulawesi,

Their size is from from is Their 4 size to cm in height. The 12 zooids are in the head the while long

Indonesia, Indonesia, Philippines and the tropical eastern and western coasts of Australia.

prolongation prolongation is well muscularized. Larvae being incubated in a thoracic brood

pouch pouch are not fully developed in the Palau colony.

ment ment with the previous descriptions Kott (see The 1990a). long post-abdominal

Material Material Examined: and and the dark color of the zooids remains All characters in fixatives. are in agree

erated erated of States Micronesia, Chuuk lagoon, Moen S. Island, sand flat, 7°24.60

Pass, Pass, 7°19.13'N; 7 m; 134°29.87'E, Stone Reef, 7°1l'N; 134°22.3'E,

Sigil/ina Sigil/ina signifera:

Polycitor Polycitor signiferus

species is cosmopolitan, cosmopolitan, is Indonesia in its presence so species not is surprising. The known

New New Caledonia.

distribution distribution comprises Brazil, Guyana, Red Sea, South Africa, Indonesia, and

here. here. The siphons, muscles and number of rows of stigmata are the same. The 214 214

Figure 38. 38. Figure

Sigillina Sigillina

.: .:

A, zooid A,

Monniot Monniot

. .

Distaplia regina regina Distaplia

& &

Monniot: New Ascidians Ascidians New Monniot:

ot 1990 Kott,

: :

B, zooid; C, detail of the gut the of detail C, zooid; B,

. . 215 215

Oka, Oka, 1926)

(Syndiazona (Syndiazona

Palau, Palau, Rock Islands, 7°20.28'N;

our our

Kott, Kott, 1990

Queensland, Queensland, Australia.

In In confused opinion, were 3 species

44e, 44e,

Kott, Kott, However 1990. these pale colonies are

A3 A3 Dis 66

Savigny, Savigny, 1816

Monniot Monniot One (1989). specimen was described by

Family Family Diazonidae

Micronesica Micronesica 29(2), 1996

from from an immature colony. other Six colonies were

Figs. 38B, C, Pl. 7 A A Figs. C, Pl. 38B, 7

Order Order Phlebobranchia

& &

S.? S.? chinensis.

Distaplia Distaplia regina?

or or

Diazona Diazona

MNHN: MNHN:

Distaplia Distaplia regina

(Oka, 1926) from from Japan (Oka, and eastern 1926) China species second A Sea.

Kott, Kott, fig. 125, 1990a:

S. S. chinensis

The The genus

Syndiazona Syndiazona chinensis

The The best known and most abundant from species the Indo-Pacific Ocean is

Previously the species was was known Previously the species only from the Great Barrier Reef.

Remarks: Remarks: The number of stigmata and the stomach correspond wall to well

The digestive tract tract digestive The a forms round the which gonads loop located are in (Figs.

The The zooids measure 4 mm in length. The oral siphon has 6 low lobes. The

The colonies are large, and and are large, colonies encrust The length coral branches. cm in 9 They reach

to to this species by Monniot

in in Millar's collection. Three other colonies from the Philippines were attributed

described as as described

studied studied by Millar from (1975) the Philippines and Indonesia (Kei Islands) and

name name of

was described from described was the eastern part of the China Sea by Tokioka with the (1955)

Diazona Diazona grandis

would would be useful in the cofirming identification.

the the description of

characters characters very No similar. seems larvae are known in any of the samples. They

(Fig. (Fig. No 38C). larvae present. were

different different from the dark purple Australian specimens. Otherwise, all anatomical

brood brood pouch begins to protrude on the right side of the bottom of the thorax

opens opens near the anus at of the level the second transverse sinus. In some zooids a

tions tions which seem due more to the gut content than to well-marked anatomical

38B, C). The The C). 38B, stomach is rather small, asymmetrical; its areolated. wall is After a

The The ovary not is developed in well the center of the male gonad. The sperm duct

short short narrow segment, the intestine remains constric cylindrical with successive

compartments. compartments. There are about testis 10 in follicles the center of the gut loop.

ring ring of two alternating orders. The branchial sac has 4 rows of 20 to 25 long

guets are guets narrow and displaced to progressively the left side.

134°26.71 'E, 'E, m. 33 134°26.71

stigmata stigmata on each side, but no parastigmatic (Fig. The vessels 38B). rapheal lan

upper rim of the cloacal siphon is produced in a wide languet (Fig. 38B) of variable variable of 38B) languet (Fig. wide a produced in is siphon cloacal the of upper rim

shape shape which contains transverse muscle fibers. There are oral 12 tentacles on a

vae vae are not present.

slightly protruding protruding slightly adheres cloacal opening. silt Some to the colony surface. Lar

and and about 1 cm in thickness. They are very soft, gelatinous, transparent with

yellow yellow zooids A). (Pl. The 7 zooids are arranged in circular systems around a

Material Material Examined:

Distaplia Distaplia regina 216 216

curved stripes according to the individuals. The intestinal regions are well well are defined regions intestinal The individuals. the to according stripes curved

esophago-rectal peduncle is rather long, often narrow (Fig. 39A) but sometimes sometimes but 39A) (Fig. narrow often long, rather is peduncle esophago-rectal

the same diameter as the stomach region. The stomach is long with straight or or straight with long is stomach The region. stomach the as diameter same the

Most of the sinuses are entire. There is about 2 long stigmata in a mesh. The The mesh. a in stigmata long 2 about is There entire. are sinuses the of Most

branchial sac has 38 rows of stigmata and 24 longitudinal vessels on each side. side. each on vessels longitudinal 24 and stigmata of rows 38 has sac branchial

languets are sharp and long, located on the left side of an unperforated area. The The area. unperforated an of side left the on located long, and sharp are languets

indentation.The flat dorsal tubercle has a slightly elongated opening. The rapheal rapheal The opening. elongated slightly a has tubercle dorsal flat indentation.The

orders of size. The prepharyngeal groove is a flat wide ribbon, with a dorsal dorsal a with ribbon, wide flat a is groove prepharyngeal The size. of orders

name. About 10 long and curled tentacles are inserted on a single ring but in two two in but ring single a on inserted are tentacles curled 10 and long About name.

either extend onto a vascular process starting from the right side of the abdomen abdomen the of side right the from starting process vascular a onto extend either

endostyle where they branch before joining a strong longitudinal muscle. The The muscle. longitudinal strong a joining before branch they where endostyle

in a short appendix (Fig. 39B). This shape of the thorax is described in the species species the in is described thorax the of 39B). shape This (Fig. appendix short a in

where they join the ventral bands to form the abdominal ribbons. The muscles muscles The ribbons. abdominal the form to bands ventral the join they where

or stop abruptly at its proximal end (Fig. 39D). The endostyle projects posteriorly posteriorly projects endostyle The 39D). (Fig. end its proximal at abruptly stop or

siphons (Figs. 39A, 39A, (Figs. siphons

bands from the cloaca! siphon extend down to the bottom of the branchial sac sac branchial the of bottom the to down extend siphon cloaca! the from bands

lowish. Zooids project slightly from the upper surface of the colony. colony. the of surface upper the from slightly project Zooids lowish.

overhanged in colorless translucent balls (Pl. 7B). In formalin they become yel become they formalin In 7B). (Pl. balls translucent colorless in overhanged

gulata, D. carnosa, carnosa, D. gulata, m and Apo Island, 16 m (N. Lindquist collection). collection). Lindquist (N. 16 m Island, Apo and m

Holotype: MNHN: MNHN: Holotype:

in other species, variable to long in in long to species, variable other in

in in

shown in the figures. The abdominal muscles end aside from the vascular process process vascular the from aside end muscles abdominal The figures. the in shown thea thea

D. labyrinthea. labyrinthea. D.

thoracic muscles are mostly longitudinal, whereas they are mostly transverse in in transverse mostly are they whereas longitudinal, mostly are muscles thoracic

easily distinguished by the following features: features: following the by distinguished easily

lamellae. lamellae.

the thoraces are also included but the colony surface is raised in a network of of network a in raised is surface colony the but included also are thoraces the

sive colonies with the thorax entirely included in the tunic mass. In In mass. tunic the in included entirely thorax the sive with colonies

Nishikawa (1991) from the Japan Sea. A third species, species, third A Sea. Japan the from (1991) Nishikawa

niot, 1987a was described from several New Caledonian colonies. colonies. Caledonian New several from described 1987a was niot,

specimens of the genus were reexamined for this present work. The species are are species The work. present this for reexamined were genus the of specimens

D. formosa, D. tenera tenera D. formosa, D.

The musculature consists of about 18 longitudinal ribbons issued from the the from issued ribbons 18 longitudinal about of consists musculature The

Description: The largest colony was 30 x 10 cm and IO IO animals 10 30 was x colony The thick. and cm largest cm The Description:

-The esophago-rectal narrowing is very short in in short is very narrowing esophago-rectal -The

-The muscle distribution. In In distribution. muscle -The

and and

There are 6 species in this collection, five of them are new. All available available All new. are them of five collection, this in species 6 are There

The colony structure. structure. colony The

. tenera D.

D. D. formosa

In In

. .

D. formosa formosa D.

and and

Pl Pl

B). B).

and and

i 18 Dia

Those from the oral siphon extend obliquely toward the the toward obliquely extend siphon oral the from Those

D. chinensis. chinensis. D.

Diazona angulata angulata Diazona

and and

D. tenera tenera D.

Monniot Monniot

D. angulata, D. carnosa, carnosa, D. angulata, D.

Figs. 39A, D, Pl. 7B 7B Pl. D, 39A, Figs.

D. labyrinthea labyrinthea D.

and and

Philippines, Siguijor Island, Tambison point, 18 18 point, Tambison Island, Siguijor Philippines,

& &

D. angulata, D. carnosa, carnosa, D. angulata, D.

D. D. formosa,

Monniot: New Ascidians Ascidians New Monniot:

have thoraces entirely free. free. entirely thoraces have

D. tenera tenera D.

new species species new

but are prolongated in it in in it in prolongated are but

the muscles have a special design, design, special a have muscles the

and especially long in in long especially and

and and

D. chinensis, chinensis, D.

Diazona textura textura Diazona

D. chinensis chinensis D.

and and

D. chinensis, chinensis, D.

D. /abyrinthea /abyrinthea D.

but elongated elongated but

D. labyrin D.

have mas have

C. Mon C.

. an D.

the the 217 217

Diazona carnosa carnosa Diazona

C, C, D, abdomens.

mm mm

2 2

muscles; muscles;

thoracic thoracic

, ,

abdomen. abdomen.

Micronesica Micronesica 29(2), 1996

the the

n. sp.: A, zooid; zooid; A, sp.: n.

sides sides of

both both

Diazona angulata angulata Diazona

n. n. sp.: E, zooid; F, G,

Figure Figure 39. 218 218

gate stomach has a few well marked longitudinal folds (no more than 9 on each each on 9 than more (no folds longitudinal few marked well a has stomach gate opens by a plain margin of the anus, at the level of the tenth or eleventh row of of row eleventh or tenth the level of the at anus, the of margin plain a by opens

side with some irregularities) (Fig. 39G). The post-stomach is present. The rectum rectum The is present. post-stomach The 39G). (Fig. side irregularities) some with

complete as in the two on each side of the dorsal lamina and the one on each side side each on one the and lamina dorsal the of side each on two the in as complete

mata divisions. There are 21 to 25 longitudinal vessels which are sometimes in sometimes are vessels which longitudinal 25 to 21 are There divisions. mata of the endostyle. In a mesh, there are 2 not very long stigmata. stigmata. long very not 2 are there mesh, a In endostyle. the of

parastigmatic vessel. The branchial sac has generally 24 rows of stigmata, but we we but stigmata, of rows 24 generally sac has branchial vessel. The parastigmatic

observed, above the first and last rows, a parastigmatic vessel and figures of stig of figures vessel and parastigmatic a rows, last and first the above observed,

the dorsal lamina has 19 languets. The first one appears at the level of the second second level the the of at appears one first The 19 languets. has lamina dorsal the

curve. The olive-shaped neural ganglion is closer to the cloacal siphon than to the the to than siphon cloacal the is to closer ganglion neural olive-shaped The curve. pn na u-hpd tubercle um-shaped an in opens

oral siphon. The neural gland is on the ventral side of the neural ganglion and and ganglion neural the of side ventral the on is gland neural The siphon. oral

down the siphon. The prepharyngeal band has 2 parallel blades without dorsal dorsal without blades parallel 2 has band prepharyngeal The siphon. the down

the lobes the

The transverse musculature is only slightly developed on the cloacal siphon and and siphon cloacal the on developed slightly only is musculature transverse The So, the body appears to have 3 processes: one vascular and two fibrous two and vascular one 3 processes: have to appears body the So,

base of the body and they are prolongated by elastic fibers in the common tunic. tunic. common the fibers in elastic by prolongated are they and body the of base

the upper part of the thorax. Both muscular abdominal bundles protrude at the the at protrude bundles abdominal muscular Both thorax. the of part upper the

is anastomosed with the longitudinal fibers. There is a single transverse muscle in in muscle single transverse is a fibers. There longitudinal the with is anastomosed ec tebs ftebacil a n mk teadmnl uclr bundles muscular abdominal the make and sac branchial the of base the reach

ogrta tezod Te hrcc ucltr (Fig musculature thoracic The zooid. the than longer

tudinal bands. Half of them ends against the endostyle while the most dorsal ones ones dorsal while most endostyle the the against ends of them Half bands. tudinal

slightly protruding. protruding. slightly

formalin. The zooids are completely included in the common tunic. The siphons siphons The tunic. common the in included completely are zooids The formalin.

and 2 cm in thickness. thickness. in cm 2 and open independantly at the colony surface without order; they are not, or only only or not, are they order; without surface colony the at independantly open

wall, 10°15.62'N; 123°59.ll'E, 53 53 m. 123°59.ll'E, 10°15.62'N; wall,

Holotype

colonies are different, in in different, are colonies

colony surface. surface. colony

differs by the thoracic muscles more numerous and less anastomosed than in in than less anastomosed and numerous more muscles thoracic the by differs rectal waist. The prolongation of the endostyle in the postero-ventral corner of of corner postero-ventral the in endostyle the of prolongation The waist. rectal

angulata; angulata; the thorax has never been observed in in observed been never has thorax the

No zooids had gonads had zooids No

(Figs. 39C, D). The bilabiated anus opens at the level of the 16th row of stigmata. of 16th stigmata. row level the of the at opens anus bilabiated The D). 39C, (Figs.

The abdomen is longer than the thorax and expanded posteriorly. The elon The posteriorly. expanded and thorax the than is longer abdomen The

The siphons are well apart. They have 6 lobes with a pigment spot between between spot pigment a with lobes 6 have They well apart. are siphons The

The zooids are less than 1cm in length, but the vascular appendage may be be may appendage vascular the but length, 1cm in than less are zooids The

Description: The colonies form large masses of 10 cm and more in diameter diameter in more 10 and cm of masses large form colonies The Description:

Remarks: This species species seems This to closely related Remarks:

. .

: :

The 12 oral tentacles, in two orders, are on a protruding crest half way way half crest protruding a on are orders, two in tentacles, 12 oral The

h psto o te aclr tln n te rsneo a esophago an of presence the and stolon vascular the of position the

MNHN: Pl Dia 26--Philippines, Cebu, south east Mactan Island, Island, Mactan east south Cebu, 26--Philippines, Dia Pl MNHN:

. .

In In

Diazona carnosa carnosa Diazona

Monniot & Monniot: New Ascidians Ascidians New Monniot: & Monniot

life the color is pink (Pl. 7C) and it disappeared in in disappeared it and 7C) (Pl. pink is color the life

. .

angulata angulata

Figs. 39E, G, Pl. 7C 7C Pl. G, 39E, Figs.

. .

For a branchial sac with 24 rows of stigmata, stigmata, of rows 24 with sac branchial a For

the zooids make obvious swellings at the the at swellings obvious make zooids the

D. chinensis. chinensis. D.

new species species new

Diazona chinensis. chinensis. Diazona

The aspect and color of the the of color and aspect The

. .

39E) has about 25 longi 25 about has 39E)

The species species The

. .

219 219

. . . .

The The two abdominal bundles lie

are are globular massive, and carti

C). C).

C) C)

(Tokioka, (Tokioka, 1955)

Fig. Fig. 40, Pl. 7D

Micronesica Micronesica 29(2), 1996

Diazona chinensis chinensis Diazona

23.52'N; 23.52'N; 124°32.57'£, 43 m and north tip of mouth Sulawesi, of

The The colonies (Figs. Philippines 40D-G) damaged were during the collection

The The Indonesian colonies (Figs. 40A, B,

oral oral siphon. In zooids of the same colony, the number of tentacles varies; they

translucent translucent and The soft. very and long thin tentacles often outburst through the

10th row row 10th of stigmata. A large part of the right side of the abdomen is occupied was also also a was break the between abdomen and the vascular appendage. The test was

test. test. All zooids were broken at the of level the esophago-rectal peduncle. There

the the intestine and the heart, the latter posterior on the left of side the abdomen. with with a trawl. The colonies seem massive with zooids embedded in the common

in in the posterior part of the abdomen (Fig. The 40B). testes are scattered around

by by a mass of globules reminiscent of oocytes, sometimes extended as far as the

anus. anus. The true with recognizable their oocytes, are refringent located cells, thecal

of the intestine are well defined. The The defined. anus opens the of are well intestine approximately at of the the level

rapidly widens posteriorly. posteriorly. short, The is rapidly esophagus the widens height. a with length twice

The The stomach has many irregular plications (Fig. 40B). The segments successive

large large flat papillae. There are 2 to 3 stigmata in a mesh. Some rows of stigmata

about about 25 stigmata on each side. The longitudinal are vessels thin, supported by were seen to be dividing. The The to dividing. seen be were abdomen with a begins pronounced narrowing and

developed developed on the right side. The branchial sac has 25 to 36 stigmata rows with

22 to to and long thin 22 30 languets are located on an unperforated area and are more

ing. ing. The neural is located complex behind the dorsal tubercle. On the dorsal line

on the prepharyngeal ring. The dorsal tubercle is urn-shaped, open circular urn-shaped, a with is The prepharyngeal dorsal tubercle on ring. the

two equal rims. There is a a In dorsal deep is V. There are fixed parasitic colony, one two equal rims. ciliates

the oral siphon. Their bases are not joined. The prepharyngeal groove lies between between joined. The prepharyngeal are lies not groove bases oral Their the siphon.

(Fig. 40A). Sixteen tentacles tentacles counted (Fig. were 40A). in Sixteen sizes 3 along in concentric 3 circles

on the left side and and extend on far side the left very along the post abdominal vascular appendage

whatever whatever the thorax contraction (Figs. 40A,

on on each oral The lobe. musculature has a characteristic the which is design same

In life, one colony was white white the was one colony In other purplish-brown life, 7D). (Pl. In formalin both

lobes, lobes, open independently at the colony surface and only make small swellings. colonies colonies with are whitish brown zooids. In one colony, the zooids had a red spot

lagineous. lagineous. The zooids are embedded completely in the test. Both siphons, with 6

of of Manado, l central central part of Luzon Island, 122°45.9'£, 13°15.3'N; m 166-172

Material Material MNHN: Examined: Pl and 23,-Indonesia, Dia 21 west north Sulawesi, Teluk Teluk Korakora °43.97'N; Inlet, 124°58.03'£, 1 33 m.-Philippines, east of the

right right The side. vascular process thin always is and long.

the anus. the anus. heart The voluminous is and the reaches post-stomach anteriorly on the

the the oviduct full was of globules as "pseudo-oocytes", up to its extremity against

stigmata. stigmata. The gonads were not developed fully in the colony. In the figured ab

domen domen (Fig. there 39F) are In by only circled another oocytes thecal cells. zooid, 220 220

contr

D, thorax; thorax; D,

Figure Figure

acted acted

40. 40.

E, E,

thorax; thorax;

Diazona Diazona

F. F.

both both

, ,

other other

d1ine11sis d1ine11sis

sides of an an of sides

side of of side

(

Tokioka

Monniot Monniot

abdomen

h boe; ,aohr thorax another C, abdomen; the

, ,

1955): 1955):

& &

; ;

, abdom G,

Monniot

A, A,

: :

New Ascidians Ascidians New

, ,

n n

of of

, ,

Indon

an an

immature zooid. zooid. immature

esia

; ;

D D

n n

to to

speci

G, Philippines Philippines G,

men

s: s:

A, A,

zooid zooid

cimens: cimens:

ith ith

221 221 a a

Syndiazona Syndiazona

is confirmed, is confirmed, although

B) B)

from from st. m 25 34, (Millar's 4) is fig.

were were not detected either in the type

new new species

B) B)

They They correspond A, to 42 Millar's figures

chinensis chinensis

? ?

and and the from specimen st. m, 54, 85 identified

Fig. Fig. 41, Pl. 7E

Micronesica Micronesica 29(2), 1996

n. n. sp.

are are damaged but appear to belong to another species,

Diazona Diazona formosa

found found corresponding specimens to the 42 figures Band C

Syndiazona Syndiazona

Diazona Diazona chinensis.

not not

chinensis, chinensis,

? ?

h~ve h~ve

Syndiazona Syndiazona chinensis,

Diazona Diazona carnosa

Tokioka, Tokioka, from 1955 the Seto Marine Laboratory Biological collection.

Description: The colony (Pl. 7E) is branched branched and From is massive. 7E) The colony mass a basal (Pl. Description:

Geographic Geographic Distribution: Previously known from eastern China Sea, Phil

We We have reexamined all described specimens by Millar They (1975). corre

Remarks: Remarks: We have reexamined zooids from the type colony of

Syndiazona Syndiazona

23.50'N; 23.50'N; 124°44.60'£, m. 20

basal test is thick, hard, yellowish, somewhat somewhat hard, thick, translucent. thin, The basal is yellowish, thoracic is test test

2 to to 2 zooids (Fig. 41A). The 10 thoraces are partially or completely isolated. The

large lobes about 2 cm in diameter irregularly divide into terminal lobes containing containing into terminal lobes divide irregularly diameter in cm about 2 lobes large

Holotype: Holotype: MNHN: Pl Dia 24-Indonesia, north Sulawesi, west of Manado,

ippines, ippines, and Japan Sea.

specimen identified as as identified specimen

perhaps perhaps to

dehydrated dehydrated and cannot be used; however it clearly belongs to another species,

identified as as identified

with with a different musculature (large bundles perpendicular to the endostyle). The

as as

structure was observed by us on some broken specimens from Philippines. Millar's Millar's from Philippines. broken on some specimens us by observed structure was

which is totally totally dehydrated is which and cannot from Mindanao, The studied. specimen be

figures figures 42B and C may correspond to a specimen collected at Doe Roa, m 30

D, D, F, E. We

showing showing muscles abruptly interrupted on the side of the abdomen. The latter

spond spond to distinct The 3 from species. specimens Kei Islands, Nochoc Roa, m 35

belong belong to the species

and and Doe Roa, m, 30 and a from specimen Doe Roa, m 25 not studied by Millar,

specimen, in in our specimen, study, nor by Nishikawa (1991).

transverse transverse (Tokioka's sinuses pl. fig. 11,

he he overlooked the transverse muscle at the base of the oral siphon. The large

chinensis chinensis

The The musculature shown by Tokioka (1955, p. fig. 11,

luminous luminous mass can extend up to the anus level.

as as "pseudo-oocytes", are very numerous in Philippines specimens and their vo

in a mesh. The gonads are well developed (Figs. 40E, F). Some oocytes were found found were F). oocytes 40E, Some (Figs. developed The gonads are well mesh. a in

Japan, also noted the presence of oocytes in the vascular vascular appendage. the in The globules, Japan, oocytes of noted the presence also

inside inside the vascular appendage. Nishikawa in (1991), a specimen from southern

In a single colony, some zooids have almost entire sinuses, when other other when only zooids almost entire zooids have sinuses, some colony, a In single

sac has has sac to of rows 30 32 stigmata with about on longitudinal each side. 22 vessels

have have T-shaped papillae, linked sometimes two by two. There about is 2 stigmata

two two orders and an anterior ones. The dorsal of ring small at may least languets 12

be very long and and long very be sharp, reaching the double the of stigmata height. The branchial

are are located in two a circles: posterior ring with large 12 tentacles alternating in 222 222

n abdomen an

ure ure

41. 41.

Dia

. .

zonaformosa zonaformosa

n. n.

Monniot Monniot

.: .:

, ,

two two

& &

zoo

Monniot: New Ascidians Ascidians New Monniot:

s; s;

, ,

C, both both C,

des des

of a thorax; thorax; a of

D, E, both sides of of sides both E, D, 223 223

new species new species

The The muscles are interrupted over the dorsal

C). C).

Figs. Figs. 42A-C, Pl. 7F

Micronesica Micronesica 29(2), 1996

Diazona Diazona /ahyrinthea

Description: Description: The colony is a firm mass cm 2 thick, with a raised network of

The The voluminous heart U-shaped is the in posterior part the of abdomen. The

The The gonads are not A protruding developed. pad gonadal of occurs globules

The The abdomens are to often the 2/3 nearly length twice the of thorax. The gut

The The tentacles are located in a line on a crest in the anterior part of the oral

Both Both siphons have 6 sharp lobes. A white spot is at the top of each lobe.

23.52'N; 23.52'N; m. 33 124°32.64'E,

thick thick lamellae arranged irregularly around deep name) cavities (species (Pl. 7F).

The The apertures, in double rows, open independently at the top of the lamellae.

Holotype: Holotype: MNHN: Pl Dia 20-Indonesia, north Sulawesi, west of Manado,

body body extended is by a long vascular appendage.

oped. oped.

intestine. intestine. zooids Some have full sperm ducts even when the testes are not devel

on on the left side of the intestinal loop (Fig. 41E). The testis is diffuse above the

pharynx pharynx to the bilabiate anus at the base of the cloacal siphon.

loop is inflated posteriorly posteriorly 41D, The smooth; long stomach E). inflated (Figs. narrows it loop is is

gradually gradually to posteriorly. the intestine The long and rectum is alongside extend the

appear, terminate against the endostyle. Posteriorly, the longitudinal vessels some appear, Posteriorly, terminate the endostyle. longitudinal against vessels the

times times are interrupted. There are or 3 4 elongate stigmata in a mesh.

tubercle tubercle area. Anteriorly about counted longitudinal were 35 on vessels each side

rows rows of stigmata. Two or 3 additional rows occur in the vicinity of the dorsal

in in the anterior area, and only 20 are present posteriorly. The which vessels dis

longitudinal longitudinal The slit. neural ganglion globular, is located the below dorsal tuber

tend tend along the whole thoracic dorsal line. The branchial sac has more than 50

cle. cle. Long and thin rapheal Ianguets, corresponding to the transverse ex vessels,

smaller smaller ones between them. The prepharyngeal band forms a high blade, very

slightly slightly indentated dorsally. The dorsal tubercle is large and protruding with a

siphon. siphon. There are about large 10 and curved ones in 2 or 3 orders of size, and

onto onto the vascular stolon.

thorax, thorax, and along extend the abdomen in two thin bundles, that do not penetrate

lamina lamina and the endostyle. The fibers become weak in the posterior part of the

same same on both sides (Figs. 41B,

Between the the thorax Between and the abdomen, a hard ring test of contains dark (Fig. cells

them to the body wall. White rings are also present close to to The the white are siphons. present rings also close White them to the wall. body

pigment pigment completely disappears after fixation. The abdomens remain yellowish.

41A). 41A). The thoracic musculature forms a pattern complex not is which the exactly

which which correspond to large transverse sinuses and to the trabeculae which link

pigmented and dearly visible. The thorax is also circled with white transverse lines transverse white pigmented with The and dearly thorax circled also is visible.

and and lines.

There There is one spot between the lobes. The neural ganglion and the endostyle are

soft, soft, and transparent. perfectly In the life, thoraces are with white colorless spots 224 224

Thoraces lie parallel to the colony colony the to lie parallel Thoraces

bundle

ing ing

work of blood blood of work

to to

it. In In it.

fte rnvre n lniuia fbr n h thorax the on fibers longitudinal and transverse the of

The musculature is is very characteri musculature The

Fig

s s

are only made of the fibers posterior to the the to posterior fibers the of made only are

life

ure ure

nte abdomen another

both both

, ,

42. 42.

the test was colorless colorless was test the

sinuses appear at at appear sinuses

Diazona Diazona

es o es

f f

n ab an

labyrinthea labyrinthea

. .

Monniot Monniot

Diazona tenera tenera Diazona

dom

en. en.

the the

n. n.

& &

and and

surface while while surface

sp.

Mon

surface. surface.

: :

A, A,

translucent

n. s n.

niot: niot:

tic tic

zooid

(Fig. (Fig.

New Ascidians Ascidians New

: :

D, D,

; ;

B, abdomen abdomen B,

zooid; zooid;

42A) 42A)

the the

; ;

fe fxto a oau net opaque an fixation after

abdomens abdomens

E, E,

neural neural

with with

thor

of of

a regular regular a

acic acic

. .

he same zooid; zooid; he same

ganglion. They They ganglion.

h to abdom two The

muscles; muscles;

are perpendicular perpendicular are

quare quare

, ,

C, C,

, ,

cross

stop stop

ina

225 225 l l

tenera) tenera)

= =

new new species

Micronesica Micronesica 29(2), 1996

Figs. Figs. 42D-G, Pl. 7G

Diazona Diazona tenera

About About 20 tentacles are located anteriorly inside the oral siphon; they are

Description: Description: The colony, hanging from a rock, forms a delicate (

The The esophago-rectal waist is long. The elongate stomach has thin regular

The The wide apart siphons have no defined lobes. About 20 tentacles in two

1°23.S0'N; 124° 44.60'E, 44.60'E, m. 12 124° 1°23.S0'N;

sharp sharp rapheal languets are curved to the right side, increasing toward in size the

is is a rather large unperforated area anterior to the first row of stigmata. The 27

opening. opening. The globular neural ganglion is posterior to the dorsal tubercle. There

prepharyngeal prepharyngeal groove has only one rim. The dorsal tubercle has a large round

short, short, curved and stout. Their size is variable and not regularly arranged. The

abdominal abdominal bundles terminate in the proximal part of the vascular process (Fig.

tracted tracted anteriorly. The arrangement of (Fig. muscles 42E) is characteristic. The

other, other, the oral one with 8 lobes, the one with cloaca} 6. All thoraces were con

dostyle dostyle and the dorsal lamina. After fixation, the pigment disappears except for 42F). 42F).

both both siphons, one above the neural ganglion, and two white along lines, the en

trude trude from common the White test. patches are on seen and the of between lobes

a a patch yellowish at the base of the endostyle. Both siphons are close to each

gelatinous gelatinous with mass transparent a colorless test (Pl. 7G). Only the thoraces pro

Holotype: Holotype: MNHN: Pl Dia 22-Indonesia, north Sulawesi, west of Manado,

loop. loop. The voluminous heart at is the bottom of the abdomen.

with with oocytes few is posterior to the stomach. The pad of globules is poorly de

siphon. siphon. The not gonads developed were in the colony fully The (Fig. 42C). ovary

grooves grooves (Figs. 42B, C). The anus with a plain edge opens close to the cloacal

the meshes contain contain the meshes short stigmata. veloped veloped also. The were empty testis follicles over lying the pole of the intestinal

stigmata stigmata has T-shaped papillae. The are transverse protruding vessels internally;

and and do not have a dorsal branch. On each of the side dorsal the first row line, of

vessels on on vessels each side. The papillae which support the are vessels short and stout

esophagus esophagus (Fig. 42A). There are about 25 rows of stigmata and 30 longitudinal

neural neural ganglion to the esophagous aperture, but the endostyle stops far from the

the the right The side. branchial sac is distorted; the rapheal part extends from the

transverse vessels; they are are short they anteriorly and trans posteriorly. first vessels; transverse longer The

protruding protruding between the siphons. About 20 dorsal languets are inserted on the

tubercle tubercle has a circular opening, it is located far anterior to the neural ganglion,

groove groove has only one undulated margin. It makes a deep dorsal V. The dorsal

of the siphon. A high undulated crest connects all the tentacles. The prepharyngeal prepharyngeal The tentacles. the all undulated connects high crest A siphon. the of verse sinuses do do not sinuses verse reach the dorsal There line. a is dorsal unperforated area on

orders of size were counted; orders counted; were of the size largest arise more posteriorly but may come out

abruptly abruptly behind the stomach, and so, do not extend in the posterior vascular

process process (Fig. 42B). 226 226

ganglion is slightly elongated and the globular neural gland is located externally externally is located gland neural globular the and elongated slightly is ganglion

tubercle is protruding, um-shaped with a circular opening (Fig. 43B). The neural neural 43B). The (Fig. opening circular a with um-shaped is protruding, tubercle

missing in some parts of the ring. A beginning of a fifth order is indicated by by indicated is order fifth a of beginning A ring. the of parts some in missing

ribbon. A triangular sheet joins it to the dorsal lamina (Fig. 43B). The dorsal dorsal The 43B). (Fig. lamina dorsal the to it joins sheet triangular A ribbon.

dorsal indentation and is made of a high anterior blade and a wide flat posterior posterior wide flat a and blade anterior high a of is made and indentation dorsal

small buds on the tentacular crest. The prepharyngeal groove does not have a a have not does groove prepharyngeal The crest. tentacular the on buds small

crest. The tentacles of fourth order have a variable development and may be be may and development variable a have order fourth of tentacles The crest.

in both siphons. About 50 tentacles are regularly intercalated in 4 orders, on a a on orders, 4 in intercalated regularly are tentacles 50 About siphons. both in

cles (Fig. 43A) consist of two sets of parallel fibers, slightly branched, posterior posterior branched, slightly fibers, parallel of sets two of consist 43A) (Fig. cles

both sides of the body; posteriorly, the muscles do not extend beyond the level level of the beyond extend not muscles do the posteriorly, body; sides the of both

the 12th row of stigmata (Fig. 43A). A short but wide velum is located anteriorly anteriorly is wide velum located but short 43A). A (Fig. stigmata of 12th row the

imn pt.Termsuaue spol dvlpd h aea toai mus thoracic lateral The developed. is poorly musculature Their spots. pigment

to the cloaca} siphon. Anteriorly, down to the anus, the fibers are continuous on on continuous are fibers the anus, the to down Anteriorly, cloaca} the siphon. to

a short stalk. The test carries only some polychete tubes. In formalin, the zooids zooids the formalin, In tubes. polychete some only carries test The stalk. short a

minal and close to each other. The zooids are linked to a stolonical network by by network stolonical a to linked are zooids The other. each to close and minal

are transparent, colorless and without patches of blood cells. cells. blood of patches without and colorless transparent, are

lagoon, reef south of Dublon Island, 7°20.92'N; 151°52.?0'E, 13 13 m. 151°52.?0'E, 7°20.92'N; Island, Dublon of south reef lagoon,

Ecteinascidia diaphanis diaphanis Ecteinascidia

Material Examined: Examined: Material

and synonymy. synonymy. and

polyhedral masses; according to the zooids, they extend from the middle of the the of middle the from extend they zooids, the to according masses; polyhedral

the gut loop. The "pseudo-ovarian" globules are not well developed; they form form they developed; well not are globules "pseudo-ovarian" The loop. gut the

bunch of testes encircle the pyloric end of the stomach and the posterior end of of end posterior the and stomach the of end pyloric the encircle testes of bunch

ovary to the esophago-rectal waist. waist. esophago-rectal the to ovary

facing the eighth row of stigmata. stigmata. of row eighth the facing

mata. The rows of stigmata do not show any division but a few small stigmata stigmata small few a but division any show not do stigmata of rows The mata.

regular plications. The intestine is not subdivided. The simple anal opening is is opening anal simple The subdivided. not is intestine The plications. regular

The vessels are close to each other and limit long meshes containing 1 to 3 stig 3 1 to containing meshes long limit and other each to close vessels are The

oviduct. The gonads (Figs. 42F, G) are diffuse, the ovary on the right side and a a and side right the on ovary the diffuse, are G) 42F, (Figs. gonads The oviduct.

are intercalated in a row. row. a in intercalated are

esl nec sd ftedra ln,ad h ls o h notl hv T have endostyle the to close 2 the and line, dorsal the of side each vessels on

trusion directed toward the dorsal line but which does not exceed the sinus level. level. sinus exceed the not does which but line dorsal the toward directed trusion

languets. The branchial sac has 28 rows of stigmata on the left and 29 to 30 on on 30 to 29 and left the on stigmata of rows 28 has sac branchial The languets.

shaped papillae more or less joined. These papillae, single or joined, have a pro a have single or joined, papillae, These less or joined. more papillae shaped

the right side. There are 20 longitudinal vessels on each side. The 3 longitudinal longitudinal 3 The side. each vessels on longitudinal 20 are There side. right the

area with raised crests which correspond to the transverse vessels vessels transverse the to rapheal their and crests raised with area which correspond

bottom of the branchial sac. On the right of the dorsal line line is wide a unperforated dorsal the of right the sac. On branchial the of bottom

The siphons are retracted into the tunic and have 8 short lobes, without without lobes, short 8 have and tunic the into retracted are siphons The

h ois . mi eghaegopd nacutr hi sposae ter are siphons Their cluster. a in grouped are length in 1.6 cm zooids The

The esophago-rectal waist is long (Fig. 42D). The long stomach has thin thin has stomach long The 42D). (Fig. long is waist esophago-rectal The

MNHN: MNHN:

Ecteinascidia diaphanis diaphanis Ecteinascidia

Sluiter, 1885: 168, Indonesia; Kott, 1985: 90, Australia, Australia, 1985: 90, Kott, 168, Indonesia; 1885: Sluiter,

Monniot Monniot

Family Perophoridae Perophoridae Family

P2 Ect 64--Federated States of Micronesia, Chuuk Chuuk Micronesia, of States 64--Federated Ect P2

& &

It It

Monniot: Monniot:

Fig. 43 43 Fig.

is accompanied by the sperm duct and the the and duct sperm the by accompanied is

New New

Sluiter, 1885 1885 Sluiter,

Ascidians Ascidians 227 227

mm mm

l l

by by Kott (1985), with very small

1996 1996

, ,

Sluiter, Sluiter, A, 1885: zooid; B, neural area.

E. E. diaphanis

Micronesica 29(2) Micronesica

Ecteinascidia Ecteinascidia diaphanis

Figure Figure 43.

Remarks: Remarks: Athough the gonads are not well developed, fully specimens these

The The oval stomach has clear spiral ridges. The post-stomach differen is well

stigmata stigmata in a to mesh, instead 6 8 2 to of Kott's 3. the anus shows 35, opening fig.

differences: differences: 20 rows of stigmata instead of 16 to 19 and a largest number of

runs runs along the dorsal lamina and ends in a small papilla posterior to the anus.

closer to closer the siphon cloaca! than in our but with specimens, regard the exact level

forms forms an angle with the transverse (Fig. vessels 43A). The gonads are not fully

correspond correspond to the description of

above above the stomach. The common sperm duct begins anteriorly to the ovary; it

tiated. tiated. The intestine begins by a widening. The anus opens with a plain margin

developed. developed. The are testis follicles arranged in a crown around the ovary located

The The oviduct in the lies right of side the body.

to to 8 regular stigmata.

at at the of level the seventh row of stigmata. The rectum is not very curved and

dorsal dorsal lamina and on one each series of side the endostyle. There are neofor few

plete longitudinal longitudinal plete 2 longitudinal vessels, flat of papillae series on each of side the

mations mations of stigmata and none of transverse The vessels. meshes are long with 6

linked linked at mid length a by thin membrane. There are no papillae at the of the level

and and on slightly the of left the ganglion. The dorsal lamina has numerous languets

posterior posterior does not reach the dorsal line. For one counted we specimen, com 18

first transverse vessel. The The branchial transverse first vessel. made sac is of rows 20 of stigmata; the most 228 228 Monniot & Monniot: New Ascidians 229 to the branchial sac is not indicated, and a larger interval separates the muscu lature of the cloacal siphon and the transverse thoracic muscles . Kott (1985) synonymized E. hataii Tokioka, 1950 from Palau with E. dia phanis, a decision with which we agree. Our samples, except for a larger size (16 mm instead of IO mm), and more stigmata rows, correspond well to Tokioka's description. We are unable to consider the largest size of the siphons of E. hataii, as this author always represents the ascidians with the condition that they may have when relaxed. Kott (1992b) synonymized the New Caledonian species E. koumai C. Mon niot,1987a and E. ndoue C. Monniot, 1991a with E. diaphanis. We cannot accept this synonymy as unlike the present species, E. koumaci has transverse muscles anterior to the cloacal siphon and a sperm duct opening at the anus level. This species is close, though, to E. diaphanis for the remaining structure. In E. ndoue the shape of the gut, with a pronounced secondary loop, is very different from E. diaphanis.

Ecteinascidia styeloides (Traustedt, 1882) Fig. 44 Phallusia styeloides Traustedt, 1882: 277, pl. 4, fig. 5, pl. 5, fig. 16, West Indies. Ascidia styeloides: Herdman, 1891a: 593. Ascidiella styeloides: Van Name, 1930: 470, fig. 42, West Indies. Ecteinascidia styeloides: C. Monniot, 1983a: 59, fig. 3, Puerto Rico and Guade loupe; Monniot & Monniot, 1987b: 128, Mediterranean Sea (Corsica). EcteinascidiagarstangiSluiter, 1897: IO, pl.I, fig. 1, pl. 3, fig. 1-4, Mozambique; Sluiter, 1900: 5, Hawaii; Sluiter, 1904: 10, Indonesia (in part); Vasseur, 1969: 922, fig. 14--16, Madagascar; Renganathan, 1984: 54, fig. I, South India; Nishikawa, 1986: 35, fig. 2., Micronesia. Ecteinascidia diligens Sluiter, 1900: 3, pl. 2, fig. 1-3, Hawaii. Ecteinascidia tokaraensis Tokioka, 1954b: 255, pl. 30, fig. 1-5, South of Japan. Ecteinascidia conklini: Goodbody, 1984: 33, West Indies. Ecteinascidia imperfecta: Nishikawa, 1984: 155 (in part), Micronesia. ? Ecteinascida rubricol/is: Kott, 1964: 146, Queensland. Material Examined: MNHN: P2 Ect 63-Palau, Koror State, Rock Islands, 7°20.28'N; 134°26.71'E, 30 m. About 10 zooids were found united by stolons as epibionts on a group of Polycarpa contecta. The largest zooids are 12 mm long. The tunic is thin and transparent, and has some hydrozoans and algae. The siphons are short, close together in erect specimens but the cloaca} siphon may lengthen to 4 mm in a zooid inserted between 2 Polycarpa. The tunic is covered with small hairy protru sions above all the surface in contact with Polycarpa. The siphons are edged in about IO triangular lobes. Inside the oral siphon are 4 cup-shaped protrusions. The lateral musculature (Fig. 44) is made of transverse fibers extending from the level of the second row of stigmata to the top of the intestinal loop (second type after the terminology of Nishikawa 1986). The fibers issued from both sides are 230 Micronesica 29(2), 1996

Figure 44. Ecteinascidiastyeloides (Traustedt , 1882): A, muscles; B, neural area; C, D, external and internal sides of the gut and gonads. connected along the dorsal axis of the body. The transverse fibers are linked together by numerous branches. The lateral muscles are independent from the siphonal musculature. About 35 long and thin oral tentacles in 3 orders are placed on concentric close circles. The prepharyngeal groove has inequal rims; the most anterior makes a protruding rod, the posterior one is a flat and larger ring. There is no dorsal indentation. The dorsal tubercle, um-shaped, inserted into the body wall, opens by a simple hole. AV-shaped structure (Fig. 44B) is anterior to 15 protruding rapheal languets united at their base by a high sheet. The first languet corresponds to the second transverse branchial sinus, the others to the successive transverse sinuses. The branchial sac has 19 rows of stigmata on each side. There are 19 longitudinal vessels on the right side and 20 on the left. Only the first sinus on the right and those against the endostyle are incomplete. There are 2 to 3

without epibionts, of an opaque blue (Pl. 7H), at the origin of the species species name. the of origin the at 7H), (Pl. blue opaque an of epibionts, without

and fixed on the rock by by rock fixed the on and

cording to the rules of nomenclature. nomenclature. of rules the to cording

Holotype: MNHN: MNHN: Holotype:

Queensland. Queensland.

ambique, Madagascar, southern India, Hawai'i, Micronesia, southern Japan, Japan, southern Micronesia, Hawai'i, India, southern Madagascar, ambique,

of of

/oides /oides

duct extending farther than the anus. The only difference observed is, is, difference in only observed The anus. the than farther extending duct

height of the membrane uniting the dorsal languets, the gut shape and the sperm sperm the and shape gut the languets, dorsal the uniting membrane the of height

to justify separate species. The name name species. The separate justify to

characters are identical: the musculature, the prepharyngeal groove, the large large the groove, prepharyngeal the musculature, the identical: are characters

collected in the mangroves of Puerto Rico and Guadeloupe and Mediterranean Mediterranean and Guadeloupe and Rico Puerto of mangroves the in collected 96 oe a e clr f h Slmn sad specimens Islands Solomon the of color red a ( 1986) noted

tation darker than that of the body. This color disappears in fixatives. in fixatives. disappears Nishikawa color This body. the of that than darker tation

have, in in have,

cavity. The colony only contained young non-differentiated larvae. larvae. non-differentiated young contained only colony The cavity.

samples of the Palau collection. The West Atlantic and Mediterranean specimens specimens Mediterranean and West collection. Atlantic The Palau samples the of

samples from Corsica. Only insignificant differences were found in the Pacific Pacific the in found were differences insignificant Only Corsica. from samples

The globular ovary is placed on the external side of the testis, in the center of the the of center the testis, in side of the external the on is placed ovary globular The beyond the rectum (Fig. 44D) at a length equal to that of one stigmata row. The The row. stigmata one of that to equal length a at 44D) (Fig. rectum the beyond onymy of of onymy

externally. The sperm duct follows the rectum and ends in a papilla extending extending papilla a in ends and rectum the follows duct sperm The externally.

are incubated inside the oviduct, and the most part in the right side side cloacal of the right the in part most the and oviduct, inside the incubated are

oviduct is prolongated toward the bottom of the right side. Some of the larvae larvae the of Some side. right the of bottom the toward prolongated is oviduct

level of the 16th row of stigmata. The stomach is globular, without ridges. The The ridges. without globular, is stomach The stigmata. of row 16th the level of

opens with a smooth-edged anus between the 10th and 11th rows of stigmata. stigmata. of rows 11th and 10th the between anus smooth-edged a with opens

mata. The growth seems to be achieved in those individuals. individuals. those achieved in be seems to growth The mata. the 16th stigmata row. The rectum is long, attached to the dorsal lamina, and and lamina, dorsal the to attached long, is rectum The row. stigmata 16th the specimens (Fig. 44D) the testis lobes cover the whole intestinal loop and protrude protrude specimens and testis lobes the whole loop 44D) (Fig. cover the intestinal

and the the even intestine, incubating In in developed and individuals gonads. with poorly

intestine has a pronounced constriction. The top of the primary curve reaches the the curve reaches primary of the top The constriction. pronounced a has intestine gut loop (Fig. 44C) (Fig. loop gut

tgaa nams.Teeaefwitrutd iue n fwitraae stig few intercalated and sinuses few interrupted are There mesh. a in stigmata

13th stigmata row, whereas the bottom of the secondary curve is at the level level of the at is curve secondary the of bottom the whereas row, stigmata 13th

23.74'N; 124°32.41'E, 124°32.41'E, 30 m. 23.74'N;

E. garstangi garstangi E.

Description: One sample only was examined, it was was 4 was it only examined, sample One Description:

Geographic Distribution: West Indies, Mediterranean Sea (Corsica), Moz (Corsica), Sea Mediterranean Indies, West Distribution: Geographic

Remarks: Nishikawa (1986) in his revision of of revision (1986) his in Nishikawa Remarks:

The gut forms a double curve (Figs. 44A, C). The esophagus opens at the the at opens esophagus The C). 44A, (Figs. curve double a forms gut The

a more regular distribution of the testis follicles than in described samples samples described in follicles testis than the of distribution regular more a

life, life,

E. garstangi garstangi E.

a characteristic aspect with long siphons with a red-brown pigmen red-brown a with siphons long with aspect characteristic a

from the Indo-Pacific region. This minor distinction does not seem seem does not Indo-Pacific distinction the region. from This minor

PS PS

. .

The testis lobes extend on the internal side of the stomach stomach the of side internal the on extend lobes testis The

and and

Ase A 262 A Ase

/3 /3

Monniot Monniot

Ascidia azurea azurea Ascidia

E. styeloides. styeloides. E.

of the left posterior side. side. test is perfectly The left smooth, the of posterior

Family Ascidiidae Ascidiidae Family

Fig. 45, Pl. 7H 7H Pl. 45, Fig.

& &

Monniot: New Ascidians Ascidians New Monniot:

Indonesia, north Sulawesi, Sulawesi, Manado, of west north Indonesia,

E. styeloides styeloides E.

We reexamined samples of of samples reexamined We

new species species new

E. garstangi garstangi E.

a pirt ad sue ac used is and priority has

x x

. .

Most anatomical anatomical Most

1 1

suggests the syn suggests the

x x

1.3 1.3 erect cm,

E. E.

styeloides styeloides

E. stye E. 231 231

Micronesica Micronesica 29(2), 1996

n. n. sp.: A, side; left muscles B, on the right side; C, internal

Ascidia Ascidia azurea

. .

side side of the gut; D, neural area.

Figure Figure 45

formalin formalin it turned whitish. No or vessels blood ampullae are seen in the firm

the the dorsal tubercle. The dorsal lamina rapidly increases in height anteriorly. It

lobes lobes but is irregularly swollen. The musculature is strong (Figs. 45A, B) and

aca} siphon siphon may lengthen aca} and extend posteriorly contracted when alive; it has no

with with papillae. The dorsal tubercle is U-shaped. The neural ganglion is far from

the the right There side. no is dorsal indentation. The prepharyngeal area covered is

is is covered with digitiform papillae. The posterior rim is high and smooth (Fig.

separated. separated. The prepharyngeal groove has a particular structure. The anterior rim

arranged arranged in 4 orders of size. They are not very long and their bases are well

forms an irregular network in the middle of each side. Forty Forty an forms each side. irregular of are tentacles network regularly middle the in

aperture aperture lined is by about undulations, 10 without basal pigment spots. The clo

and and thick test mm (1 thick on the and left mm 2 thick on the right The side). oral

In In

45D). 45D). The prepharyngeal groove closer to is the tentacles on the than left side on 232 232 pheres

nrneo h ih ie ntebacilsc ecutd6 ogtdnl vessels 62 sac,we longitudinal branchialcountedside, the in right the on entrance

cesses. The branchial meshes are longer than wide. Minute plications are accen are plications Minute wide. than longer are meshes branchial cesses. The

forms 2 blades from its origin to the level of the neural ganglion. Posteriorly, therelevel 2 the forms its blades originto from Posteriorly, ganglion. of neural the sol n ihset t egtarpl dmnse eoeteeshgs open eosphagus the before diminishes abruptly height sheet; its high one only is ing, and there, it is replaced by papillae, united by a low membrane. In the centre the In low membrane. a by united papillae,is by replaced it there, and ing,

lamina ending in papillae. At the esophagus entrance, on the right side, the trans side, the right the on entrance, esophagus the At in papillae. ending lamina es esl n naln fppla, hre pseiry A telvlo h gut the level of the At posteriorly. shorter papillae, of line a vessels in verse end

developed on the external side the external developed on (Figs. 45A, testisC). The follicles over scattered are pro lateral without flat, high, are papillae branchial side. The left the on 45 and

utd ntemda pr o h rnha sc hyfr puhsmd o two of made pouches form they sac; branchial the of part median the in tuated successive meshes with a depth the double height of a papilla. There are no par no are There papilla. a of height double the depth successive a withmeshes astigmatic vessels and no intermediate papillae. All the external side of the bran the ofside external the All papillae. intermediate no vessels astigmatic and chial tissue is covered with small papillae.small with istissue covered chial me/anostoma dorsal the of side left the vessels onto transverse extend the sac, branchial the of

margin. The ovary is a mass of tubules in the primary intestinal loop and is more and loop intestinal primary the in tubules of mass is a ovary The margin.

h woegt op Te oout, ulo octs n semtzod, open spermatozooids, and oocytes of full gonoducts, The loop. gut whole the with tubules overlapping the limbs of the gut loop and becoming visible from the becomingvisible from and loop gut the of limbs the overlapping tubules with a in gathered vesicles not small, are level. accumulation The anus the at together external side. external defined outline. All side the internal of the body wall is covered with small papillae. rmr itsia loop. intestinal primary

n tikts, h aemsuaue atr, erl agin t mid-distance at ganglion neural a pattern, musculature same the test, thick and has few grooves stomach visibleThe siphon. glandularvoluminous the on external from the base of the siphons, the same shape of the gut with a cylindrical a intestinewith gut of the shapesame the siphons, the of base the from undulated but plain a having anus an with ends intestine cylindrical The side. and a prepharyngeal area covered with papillae. However, all these species were these all However, papillae. with covered area prepharyngeal a and observed when living and none had a blue color. Additionally all these species these all Additionally color. blue a had none and living when observed Phallusia arabica Phallusia tract digestive the of side internal the on developed more gonad female a have ymy.

ao l nado, neir ieo h rpayga groove. prepharyngeal the of side anterior yhl o h pseir etsd.I lf h aia ws fa nfr pl blue pale uniform a of was animal the life In side. left posterior the of half by color.

Material Examined: Material

The gut (Fig. 45C) describes a double loop reaching the level of the cloaca! the level of the reaching loop double a describes 45C) (Fig. gut The

eak: hsseishs hrces iia o or other four to similar characters species has This Remarks:

The studied sample measures 51 x 55 x 18 mm, it was attached on the rock the on attached was it 18 mm, 55 x 51 x measures sample studied The

Sluiter,1895,

65'; 124°44.48'E, 20 m. 26.57'N;

a rtuig ihn ih1 oe ihwiesos etrmre on 12 marked better with lobes spots, white siphons with protruding had

Sluiter, 1885. All have in common the same body shape, a smooth a shape, body same the common 1885.in have All Sluiter,

is not the case in the is not

ain,11:14 ot 18:6,fg2ad ll, o synon not 1985: fig.24a-d, 61, pl.le, 1816: 164; Savigny, Kott,

A. gemmata A.

MNHN:

hlui arabica Phallusia

. melanostoma A.

Monniot

5Pa7-noei, ot Sulawesi, west ofMa north 73-Indonesia, P5 Pha

Sluiter, 1895, Sluiter,

Fig. 46, PL 8A 46, Fig.

. azurea A.

Monniot: New Ascidians New Monniot:

Savigny, 1816 Savigny,

whose ovary is the strictly inenclosed whose ovary

. azurea A.

A. g/abra A.

osse aile n the on papillae possesses

atee, 1921, and Hartmeyer,

sii: . em A. Ascidia:

233

A. contains

of the distance between the ganglion distance the the betweenof

A, left side; B, internal side of the gut.

Micronesica 29(2), 1996

Savigny, 1816

has no striking characters. About 60 short and well

orders of size, are inserted above a muscle. The pre

Pha/lusia arabica Pha/lusia

Figure 46.

After fixation, the siphons are contracted and not so protruding. The firm

234 and thick test is 1 to 3 mm in thickness and has a granular surface.

of the of gut loop. The depressed test interiorlyslightly is against the tract. digestive

the left side (Fig. 46A). the cloacal siphon (Pl. 8A). All colors disappear in formalin and the animal is pharyngeal groove, with two laminae, close is to the closer tentacle on ring the ( yellowish in a yellowish translucent tunic. left side); the area between them is covered with papillae. There is no dorsal indentation. A dorsal tubercle was not found in the studied The specimen. neural numerous blood star-shaped diverticula, whichare colorless. Internally a net lies at the base is, ganglionsiphon. of cloacalthe neuraltheopenings of The accessory work of large blood sinuses originating from leavinga the body vessel at the top gland are placed in a cluster, near the neural ganglion, they are more isolated and less numerous anteriorly, less and disappear at and the prepharyngeal groove. The dorsal lamina doubled is anteriorly. Posterior dentate. The musculature is strong on the right side and on the anterior part of to the neural ganglion, it is a sheet ofsingle medium height. On the left side, the

spaced tentacles, in 4 or

The body wall is thin. The siphonal lobes are not marked well and their edge is Monniot & Monniot: New Ascidians 235 transverse sinuses extend onto the dorsal lamina in a small papilla. On the right side, the dorsal lamina is covered by digitiform papillae. It curves around the esophageal opening and then diminishes in height. On the right side, close to the esophageal opening, the transverse sinuses form a low plain lamina; posteriorly they only form decreasing papillae. The branchial tissue is neatly pleated. It has about 70 longitudinal sinuses on the right side and 60 on the left. The main pa pillae are not high and have small lateral extensions. Many intermediate papillae like small buttons become visible only after staining. The square meshes contain at least 10 stigmata. Parastigmatic vessels only exist when a row of stigmata is dividing. The gut forms a closed loop (Figs. 46A, B). The rectum is dilated. The anus is attached to the body wall, it has low lobes. The ovary is in the primary gut loop, spread over the internal side (Fig. 46B). The testes are in small dense masses over the digestive tract. The gonoducts open together under the anus. The accu mulation vesicles are small and contain brown grains. Geographic Distribution: The species is recorded from the Red Sea, Mada gascar, Sri Lanka, Philippines, and north eastern Australia.

Phallusiajulinea Sluiter, 1919 Pl. 8B Phallusiajulinea Sluiter, 1919: 7; Kott, 1985: 65, fig.25, pl.If, and synonymy. Materiel Examined: MNHN: PS Pha 74--Papua New Guinea, North coast, Na gada Harbor, 5°09.35'N; 145°48.00'E, l m. This species is very distinctive (Pl. 8B) and recorded from the north coast of Australia, Indonesia, Palau Islands, and New Caledonia.

Family Plurellidae Plurella kottae new species Fig. 47, Pl. 8E Holotype: MNHN: P7 Plu I-Philippines, Cebu, Mactan Island, Buyong, 10°17.19'N 124°00.15'E, 27 m. Description: This species forms masses of aggregated zooids 3 cm thick (Pl. 8E). The siphons emerge from this mass without peculiar disposition. The siphons are almost equal, some of them a little shorter or lower, are the cloacal apertures. The individuals are included for 2/3 in a common mass. It is not possible to isolate the individual tunics. The appearance of this species is similar to that of the ag gregated Polycarpa tokiokai. The tunic is entirely encrusted with sand and has a reddish color in life. The siphons have internal chalk-white strips separated by red or brown lines (Pl. 8E). There are 16 to 20 lobes at each siphon, slightly protruding. In formalin, the colors disappear. The tunic is divided in two layers, the external one contains much sediment, the internal is thin, without inclusions and easy to separate from the external one in the branchial region. It is thicker at the level of the gonads rapheaJ

the

C, ventral of side

above

fold

specimen;

"' 3 3

or a

E, branchial

sides

both

siphon;

accessory openings of the neural gland; G,

internal side or the ovary.

A, B

and

Micronesica 29(2), 1996 Micronesica

n. sp.:

opened oral

n !!!

....

D, ganglion

In the oral siphon, the musculature fades in front of the

neural

specime~

.':;•

Plurella kottae

47.

another groove; detail of the branchial sac;

Figure

236

tentacle ring. On the right side of the body is an area of thin crowded muscles (Fig. 47B). This (Fig. 47B). area does not cover the whole width of the right There side. a is

which are embedded in it. The siphonal musculature is reduced to some circular and longitudinal fibers

considerable assymetry of the two of sides the body. The body wall extremelyis Monniot & Monniot: New Ascidians 237 thin and is difficult to separate from the internal layer of the tunic. At the level of the siphons, the right side may be three times wider than the left side (Fig. 47D). Sixteen large, long and thin tentacles are evenly spaced along a crest, some times alternating with smaller ones. Small papillae cover the area anterior to the tentacle ring. The prepharyngeal groove has two equal low crests. On the left, it makes a fold characteristic of the family, which is linked to the dorsal lamina. Anteriorly, the dorsal lamina is a double membrane. In the bottom of the furrow is an elongate hollow (Fig. 47E) linked with cells which stain more than the dorsal lamina. In its half way down, the dorsal lamina is a single sheet with a plain margin and thickenings on its left side. The dorsal lamina is interrupted at the esophagus opening. Posteriorly it is replaced by two parallel high blades which represent the retropharyngeal groove. The neural ganglion (Figs. 47C, F) is I mm above the base of the cloaca} siphon, inside the internal tunic, but also projecting into the external tunic in an area where the two layers are tightly united. The neural gland is anteriorly prolonged by a thin channel, branched and ended in a small number of um-shaped ciliated openings at the surface of the internal tunic layer (Fig. 47F). No communication was seen between the neural ganglion and the anterior groove of the dorsal lamina. The branchial sac is flat, extending along the whole body, overlying the gut and the gonads. It has a fold on the left side along its whole length, corresponding to the indentation of the prepharyngeal groove. Anteriorly this fold covers the dorsal lamina, but posteriorly it separates from it. On the right side the branchial wall is flat. There are more than 110 longitudinal vessels on the left and less than 50 on the right, without the IO to 25 crowded vessels making the fold. The vessels are thin with protruding papillae. In opposition to all other families in the Phle bobranchia, the first row of stigmata is overlapped by the longitudinal vessels as in the Stolidbranchia (Fig. 47G). There are 1 or 2 stigmata in a branchial mesh, rounded or elongate, according to the branchial area. The gut is not included in the tunic as in Microgastra granosa (Sluiter, 1904). It is only linked to the branchial sac by the esophagus and seems independant from the body wall. It forms an open loop (Fig. 47A). Its shape is distorted by its content and no differentiations can be seen. The anus is slightly bilabiated. The gonads are typical of members of the Plurellidae, embedded in the internal layer of the tunic. The single testis is enclosed in a pouch at halfway between the cloaca} siphon and the posterior end of the body. The single sperm duct branches close to the group of ovaries at the base of the cloaca} siphon. The ovaries are variable in number and are not arranged in lines but grouped in clusters (Fig. 47A). Each ovary associated with a branch of the sperm duct dilated in an ampulla, generally full of sperm (Fig. 47H). This ampulla is present even empty. Remarks: This species is perhaps the same as the species mentioned by Kott (1985) but not named and only partly described. It differs from Plurella e/ongata Kott, 1973 by numerous characters: the location of the cloacal siphon close to the oral siphon, a loop of the gut, and ovaries in clusters and not in lines. Kott (1973, 1985) estimated that the Plurella species were colonial. At this stage the budding mechanism is not known. She only said that the largest zooids were placed in the

longer longer

species, a a species,

are are

they they

Botryllus Botryllus

Bot Bot B 109-Indonesia, north

new new species

MNHN: MNHN: SI

species remains remains species to be demonstrated.

Family Family Styelidae

Fig. Fig. 48, Pl. 8C, D

Micronesica Micronesica 29(2), 1996

27.08'N; 27.08'N; 124°44.59'£, m. 20

Order Order Stolidobranchia

Botryllus Botryllus firmus

Plurella Plurella

The digestive tract (Fig. 48C, D) is posterior and describes a well pronounced pronounced tract posterior digestive D) The (Fig. well 48C, is a and describes

The The oral siphon not is lobed. tentacles orders in 3 regularly alternate. 12 The

Description: Description: In life, the two colonies had a very different aspect. The type

The The colony structure characteristic. is The holotype forms a thick hard pad

ing) with the the with heart ing) somewhat forming posterior a large vesicle to the The caecum.

region; but but region; on against the side, left the intestine, they are short delimited. and well

the the right side the folds are elongate and progressively disappear in the pyloric

The The small, caecum is stalked, to and difficult see may confused be (without stain

double double loop. The stomach has about protruding 12 folds the in anterior part. On

verse stigmata stigmata verse in the posterior part.

second second row of stigmata which does not reach the dorsal lamina, nor oval trans

the the endostyle and dorsal lamina. We did not find, as in many

vessels are are vessels close to each other, separated by only 2 or 3 stigmata, and far from

branchial branchial sac has to 16 18 rows of 18 to 20 long stigmata. The 3 longitudinal

which are are which not regularly alternated are completely embedded in the common test.

angle angle along the endostyle. There are no dermato-branchial trabeculae. The buds,

agus agus opening (Figs. 48B, C, D). The body wall is thick, in two forming sheets an

languets languets only are gold-yellow in the colony SI Bot B The 109. common test is

at at the of level the common openings, as cloaca} they extend up to its rim. These

zooids. zooids. The siphon zooid very large cloaca} is and reaches of the the level esoph

and and above the neural ganglion. These patches of pigment are not on visible living

zooids zooids tum pale ochre with two brown patches at the upper part of the endostyle

hard, hard, full of blood Its vesicles. surface is smooth and shining. In formalin, the

Sulawesi, west west of Manado, Sulawesi, l

perpendicularly perpendicularly to the surface and regularly spaced on each side of the common

(7 (7 mm) giving the species name. The long zooids (up to 4 mm) are arranged

The The pattern corresponds to the common cloacal channels. After fixation, the two

nian. The other colony was black with a gold yellow meandriform meandriform nian. The 8D). other (Pl. yellow black with design gold was colony a

colony colony was uniformly gold and yellow opaque (Pl. 8C) encrusting a dead gorgo They They are short when the zooids are facing in two parallel rows;

48A). The The 48A). roof of the channels suppported is by the atrial languets of each zooid.

cavity is enlarged enlarged and protruding a is opens by cavity opening (Fig. round common cloaca}

cloacal cloacal channels of the colony. In some places of the meandriform network, the

Holotype: Holotype: MNHN: SI Bot B 108-Papua New Guinea, Port Moresby, Basilisk

colonies colonies became a little translucent and pale brown.

passage, passage, m. 28 9°32.32'S; 147°08.04'E,

colonial colonial structure of the

part part in the budding. We have not observed any indication of budding, and the

center center of the colony and that the vascular ampullae of the tunic should play a 238 238

The buds of the the of buds The

to to

of the third last last third the of

the base at the the at base the

intestine intestine

testes at the the at testes

the the

stomach (Fig. (Fig. stomach

Figure 48 Figure

describes a closed loop, its top at the level of second last last level second of the at top its loop, closed a describes

order bud. bud. order

level level

, ,

. .

stomach stomach

, ,

first first

left left

Botr

row row

of the intestinal loop. One regularly is on the right right the on is regularly One loop. intestinal the of

side of the gut in in gut the of side

generation, whose siphons are not yet yet open, not whose are siphons generation,

48B) and sometimes another is on the left, left, the on is another sometimes and 48B)

llus llus

of stigmata. The feeding zooids only have have only zooids feeding The stigmata. of

level. The anus is wide, with a plain rim, located located rim, is plain a wide, level. with anus The

firmus firmus

Monniot Monniot

n. n.

p.: A, A, p.:

& &

two two

Monniot

0 0

l{l l{l

e e

part part

ooids

of the colony the of

: :

; ;

New New

E, E,

F, F,

Ascidians Ascidians

right and and right

; ;

B, B,

right side of a zooid a of side right

dorsal dorsal

......

e e

!i !i

sides of a first first a of sides

have have

in in

very very

stigmata row, row, stigmata

the gut loop. loop. gut the

t t

side, side,

or or

posterior posterior

in in

; ;

2 2

front front

close close

well- 239 239

by by

from from

Herdman, Herdman,

sensu sensu Kott,

Botryllus Botryllus auran

species, species, colonies

Botrylloides Botrylloides vio/aceus

Botryl/oides Botryl/oides vio/aceus

Botryllus Botryllus

B. B. perspicuum rubicundum)

Botrylloides Botrylloides perspicuum

and and

n. n. sp. looks like some Australian spec

Ritter Ritter & Forsyth, 1917 were seen only

(Botryl/oides (Botryl/oides magnicaecum

n. n. sp., but the zooids are smaller, have a

Tokioka, Tokioka, from 1967 Palau has an external

(1981a, (1981a, b) and Saito & Watanabe (1985) have

Micronesica Micronesica 29(2), 1996

(B. (B. perspicuum

Botryllus. Botryllus.

et et al.

in in the colony and zooid structure. The specimens de

Botryllusfirmus Botryllusfirmus

(Herdman (Herdman 1891)

Botryllus Botryllus firmus

Saito Saito

n. n. sp. differs from the 5 Japanese species included in

Botryllus Botryllus tuberatus

. .

considered considered by Tokioka (1953) as the commonest Japanese

B. firmus firmus B.

n. n. sp. is characterized by the oocytes located in the buds of the

B. B. aurantius.

is is incomplete

By its its By external aspect,

Botryllus Botryllus anceps

Botrylloides Botrylloides marginatus violaceus

Botryllus Botryllus firmus

Remarks: Remarks: The knowledge about the Indo-Pacific species of the genus

(Oka, (Oka, C. (see 1927b) Monniot In 1988: 169). many

1886. Her Her 1886. description, however, includes at least 5 specimens showing important

1985) also also 1985) has a large cloaca} aperture, an oval stomach with a large caecum, the

the the Philippines have a secondary intestinal loop little pronounced. No indications

scribed scribed by Herdman (1886)

are are given concerning the cloaca} aperture and gonads.

imens imens that Kott, grouped 1985 under the name

differences differences from

feeding feeding zooids.

intestine intestine without a second loop, an ovary anterior to the testis and occurring in

lamina, lamina, rudimentary stigmata in the posterior part the of branchial sac. The gonad

uncommon uncommon condition for a

posterior posterior gonads and figured a young gonad which seems hermaphrodite, a very

of of the type specimen is not known, but in another sample Tokioka noted very

large large cloacal aperture and a second stigmata row which does not reach the dorsal

aspect aspect very similar to

belong belong to

at at the middle of the branchial sac. Although the gonads are unknown, it cannot

Kott, Kott, have 1985 the intestine without a neat secondary loop, and the anus opening

trylloides trylloides vio/aceus,

second second stigmata row not reaching the dorsal lamina. According to the species, the

the the bud of first order, a postero-dorsal ovary, an antero-ventral testis and the

curved, curved, but always present. The specimens identified as

species. species. All these species have in common the appearance of the female gonad in

cloacal cloacal aperture may be large or reduced and the secondary gut loop more or less

twice). twice). The species identification requires the presence of gonads: for example

occurs occurs (the testes of

second second generation.

Botryllus Botryllus firmus

tius tius

tryllus tryllus (Oka, (Oka, This 1927b). species name is not valid and was changed to

with with ovaries are rare but almost all samples have testes. The reverse sometimes

development, development, could help to isolate 5 Japanese species from

demonstrated demonstrated that the biological cycles, and particularly the period of oocyte

the the beginning of the branchial differentiation. The bud, left smaller, does not seem

to to contain oocytes. The studied colonies did not contain ripe oocytes nor tadpoles.

48E, 48E, F). The latter, on the right side, have on each side a group of oocytes before

developed developed testes, protruding exteriorly, and 2 buds of second generation (Figs. 240 240 Monniot & Monniot: New Ascidians 241

Amphicarpa agnata (Kott, 1985) Fig. 49A Stolonica agnata Kott, 1985: 232, fig. I 10, Queensland, West Australia. Amphicarpa agnata: C. Monniot, 1988: 182, fig. 8, New Caledonia. Material Examined: MNHN: SI Amp 14-Federated States of Micronesia, Chuuk Atoll lagoon, patch reef north of Fefan Island, 30 m. The samples from Micronesia were attached on flakes of calcareous Halimeda sediment or on shell debris at 30 m depth. The colony studied consists in numerous zooids united by stolons. The tunic is slightly covered with sand. The color in formalin is light brown. Some zooids shelter in their tunic bivalve molluscs whose size sometimes reaches that of the ascidian host (7 mm). The presence of the bivalve does not affect the ascidian development. Some small differences can be noticed with previous descriptions. In the Chuuk specimens, the branchial sac may have 17 rows of stigmata (13 in Aus tralia, 12 in New Caledonia). The second branchial fold on the left side is replaced by a single sinus higher than the others. Kott (1985) found l to 3 sinuses on this fold. In our material, in the anterior part of the branchial sac, on the sides of the higher sinus the two lateral sinuses are closer, but do not make a fold. The shape of the stomach is slightly different. Here (Fig. 49A) the stomach is larger in its pyloric region and has 13 plications. Kott describes "a cylindrical rather a pyri form stomach", with 12 folds. In New Caledonia the stomach has only 6 folds. The pyloric caecum is more developed in the Chuuk colony. The structure, num ber and disposition of the gonads are everywhere identical. Geographic Distribution: Queensland, New Caledonia.

Eusynstyela latericius s. str. (Sluiter, 1904) Fig. 49B-D, 50, Pl. 8F Gynandrocarpa latericius Sluiter, 1904: 94, pl. 15, fig. 8- 11. Eusynstyela latericius, C. Monniot, 1991b: 14, fig. 4 A-C. Material Examined: MNHN: SI Eus 13 and IS-Indonesia, north Sulawesi, west of Manado, 1°23.SO'N; 124°44.60'E, 12 m. The colony forms a sheet about IO cm across. The zooids have 2/3 of their body included in the common test. They are purple-red with a large white strip between the siphons (Pl. 8F). Fixed they become brownish, the white strip remains as a paler mark. The test is naked and thin. The zooids are horizontal, the largest reaches 8 mm. The siphons are well apart, separated by 4 to 5 mm. The thin body wall is linked to the test with some elastic fibers around the siphons. The muscles are diffuse. About 12 tentacles of variable size, not regularly alternating, are inserted along a high crest. The prepharyngeal groove has only one crest and outlines a dorsal V. The dorsal tubercle is low with a longitudinal slit. The dorsal lamina has a plain edge, its height progressively increases from the dorsal tubercle to the esophagus opening. The endostyle is not directly at tached to the body wall but joined to it by a thin membrane. 242 Micronesica 29(2), 1996

Figure 49. Amphicarpa agnata (Kott, 1985);A , specimen opened along the ventral side. Eusynstye/a latericius (Sluiter, 1904): B, zooid opened along the ventral side; C, detail of a polycarp; D, different stages of the polycarps of the Figure 50 zooid. Monniot & Monniot: New Ascidians 243

Figure 50. Eusynstye/a /atericius (Sluiter, 1904): A, specimen opened along the ventral side; B, C, external and internal views of the ventral side. 244 Micronesica 29(2), 1996

The branchial formula is: R.E. 2 6 5 12 5 (5) 5 8 2 R. 1 JO 5 (5) 5 12 4 8 2 EL In the anterior part of the branchial sac there is an indication of 4 folds. Folds 1, 3, and 4 are obvious, the second on each side is reduced to 5 grouped sinuses which are not protruding. In the posterior part of the branchial sac, folds 2 and 4 on the right and fold 2 on the left disappear. This is not the result of a :flattening but in the median part of the branchial sac the sinuses interrupt. The first longitudinal vessel on the right of the dorsal lamina posteriorly moves aside from it (the interval being of 7 or 8 stigmata anteriorly up to 20 posteriorly). On the left side, the first longitudinal vessel is almost parallel to the dorsal lamina (interval of 8 stigmata anteriorly and 5 posteriorly). Between the folds the meshes are longer than wide, with 3 to 5 stigmata. On the folds the meshes contain 1 to 2 stigmata. Even where the rows of stigmata sometimes double, there are no parastigmatic vessels. The gut makes a small closed loop (Fig. 49B). The globular stomach has 12 folds. The caecum is straight and short. The intestine only forms a short loop and the bilabiated anus opens close to the cloacal siphon. The gonads are arranged in two irregular lines on each side of the endostyle. Each polycarp is made of an oval ovary and 2 testis follicles placed externally and anteriorly from it. The ovi duct opens above the testis by a flat aperture. No sperm ducts were observed but the testis follicles seem to open by a slit on each side of the female papilla. On the left side of the body, the most posterior polycarps are overlaid by the intestine. The endocarps are scattered over the body wall. No cloacal tentacles were ob served (Fig. 49B). In another colony we noted some differences. In formalin the common test is yellowish and the zooids are pale pink with a yellow spot between the siphons. The rim of the siphons is a thin red line. There are only few elastic fibers uniting the tunic and the body wall. The branchial sac has fewer longitudinal vessels, with higher folds number 2, almost disappearing in the posterior part. The branchial formula is: R.E. 1 7 4 9 2 7 3 9 1 R. 1 9 2 6 3 9 3 6 1 E.L. The stomach (Fig. 50) is globular and only has some incomplete ridges on the ventral side. Its pyloric limit is clearly seen. The caecum is small, curved, half hidden in a stomach depression. The anus is largely opened with a folded and undulated rim. The gonads are distributed in 2 lines on each side of the endostyle. The left one turns around the gut (Figs. SOB,C). The polycarps are not all at the same developmental stage. The youngest are encountered at the anterior and pos terior extremities of the lines and also ventrally. They are initially formed of 2 testes with short sperm ducts joined in a common papillae. Then, on the most ventral testis, develops a cluster of small oocytes. At last, the ovary extends and covers the ventral testis. It can extend ventrally on the body wall and partially cover the dorsal testis (Figs. 49C, D) but never totally. We have not seen the oviducts.The cloacal aperture is surrounded by a short velum with small tentacles. Remarks: Kott (1985) proposed only one species of Eusynstyela for the Indo Pacific region; E. latericius. In New Caledonia C. Monniot (1991b) recognized 3

(Figs. 51 E, F, G) and finally cross the branchial tissue to open inside the branchial branchial the inside open to tissue branchial the cross finally and G) 51 F, (Figs. E,

hermaphrodite gonads lengthen according to the maturity stage of the ovaries ovaries the of stage maturity the to according lengthen gonads hermaphrodite

to 10 hermaphrodite gonads on a single line, on the right (Figs. 51A, B, C). C). B, 51A, (Figs. right the on line, single a on gonads hermaphrodite 10 to

to the posterior part of the body. There are 2 male gonads on the left side and 8 8 and side left the on gonads male 2 are There body. the of part posterior the to

in a conjunctive tissue. The anus is smooth, slightly curled. The gonads are limited limited are gonads The curled. slightly smooth, is anus The tissue. conjunctive a in

complete, the anterior ones very short. There is a large curved caecum embedded embedded caecum curved large a is There short. very ones anterior the complete,

o od nec sd orsodn t h od l 3ad 4 3 and l, folds the to corresponding side each on folds low

lamina is low anteriorly and high posteriorly. The branchial sac has 3 rod-like, rod-like, 3 has sac branchial The posteriorly. high and anteriorly low is lamina

is no more than 1 sinus in a mesh. mesh. a in 1 sinus than more no is

and contain 2 or 3 stigmata crossed by a parastigmatic vessel. On the folds there there folds the On vessel. parastigmatic a by crossed stigmata 3 or 2 contain and

of the space between 2 consecutive sinuses. Between the folds the meshes are long long are meshes the folds the Between sinuses. consecutive 2 between space the of

teriorly . The extra sinuses form either against the dorsal lamina or in the middle middle the in or lamina dorsal the against either form sinuses extra . The teriorly

Sometimes the first or the last of the latter are only male. The sperm ducts of the the of ducts sperm The male. only are latter the of last the or first the Sometimes

Between the first fold and the dorsal lamina are 3 sinuses anteriorly and 9 pos 9 and anteriorly sinuses 3 are lamina dorsal the and fold first the Between

dorsal convergence of 2 to 3 sinuses indicates the remains of the second fold. fold. second the of remains the indicates sinuses 3 to 2 of convergence dorsal

formula is: is: formula

shape of a trapezium, marked with about 40 grooves; those on the posterior side side posterior the on those grooves; 40 about with marked trapezium, a of shape

caecum which is not, or only slightly, curved. Sluiter's drawings (pl. XV, figs 9, 9, figs XV, (pl. drawings Sluiter's curved. slightly, only or not, is which caecum

with round testes placed dorsally to the ovary, short gonoducts, and a short gastric gastric short a and gonoducts, short ovary, the to dorsally placed testes round with

in a transverse slit. The neural ganglion is close to the dorsal tubercle. The dorsal dorsal The tubercle. dorsal the to is close ganglion neural The slit. transverse a in

present colonies conform with the type having an irregular distribution of gonads gonads of distribution irregular an having type the with conform colonies present

with a slight dorsal indentation. The neural gland opens on the dorsal tubercule tubercule dorsal the on opens gland neural The indentation. dorsal slight a with

The type colony was partially figured in 1991 but not completely described. The The described. completely not but 1991 in figured partially was colony type The

the tentacles of the third order is variable. The prepharyngeal groove has two rims rims two has groove prepharyngeal The is variable. order third the of tentacles the

and regular. About 30 oral tentacles, in 3 orders, regularly alternate. The size of of size The alternate. regularly orders, 3 in tentacles, oral 30 About regular. and

Japan, Sri Lanka and Mauritius. Mauritius. and Lanka Sri Japan, In formalin the color turned brownish. The tunic is thin. thin. is tunic The brownish. turned color the formalin In

siphons are slightly protruding, the oral one larger, both with eight white spots. spots. white eight with both larger, one oral the protruding, slightly are siphons

stolons and covered with some sand. When living the color was red (Pl. 8G). The The 8G). (Pl. red was color the living When sand. some with covered and stolons

Holotype: Holotype:

Monniot, 1991 b, closely allied with, but distinct from, from, distinct but with, 1991 allied closely b, Monniot,

species; species;

tralia, Queensland, Arafua Sea, Indonesia, Viet Nam, Gulf of Siam, Philippines, Philippines, Siam, of Gulf Nam, Viet Indonesia, Sea, Arafua Queensland, tralia,

11) are wrong. wrong. are 11)

134°26.7l'E, 30 m. m. 30 134°26.7l'E,

The gut (Figs. 51 A, C, D) is located posteriorly with a large stomach in the the in stomach large a with posteriorly located is D) C, A, 51 (Figs. gut The

The organs can be seen through the thin body wall. The musculature is fine fine is musculature The wall. body thin the through seen be can organs The

The space between the two first folds is wider and in some zooids, a slight slight a zooids, some in and wider is folds first two the between space The

Description: The colony is made of a group of erect zooids, connected by by connected zooids, erect of group a of made is colony The Description:

Geographic Distribution: Distribution: Geographic

E. misakiensis misakiensis E.

MNHN: SI SI MNHN:

D.E. 2 5 5 5 5 2 D.E.

Watanabe & Tokioka, 1972; 1972; Tokioka, & Watanabe

Sto 19-Palau, Koror State, Rock Islands, 7°20.28'N; 7°20.28'N; Islands, Rock State, Koror 19-Palau, Sto

Stolonica limhata limhata Stolonica

Monniot Monniot

9 9

E. latericius latericius E.

Fig. 51, Pl. 8G 8G Pl. 51, Fig.

11 11

& &

Monniot: New Ascidians Ascidians New Monniot:

JO JO

2 R. 3 3 R. 2

new species species new

swdl dsrbtd n etr Aus western in distributed widely is

9 9

8 8

5 5 2 E.I. E.I. 2 5 5

E. a/iena a/iena E.

. .

latericius latericius

. .

In the anterior partthe partthe anterior the In

and and

(Sluiter 1904). 1904). (Sluiter

E. phiala phiala E.

245 245 C. C. 246 Micronesica 29(2), 1996

Figure 51. Stolonica limbata n. sp.: A, B, C, ventral, right and left sides of a spec imen; D, same specimen ventrally opened; E, F, G, successive development stages of the right side hermaphroditic gonad; H, I, J, left side testes at the same stages. cavity. In contrast to the situation in most Stolonica species, the sperm ducts of the left side gonads also lengthen (Figs. 51H, I, J). Larvae are incubated on the right, in the space limited by the gonads and a large transverse endocarp. The Monniot & Monniot: New Ascidians 247 characteristic location of the large endocarp which limits the extension of the larvae justifies the species name S. limbata (limbatus = edged). Other endocarps are present; they do not show the characteristic symmetrical disposition. The clo acal siphon has a row of small tentacles (Fig. 51D). Remarks: The genus Stolonica sensu stricto, is homogeneous, characterized by 1 line of gonads on each side, male on the left and male and hermaphroditic on the right. We do not follow Kott (1985, 1990b), who successivelymixed species of the genera Stolonica and Amphicarpa. Sto/onica aluta Kott, 1985 which has on the right side "sac-like ovaries crowded in amongst male folicles" (p. 233) instead of hermaphroditic gonads, is not a Stolonica but an Amphicarpa. The tropical species of Stolonica have the following characters: S. inhacae (Millar, 1956) has only one branchial fold and a spherical stomach. S. reducta (Sluiter, 1904) has 3 branchial folds, an long stomach, and always short sperm ducts. S. sabulosa Monniot, 1972, from Brazil and Bermuda, is close to the Palau species. It has 3 branchial folds, the same kind of stomach and gonads. But its gonads are located in the middle of the ventral side instead of the posterior part, and there is no endocarp limiting the brooding of the larvae. S. truncata Kott, 1972 has 3 branchial folds, a short stomach, and very elon gate male gonads with short sperm ducts. S. variata C. Monniot, 1988 is reminescent of the Palau species, but the stomach is more elongate with a maximum of 25 grooves, the male gonads are more numerous on the left side (5 at least) with always a short sperm duct, whereas the sperm ducts of the right gonads are always long. S. vesicularis Van Name, 1918, as described by Kott (1985), is characterized by 3 branchial folds on the right side and 2 on the left, a triangular stomach with 14 very short grooves. The 3 male gonads on the left and 9 gonads on the right (the number of hermaphrodite ones among them being not precised) have a long sperm duct. Kott's description is rather different from the original one by Van Name (1918) and from that by Millar (1975). The latter authors reported the species has a short oval stomach with about 30 complete grooves. The left gonads are more numerous than the right ones and are in a V along the endostyle and intestine. No true hermaphrodite gonads were described. This species probably belongs to the genus Amphicarpa. Actually, the species described by Kott (1985) cannot be Stolonica vesicularis.

Polycarpa argentata (Sluiter, 1890) Fig. 52, Pl. 8H Styela argentata Sluiter, 1890: 340 Polycarpa argentata: Kott, 1985: 148 fig. 66, and synonymy; C. Monniot, 1987b: 277 fig. 1 A,B, New Caledonia. Material Examined: MNHN: Sl Pol B 307,308, 309-Palau, Mutremdiu, Barrier Reef, 7°16.51'N; 134°3l .55'E, 5 m.-Papua New Guinea, Madang, Nagada Har bor, 5°09.35'S; 145°48.00'E, 1 m.

be be

tough tough

tunic tunic is

body body

A, A, specimen opened along the ven

: :

1990)

(Sluiter, (Sluiter,

Microncsica 29(2), I 996 996 Microncsica 29(2), I

argentata argentata

A A

gonad. gonad.

B, B,

; ;

Polycarpa Polycarpa

3 3

a a

V, V,

0 0

tral tral side

a a

n n

......

Figure Figure 52.

The The body wall is not thick and slightly translucent. The muscular are fibers

The The specimens were erect, brownish red and covered with epibionts in life

brownish-red. brownish-red. The gonads that are included in the body wall thickness can

exteriorly, exteriorly, and greyish softer interiorly.

ored. ored. The tunic lining pink, is without ornamentations. The

stripes stripes and greenish-white spots. In formalin, the test is wrinkled and col silver

(Pl. (Pl. 8H). The siphons are long, with a circular opening and internally 8 white 248 248

60, Indonesia, Saleyer, Timor, and Ambon Islands. Islands. Ambon and Saleyer, Timor, Indonesia, 60,

Styela captiosa captiosa Styela

una Philippines Guinea,

cover each other. The sinus number is the same but the space between the folds folds the between space the but same the is number sinus The other. each cover

Material Examined: MNHN: Sl Pol B 303-Chuuk lagoon, 7°22.63'N; 7°22.63'N; lagoon, 303-Chuuk B Pol Sl MNHN: Examined: Material

Synonymy: see Kott, 1985: 1985: 149 see Kott, Synonymy:

grows in larger specimens larger in grows

h gnd (Fig gonads The

specimen are low but in New Caledonia and Australia they are higher and slightly slightly higher and are they Australia and in New Caledonia specimen low but are

gonads is also greater than in other regions. The branchial folds of the present present the of folds branchial The regions. other in than greater also is gonads

mata in a mesh, and 1 to 2 on the folds. There are no parastigmatic vessels. vessels. parastigmatic no are There folds. the on 2 1 to and mesh, a in mata

and New Caledonian samples and to that of of that to and samples Caledonian New and

segmentation, were found in the cloacal cavity. cavity. cloacal the in found were segmentation, h is iu ntergtprs sd rm h osl aia 8simt ante stigmata (8 lamina dorsal the from aside parts right the on sinus first The

numerous, up to 250 on the right and 150 on the left. Some eggs, without visible visible eggs, Some without left. the 150 on and right the 250 on to up numerous,

Marshall Islands, Western Australia, Queensland, New Caledonia. Caledonia. New Queensland, Australia, Western Islands, Marshall

isodiametric intestine forms a regular curve forms a regular intestine isodiametric has 4 low, widely separated folds. We counted: counted: We folds. widely low, separated 4 has

an olive-shaped stomach with few and not well defined internal grooves. The The grooves. internal defined well not and few with stomach olive-shaped an dostyle is high, attached to the body wall by a thin membrane. The branchial sac sac branchial The membrane. thin a wall by body the to attached is high, dostyle

ipehl.Te o osl aia nice h spau etac. h en The entrance. esophagus the encircles lamina dorsal low The hole. simple a

than specimens recorded elsewhere, which do not exceed 3 cm. The number of of number The cm. 3 exceed not do which elsewhere, recorded specimens than

it draws a deep dorsal V. The dorsal tubercle is slightly protruding, its opening is is opening its is slightly protruding, tubercle dorsal The V. dorsal deep a draws it

the Palau Islands. The present specimen and those described from Palau are larger larger are Palau described those from specimen present and The Islands. the Palau

agus opening progressively lowering. Between the folds there are about 10 10 stig about are there folds the Between lowering. progressively opening agus and small ones intercalated). The prepharyngeal groove has only one high crest; crest; high one only has groove prepharyngeal The intercalated). ones small and

riorly and more than 40 in the medium part). The branchial folds join the esoph the folds join branchial The part). medium the 40 in than more and riorly

On the right side the 3 first sinuses are more brought together than the others. others. the than together brought more are sinuses first 3 the side right the On

truding in the cloacal cavity in the cloacal truding

seen in transparency. There are 16 tentacles in 3 orders (2 orders of large ones ones large of orders (2 orders 3 in 16 tentacles are There transparency. in seen

151°50.70'E, 151°50.70'E, 22 m.

This species is recorded from the north coasts of Australia, Indonesia, New New Indonesia, Australia, of coasts north the from species recorded is This

The gut (Fig. 52A) has a translucent wall. The short esophagus opens into into opens esophagus short The wall. translucent a has 52A) (Fig. gut The

Remarks: This specimen well corresponds to the descriptions of Australian Australian of descriptions the to well corresponds specimen This Remarks:

Geographic Distribution: Previously known from Palau, Gilbert Islands, Islands, Gilbert Palau, from known Previously Distribution: Geographic

In the posterior half the first folds lower and become only a group of sinuses. sinuses. of group only a become first lower the and folds half posterior the In

R.E. 6 6 R.E.

12 12

Sluiter, 1885: 1885: 202 Sluiter,

. .

6 6

52B) are included in the body wall and are only slightly pro slightly only are and wall body the in included 52B) are

Polycarpa aurata aurata Polycarpa

12 12

, ,

Chuuk and Palau Islands. Islands. Palau and Chuuk

7 7

Polycarpa captiosa captiosa Polycarpa

JI JI

. .

Monniot & Monniot: New Ascidians Ascidians New Monniot: & Monniot

6 6

. .

The gonoducts do not protrude. The gonads are very very are The gonads protrude. not do gonoducts The

13 13

, ,

or or

fig. 67, pl. 3c. 3c. fig. 67, pl.

Fig. 53, Pl. 9A 9A 53, Pl. Fig.

, ,

pl. 9, fig. 4-7, Indonesia, Billiton Island; 1904: 1904: Island; Billiton Indonesia, fig. 9, 4-7, pl.

(JO (JO

(Quoy (Quoy

1 3) 0.R. 3 3 1 3) 0.R.

. .

(Sluiter, 1885) 1885) (Sluiter,

The rectum opens opens in rectum a The scalloped anus

& &

Gaimard

P. iwayamae iwayamae P.

JO JO

4 4

12 12

, ,

1834) 1834)

7 7

12 12

Tokioka, 1950 1950 from Tokioka,

6 6

13 13

5 5 E.L.

249 249 . .

mm mm

5 5

n n

,_. ,_.

internal internal side of the body wall; B,

1 1

/ /

' '

, ,

Palau, Palau, Rock Islands, Marine Lake,

/ /

,, ,,

,--" ,--"

1885): A 1885):

, ,

------

'cf--

Pol Pol B 299

\)(\ \)(\

I I

(Sluiter

Micronesica 29(2), 1996 1996 29(2), Micronesica

aptiosa aptiosa

MNHN: MNHN: SI

Kott, Kott, part: Australia. 1985? 184,

Tokioka, 1950: 136, fig. 14, pl. 10, fig. 1, Palau Palau fig. Islands; 1, pl. 1967: 10, fig. 14, Tokioka, 136, 1950:

Polycarpa Polycarpa

. .

dorsal dorsal tubercle.

The siphons are 4 lobed, with gold yellow patches, The patches, siphons are on 4 lobed, with 4 the gold yellow oral siphon

. .

igure igure

The The tunic is about 2 mm thick. The body wall is opaque, l mm thick and

The The specimen of 8 cm in length was attached by the posterior half of the

m. m.

0 0

171, fig. 70, Palau Palau 70, fig. and 171, Mariana Islands.

1 1

the body wall as as the body they pale, remain wall devoid granules. pigment of The oral tentacles

turned turned black-brown, the tunic yellowish.

and and 2 on the cloacal siphon, and uniformly red inside. In formalin the siphons

Material Material Examined:

(Pl. 9 A) 9 (Pl. granules, granules, is wrinkled with polygonal areas. Gonads and endocarps contrast on

Po/ycarpa Po/ycarpa papil/ata:

long long and well separated, the deep red test is tuberculated, with some epibionts yellow. yellow. The musculature cannot be seen. Internally the body wall, full of gold

ventral ventral the side, upper part of the body was erect. When the living, siphons are

Polycarpa Polycarpa captiosa: 25 Monniot & Monniot: New Ascidians 251 are short, not pigmented, about 20 in 3 orders of size. The prepharyngeal groove has 2 high equal rims against the tentacle ring and the branchial sac. The dorsal V is filled with a protruding dorsal tubercle with multiple openings (Fig. 53B). The dorsal lamina, branchial sac and gut have been eviscerated. A part of the branchial sac remaining attached to the prepharyngeal groove, allows determi nation that the branchial folds were high and not pigmented. On the left side of the body remains the shape of the gut with a double curve, a well marked sec ondary bend (Fig. 53A). There, the body wall lacks the polygonal pattern present elsewhere. The endocarps which were close to the gut stayed in place. The gonads (Fig. 53A) are grouped in an irregular row on each side. They are protruding into the cloacal cavity. The testis follicles and oocytes are poorly developed. The en docarps are numerous and upright, those which are close to the cloacal siphon are long tending to penetrate into it. Remarks: Several authors, including Tokioka (1952) and Millar (1975), de scribing P. captiosa had emphazised the similarities of this species with P. papil /ata, but separated the two species. Kott (l 985) considered them as synonyms and thought that specimens attributed to P. captiosa were particularly large specimens of P. papillata in which the "concave shape" and widely spaced siphons corre sponded to animals that "lie horizontally". The examination of the type specimen of P. captiosa has shown it to be identical to the Palau sample examined here. Sluiter's (1885) description has numerous errors. Sluiter (1885 pl. IX fig. 6) mis interpreted the narrow groove located between the prepharyngeal groove and the dorsal tubercle as the dorsal tubercle. The latter is actually very large, with several openings in transverse slits. The anus is not constricted, but widely opened and without lobes. P. captiosa and P. papilla ta differ in gonad distribution, properties of the test, the body wall (thin, translucent and brittle in P. papillata; thick, opaque, muscular with a pigmented internal layer, in P. captiosa), the gut loop and stomach shape, and the anal border (deeply lobed in P. papillata, smooth in P. captiosa). Geographic Distribution: Indonesia, Palau, Mariana Islands, and perhaps present in Australia.

Polycarpa cryptocarpa (Sluiter, 1885) Pl. 9C Material Examined: MNHN: Sl Pol B 298- Palau, Rock Islands, Topkukau, 7°21.03'N; 134°27.0l'E, 7 m. The sample in this collection conforms to the description of specimens of this species by Tokioka, 1950. The living specimens have a dark brown tunic with wide siphons (Pl. 9C). The siphonal rim is underlined with white dots. This aspect is somewhat different from that observed in New Caledonia, where the internal tunic of the siphons was uniformly white. Geographic Distribution: Japan, Philippines, Indonesia, Indies, Palau, So lomon Is., Gilbert Is., Marshall Is., Queensland, New Caledonia.

6 6 E. L.

20 20

9 9

22 22

9 9

20 20

9 9

15 15

. .

is is partly encircled by a deep hollow

(Sluiter, (Sluiter, 1885)

(Herdman, (Herdman, 1906)

It It

0 0 R. 3

8 8

Pl. Pl. 9D

4 4

13 13

Micronesica Micronesica 29(2), 1996

Sl Sl Pol B 304--Papua New Guinea, North coast,

10 10

Figs. Figs. B, Pl. 54A, 9E

Sluiter, Sluiter, 192 1885:

22 22

8 8

Polycarpa Polycarpa papillata

Po/ycarpa Po/ycarpa captiosa.

Kott, Kott, C. 89; fig. Monniot, 192, 1985: fig. 3. 283, 1987b:

20 20

MNHN: MNHN:

Polycarpa Polycarpa pigmentata

Kott, Kott, fig. 85-86 184, and 1985: synonymy except references

8 8

Herdman, Herdman, 318. 1906:

and and

22 22

Styela Styela

R. R. E. 6

The The gut (Fig. is very posterior. 54B)

The The ventral folds are relatively high but do not cover each other; they are

The large specimen is entirely black black the except pure entirely is The internal white large specimen of the side The The body wall is black, opaque, to 1 2 mm thick. The muscles do not form

This species has has recorded New been Caledonia, in This species Australia, Indonesia, Phil

has has 5 folds on the right side and 4 on the left.The longitudinal vessels are

of of the body wall. When the intestine up, is lifted its trace on clearly visible is the

body body The wall. short enlarged in a stomach esophagus slightly gives with internal

it, separated it, separated about by stigmata, to back 20 coming near before it, close the esoph

sinus on on sinus the right anteriorly is to closer the dorsal lamina, then it runs apart from

less protruding protruding less dorsally. The additional fold on the right is complete. The first 2 2 or on 3 the folds. There are no parastigmatic vessels.

agus. The disc-shape tissue, circling the esophagus opening, is very thick; thick; the very folds esophagus the opening, is circling The agus. disc-shape tissue,

arranged arranged as follows:

only, only, irregularly alternated and more numerous dorsally. The prepharyngeal It It

dorsal lamina is long, smooth, low and only slightly increases in height posteriorly. posteriorly. height in increases and slightly only long, smooth, low dorsal lamina is

pigments. pigments. The oral tentacles are short, colored ochre in They life. are about 10 The endostyle is attached attached The is endostyle to body the The branchial wall. contains sac black cells.

diameter, diameter, sometimes with hollow whitish concretions fragile and patches of dark

internal internal part. The body contains wall large translucent of 0.5 to vesicles 0.6 mm flat flat dorsal tubercle all occupies the V and has multiple openings (Fig. The 54A).

groove groove has only one crest and forms a deep prolonged V by a small groove. The

Kavieng, Kavieng, west of Albatross Channel, m. 2°45.06'S; 150°42.84'E, 10 are are present, mostly encrusting bryozoans.

Material Material Examined: MNHN: SI Pol B 317-Papua New Guinea, New Ireland, animal. animal. The tunic remains soft and black in its whole thickness. Some epibionts

Polycarpa Polycarpa pigmentata: black. black. The siphons obvious on the animals living are 4 cm apart in the retracted

siphons siphons and their ochre margin (Pl. 9E). When fixed the reflex tunic becomes a a thin layer but are distributed in the whole thickness, especially in the most

join join it without There lowering. are 7 to stigmata 10 in a mesh between the folds,

Styela Styela pigmentata:

Barrier reef of Madang, 5°10.21 'S; 'S; Barrier m. reef of 10 Madang, 145°50.37'E, 5°10.21

ippines, ippines, Palau Islands, Lanka, Sri and Madagascar.

Materiel Materiel Examined:

Polycarpa Polycarpa papillata:

Styela Styela (Polycarpa) papilla/a

concerning concerning 252 252

more than 10 on each side, developed enough to be seen in the body wall, but but wall, body the in seen be to enough developed side, each on 10 than more siphon possess a thick crest bordered with small tentacles. tentacles. small with bordered crest thick a possess siphon

follicles are lobed. There are 2 large foliated endocarps in the gut loop. The cloacal cloacal The loop. in the gut endocarps 2 are foliated large follicles There lobed. are

many more if some of the pigment masses represent very young gonads.The testis testis gonads.The young very represent masses pigment the of if some more many

body wall and the ducts are not protruding. The gonads are not numerous, no no numerous, not are gonads The protruding. not are ducts the and wall body

gonads are not functional in this specimen; they are completely included in the the in included completely are they specimen; this in functional not are gonads

gut is only attached to the body wall at the level of the endocarp and rectum. The The rectum. and endocarp level the the of at wall body the to attached is only gut

linked to the dorsal lamina but to the body wall. The anus is slightly lobed. The The lobed. slightly is anus The wall. body the to but lamina dorsal the to linked

grooves that are not visible exteriorly. There is no caecum. The rectum is not not is rectum The caecum. no is There exteriorly. visible not are that grooves

. .

pigmentata pigmentata

Remarks: This specimen better correspond to Kott's ( 1985) description of of 1985) description ( Kott's to correspond better specimen This Remarks:

Figure 54. 54. Figure

lycarpa rima rima lycarpa

than to that of C. Monniot (1987b). It differs from the New Cal- New the from differs It (1987b). Monniot C. of that to than

Polycarpa pigmentata pigmentata Polycarpa

n. sp n.

Monniot Monniot

: C, dorsal area; D, specimen opened ventrally. ventrally. opened specimen D, area; dorsal : C,

& &

(Herdman, 1906): A, dorsal tubercle; B, gut. gut. B, tubercle; 1906): dorsal A, (Herdman,

Monniot: New Ascidians New Ascidians Monniot:

Po 253 253

4 4 E.L.

21321418 21321418

3 3

crack). crack).

25 25

= =

new new species

(rima (rima

7) 7) 0 R. 0

(22+ (22+

Micronesica Micronesica 29(2), 1996

Figs. Figs. D, 54C, Pl. 9F

2 2

21 21

Polycarpa Polycarpa rima

3 3

21 21

4 4

17 17

4 4

The The dorsal lamina is long, low with a plain margin. The branchial sac

R.E. R.E.

. .

The The first on sinus to the close left is side the dorsal lamina but the fold first

About About 40 tentacles are distributed in 3 or 4 orders on a single circle. The

The The body opaque. is wall bands The muscles are dark brown and they form

Description: The specimens were erect, 8 x 3 cm, cm, 3 attached x erect, 8 were a by Description: part small The specimens of

Geographic Geographic Distribution: Lanka, for Except Sri location, the type and West

folds are are folds not to close high very with each sinuses other. The between the sinuses

is doubled soon in the anterior part of the branchial sac. Posteriorly, all the sinuses sinuses the all anterior the Posteriorly, part branchial doubled the soon in sac. of is

is thick thick and is tough. On are external the the large transverse side and sinuses mus

constituting constituting the first fold part but remain in two distinct groups. The branchial

mula mula is:

cularized. Each cularized. dermato-branchial trabeculum has The a branchial muscles. for

openings

The The dorsal tubercle is large and protruding with a number small of independent

posteriorly posteriorly into to a mm 2 groove in front long 3 of the dorsal lamina (Fig. 54C).

tract. tract. This divided surface by layer furrows is deep in more quadrangular or less

prepharyngeal prepharyngeal crests. made equal It of two low has is groove a dorsal extended V

fields, as suggested by the species name name by as suggested the species fields,

numerous numerous on the the endostyle, branchial tissue, the endocarps and the digestive

cloacal surface surface the of cloacal body are wall scattered rare black are which more granules

and and contains numerous blood lacunae, the In gonads the and vesicles. clear large

a a dense network. The body wall is to 1 2 mm in thickness, with two parts: one

brown brown in its total and thickness soft. is

external external containing the musculature, the internal one of a light brown is fibrous

externally golden and white internally, internally, golden and but white externally tunic in formalin turns whole the dark

siphon siphon is 3 cm from the oral siphon; when alive (Pl. 9F) the siphons are bare,

the the posterior with strong side left hairs. The oral aperture terminal, is the cloacal

(Coral (Coral km Sea), southwest of 140 Port Moresby, m. 30

Holotype: Holotype: MNHN: SI Pol B 318-Papua New Guinea, Eastern Fields Atoll

Caledonia. Caledonia.

Japan, Japan, Indonesia, Palau, Philippines, Marshall Is., Solomon Chuuk, Is., New Fiji,

tract tract between the Austalian and New Caledonian samples.

anus. The present specimen is intermediate intermediate for anus the the is and shape of digestive anus. present The specimen

ern ern Australia, the geographic distribution of this remains species in the Pacific:

sinuses contain contain sinuses The fibers. white gut forms a larger loop with a smooth rimmed

body body wall has the same structure but is pigmented. less The tentacles are more

perhaps perhaps related to the body shape). The branchial sac is more resistant and the

numerous, numerous, the dorsal tubercle more protruding, the dorsal lamina shorter (this

massive massive and quadrangular, there is a stiff external and soft internal tunic. The

edonia edonia specimens by many small details. In New Caledonia the shape is more 254 254

pigmented bands are limited to the upper part of the siphons. The surface of the the of surface The siphons. the of part upper the to limited are bands pigmented

has 8 brown stripes alternating with 8 paler ones with bright yellow dots. The The dots. yellow bright with ones paler 8 with alternating stripes brown 8 has

imen. imen.

eyyugseieswoetnc eent ue otespotn aut spec adult supporting the to fused not were tunics whose specimens young very

cm, their tunics fused at their base only or along almost their whole length. Out Out length. whole their almost along or only base their at fused tunics their cm,

found. found.

of the tunic, the individuals have the same size, no young specimens or buds were were buds specimens or young size, same no the have individuals the tunic, the of

MNHN: MNHN:

intertidal; intertidal;

1.5 cm, with tunics fused at their base. On the adult specimens we found some some found we specimens adult the On base. their at fused tunics with 1.5 cm,

MNHN: SI SI MNHN:

Material Examined: Holotype: MNHN: Sl Pol.B 316--Palau, Marine Lake; Lake; Marine 316--Palau, Pol.B Sl MNHN: Holotype: Examined: Material

Polycarpa nigricans: nigricans: Polycarpa

Polyandrocarpa nigricans nigricans Polyandrocarpa

? ?

length of the genital ducts. ducts. genital the of length

molguloides, molguloides, rim and not protruding genital ducts. ducts. genital protruding not and rim

1 1 m;

a simple dorsal tubercle. It differs from from differs It tubercle. dorsal simple a

and sand, fewer vessels in the folds, lobed testis follicles, short genital ducts, and and ducts, genital follicles, testis short lobed folds, fewer vessels the in sand, and

Herdman, 1882 (see Kott, 1985, C. Monniot, 1987b). 1987b). Monniot, C. 1985, Kott, (see 1882 Herdman,

in polygonal areas, is similar in in similar is areas, polygonal in

tentacles on its margin. The tentacles are scattered inside the siphon. siphon. the inside scattered are tentacles The margin. its on tentacles

loop. The cloacal aperture is circled by a fleshy undulated crest having small small having crest undulated fleshy a by circled is aperture cloacal The loop.

and long, they encircle the ovary. There are 2 very large endocarps in the gut gut the in endocarps large very 2 are There ovary. the encircle they long, and

button located at the base of the female papilla. The testis follicles are numerous numerous follicles testis are The papilla. female the of base the at located button

layer and by large female papillae. The male papilla, much smaller, opens on a a on opens smaller, much papilla, male The papillae. female large by and layer

bedded in quadrangular fields of the body wall, and each field may contain up to to up contain field each may and wall, body fields the of quadrangular in bedded

in an anus with has finger-like lobes on the margin. The gonads are deeply em deeply are gonads The margin. the on lobes finger-like has with anus an in

larged and has few internal grooves, hardly visible. The cylindrical intestine ends ends intestine cylindrical visible. The hardly grooves, few internal has and larged

3 of them. They are only visible by a light color through the body wall internal internal wall body the through color light a by visible only are They them. of 3

are in a mesh between the folds. There are no parastigmatic sinuses. sinuses. parastigmatic no are There folds. the between mesh a in are

folds correspond to the most ventral sinuses of the preceeding fold. 10 10 stigmata fold. preceeding the of sinuses ventral most the to correspond folds

Polycarpa pedunculata: pedunculata: Polycarpa

27.07'N; 124°44.59'E, 124°44.59'E, m. 7 27.07'N;

In life, the test is orange, the rim of the siphons blackish. The siphon lining lining siphon The blackish. siphons the of rim the orange, is test life, the In

The specimens collected in the intertidal area (Pl. 9G) are much smaller, 1 1 to smaller, much are 9G) (Pl. area intertidal the specimens in collected The

The specimens collected on a ship hull were grouped each measuring 2 to 3 3 to 2 measuring each grouped were hull ship a on specimens collected The

Remarks: The gonad disposition, in an internal layer of the body wall wall layer of divided body the internal in an disposition, gonad The Remarks:

MNHN: SI SI MNHN:

h dgsietat Fg 5D om obe op h tmc i o en not is stomach The loop. double a forms 54D) (Fig. digestive tract The

Sl Pol.B 296--Indonesia, west ofManado, 1 km West ofMurex Resort, Resort, 1 West ofMurex km west ofManado, 296--Indonesia, Pol.B Sl

MNHN: MNHN:

Pol B 301-Palau, Airai Channel, ship hull, 7°21.03'N; 134°30.95'E, 134°30.95'E, 7°21.03'N; hull, ship Channel, Airai 301-Palau, B Pol

a species from southern Australia, by the tunic covered with hairs hairs with covered tunic the by Australia, southern species from a

Pol.B 302, Koror State, Rock Islands, 7°18.48'N; 134°28.12'E, 134°28.12'E, 7°18.48'N; Islands, Rock State, Koror 302, Pol.B

Monniot Monniot

SI Pol.B 344, Palau, Marine Lake, 7°17.7'N;l34°27.3'E; 7°17.7'N;l34°27.3'E; Lake, Marine Palau, 344, Pol.B SI

Polycarpa tokiokai tokiokai Polycarpa

C. Monniot, 1987: 1987: Caledonia. fig.6A-C, 288, New Monniot, C.

non Heller, 1878, Tokioka, 1979: 1979: 91, fig. 5, 1878, Philippines. Tokioka, Heller, non

Monniot & Monniot: New Ascidians Ascidians New Monniot: & Monniot

& &

P. mytiligera mytiligera P.

Monniot, 1987: 1987: 234, Philippines. fig. 5, Monniot,

Fig. 55, Pl. 9G 9G 55, Pl. Fig.

P. rima rima P.

P. mytiligera mytiligera P.

new species species new

(Savigny, 1816) 1816) and (Savigny,

is essentially characterized by the the by characterized essentially is

which has a smooth anal anal smooth a has which

P. rima rima P.

differs from from differs

P. molguloides molguloides P.

255 255 P. P.

detail detail of the

, ,

2 2 EL

8 8

2 2

2 2 EL 7 1

8 8

JO JO

2 2

3 3

· ·

9 9

JO JO

.--:/ .--:/

4 4

{ {

6 6

\o, \o,

-~~--

0 0 D.L. 2 1 7

: A, A, holotype : opened ventrally;

0 0 D.L. 1

9 9

2 2

8 8

A A

n. n. sp

3 3

9 9

Micronesica 29(2), 29(2), 1996 Micronesica

8 8

2 2

3 3

10 10

is is about mm thick, 1 soft and nacreous internally.

9 9

3 3

It It

6 6

Polycarpa Polycarpa tokiokai

R.E. R.E. 2 2 7

R.E. R.E. 3

gonad. gonad.

moves moves aside from the dorsal lamina, separated from it anteriorly by 4

It It

Figure Figure 55.

The The first sinus on the right of the side dorsal lamina more is developed than

While While in a counted: we specimen small

The The thin musculature is mostly longitudinal. The internal side of the body

6 6

others. others.

other. other. The branchial formula in a is: large specimen

high. The The high. branchial not do is sac it has not folds pigmented; which 4 each recover

others small and and others small The prepharyngeal irregular in size. groove has equal and 2 rims

crypt. crypt. The neural ganglion is long. The dorsal lamina is long, smooth edged and

to to 30 long and thin tentacles in 2 or 3 orders of size and, between them, some

curves in in a The V. curves deep dorsal has tubercle a large irregular opening from a wide

encloses a deep dorsal V. The The dorsal V. a deep anterior the flattens dorsal in part, one encloses posterior the

test is smooth smooth is test or it has small There tubercles. are epibionts. some The test turned

wall contains contains wall whitish granules more abundant in the anterior part. There are 16

dark dark brown in alcohol. 25 mens an unperforated area may develop posteriorly, on the right side the right onof the dorsal developmay posteriorly, area mens unperforated an

edge, somewhat curled or fringed with small lobes. We have not seen a caecum. a seen not have We lobes. small with fringed or curled somewhat edge, lreedcr ocpe h etr itsia lo; t satce t te body the to attached is it loop; intestinal entire the occupies endocarp large A wall, the stomach and the intestine. The gonads (Fig. 55B) (Fig. They gonads intestine. the The are characteristic.and wall, stomach the speci large In opening. esophagus the at 30 stigmata to 20 by and stigmata 6 to

They areThey erect in the cloaca) cavity. in areThey a embedded tissue thick containing transver are meshes branchial The stigmata. of number the diminishing lamina, sally elongated. 8 to 12 stigmata were counted in a mesh between the folds and 6 and folds the between mesh a in were 12 counted stigmata to 8 sally elongated. o3o h od.Teeaen aatgai vessels. parastigmatic no are There folds. the on 3 to

number of polycarps varies from less than 10 to about 40 in adult specimens. The 40 in adult 10 about less to fromvaries than polycarpsof number smooth a with anus widely opened a in ended wall, body the to attached is long, cloacal tentacles are thread-like on small crests scattered around the siphon. the around scattered crests small on thread-like are tentacles cloacal

r lne t tebd wl n a emt-rnha snsb ter itl part. distal their by sinus dermato-branchial a and wall body the to linked are

granules. They possess 2 rows of external testes and an internal ovary. The sper The ovary. internal an and testes external of rows 2 possess They granules. matozooids are collected in 2 sperm ducts joining in a single papilla, slightly pro single a papilla, in ducts joiningsperm2 collected in are matozooids

ail.hr ae ueos rnls n h bd wl rud h gnd. The gonads. the around wall body the in granules numerous are papilla.There

carpa nigricans carpa & h niiul a h ruet oik' ue oicueti pce no the species into this include to used Tokioka's argument the was individuals the genus

zooids more or less fused by a part of their tunic into the genus the into tunic their of part a by less fused or more zooids male the from far opens is andwide oviduct The ovary. the to anteriorly truding when they were not issued form a simple egg, but budded from lysed zooids. The lysed zooids. from budded simple egg, a but issued form not werethey when set f h gnd i seiesfo Piipns Idnsa n Plu pro Palau, and Indonesia Philippines, from specimens in gonads the of aspect rdn it tecoc)cvt,de o crepn ete o te descriptions other to either correspond not does cavity, cloaca) the into truding of h ohr ye pce f hs uhr n i o ctd n h 1905 Michaelsen's the in cited not is and author this of species type other the eiino elrstps atee (1905) redescribed Hartmeyer types. Heller's of revision the aggegates collected at Mauritius by Dr. by Mauritius aggegatescollected the at in Hrmyr oe ta teseiswsaudn i artu ad ht nu that and Mauritius in abundant species was the that noted Hartmeyer tion. having species the placed systematically have (1987b) Monniot C. and (1985) merous specimens were stored in the museums of Kiel and Berlin. aggregated specimens Kiel of The and museumsin the merous were stored specimens observed by Hartmeyer (1905) were probably dispersed and some zo some and dispersed (1905) probablywere Hartmeyer by specimens observed oids are at the Zoological Museum of Copenhagen, in the Senckenberg Museum Senckenberg the in Copenhagen, of Museum Zoological the at are oids rectum closed thecurve; a forms intestine 15 the outside; groovesvisible the from in Frankfurt and in the Museum of Wien. We have examined all these Wien. of specimens,examined We have Museum the in and in Frankfurt l aee atee, 1905 Hartmeyer, labeled all ical Museum of Copenhagen as the holotype. We believe that all specimens can specimens all believe that We holotype. the as Copenhagen of Museum ical ecniee sblnigt h yesre xmndb elr(88,ad are (1878), and Heller by series examinedtype the to belonging as considered be paratypes. The species from Mauritius is different from the species described species the from different is Mauritius from species The paratypes. above.

ono (1991) as Monniot

P. nigricans P.

The digestive tract is not voluminous (Fig. 55A). about (Fig.digestive has The is long voluminousThe stomachnot tract

Remarks. This two populations, correspond to the specimens of the to correspond populations, two This Remarks. We have searched for Heller's type of type Heller's for searched have We

Three different species have been assigned to assigned been species have different Three

Polyandrocarpa,

or to Heller's specimens. Heller's to or

ecie rmtePiipnsb oik (90)ad Monniot (1970b) and Tokioka by Philippines the from described

. nigricans. P.

even without any budding evidence. Since this time, Kott time, evidence. Since this budding any without even

Monniot

Kott (1985) considered the specimen in the Zoolog thespecimen in (1985)the considered Kott

The complete fusion of the test at the base of base the at test the of fusion complete The

Monniot: New Ascidians New Monniot:

Mobius

. nigricans, P.

. nigricans. P.

and

used for Heller's descrip Heller's for used

. nigricans P.

ti o lctd among located not is it

Polycarpa,

after one of one after

Polyandro

even

257 258 Micronesica 29(2), 1996

(1) Polycarpa nigricans stricto sensu: Polycarpa nigricans Heller, 1878: 174, fig. 80, Mauritius; Hartmeyer, 1905: 390, pl. 13, fig. 1, 12-14, Mauritius; Vasseur, 1967: 115, pl. 5, fig. 37--40, Mauritius. This species has aggregated or solitary zooids, 2 to 3 cm high with gonads included in the body wall and only slightly protruding in the cloacal cavity with large projecting gonoducts (Hartmeyer 1905, fig. 14). Each polycarp is enclosed in a mass of clear vesicles contrasting with the dark granules of the body wall and the area around the dorsal tubercule. The testis lobes are round, small, not regularly arranged as noted by Vasseur (1967, fig. 40). (2) Polycarpa nigricans non Heller, 1878: Kott, 1985: 174, fig. 80, pl. 3f, West Australia; C. Monniot, 1987: 286, fig. 5, New Caledonia. This species is solitary or aggregated, with a black tunic in life, and gonads deeply embedded in the body wall which cannot be seen on the internal surface. The gonoducts are not or only slightly protruding. The testis lobes are long on each side of the ovary. The West Australian samples have a complex dorsal tubercule. This is not the case in New Caledonian specimens. (3) Po/ycarpa tokiokai n. sp. The polycarps are clearly protruding into the cloacal cavity. The testis follicles are long, regularly arranged around the ovary. The polycarps are encircled by whitish granules. Po/ycarpa pedunculata C. Mon niot, 1987 may belong to this species. It is not possible to insert in one of these three species the reference of P. nigricans from the Comoro Islands, given without description by Lafargue & Vasseur (1989: 65).

Polycarpa sp. A Fig. 56 Material Examined: MNHN: SI PolB326-lndonesia, north Sulawesi, west of Manado. A specimen measuring 85 x 40 mm, entirely black in formalin, slightly flat tened laterally, was erect in life. The siphons are slightly protruding. The oral aperture is apical, the cloacal aperture 30 mm from it. The relatively soft tunic is wrinkled. Its anterior part is partially covered with epibionts, mostly calcareous encrusting algae. The tunic is somewhat translucent about 2 mm thick. The light brown body wall reveals the gonads in transmitted light. The muscles are colorless in a regular network in the external layer of the body wall. The internal layer is thicker, with large clear vesicles. The total thickness is 1 to 2 mm. We counted 14 large tentacles in 3 orders and numerous small ones inter calated. The prepharyngeal groove has an anterior groove and a posterior crest (Fig. 56B). The deep dorsal V is prolongated by a groove to reach the dorsal lamina. The dorsal tubercle is not protruding and has 2 openings in the present sample. It is circled with non-protruding blood vesicles (Fig. 56B). The dorsal lamina is low. On its right side and posteriorly lies a large unperforated area covered with papillae. The endostyle is only attached to the body wall by a sheet of tissue. It is very long and ends against the esophagus opening. Consequently, Monniot & Monniot: New Ascidians 259

Figure 56. Polycarpa sp. A: A, specimen opened ventrally; B, dorsal area. the branchial sac forms a large posterior bag. The branchial tissue is of a pale ochre. The 4 folds are low and a pseudo-fold is added on the right side. In the posterior part the formula is: R.E. 3 14 5 16 5 17 7 (12 3 4) 0 D.L. 3 JO 6 17 5 17 6 17 4 E.L. Anteriorly, the first sinus on the right is more developed. Posteriorly, it draws near to the dorsal sinuses of the first fold and makes a protruding pseudo-fold. There is no additional sinus there and the space between the pseudo-fold and fold 1 remains constant, so the dorsal sinuses of the first fold move apart in this space. At the same level the first fold lowers. We counted about 20 stigmata between the dorsal lamina and the first sinus on the right side anteriorly, and posteriorly 20 to 30 stigmata. The meshes contain 6 to 8 stigmata between the folds, and 2 to 3 on the folds. Parastigmatic vessels are only present in the case of a division of the rows of stigmata. The gut forms a closed loop, without a secondary curve (Fig. 56A). The stomach is distended, with some grooves seen in transparency. There is no caecum. E.L.

(Sluiter, but 1885),

is necessary to have

16 5

is clear.

P. cryptocarpa

sp. B

P. viridis

4 R. 3

Herdman, from 1882 the southern and eastern

has 4 folds which do has whichfolds 4 not recovereach other. The

Po/ycarpa

Micronesica 29(2), 1996

Figs. 57 A, B, Pl. 9H Figs. 57

makes a clear dorsal prolongated V a in narrow posterior

P. viridis

is alwaysis circonvoluted. There is also one endocarp single

orders, they are long but not curved. The prepharyngeal groove

R.E. 3

P. cryptocarpa

also looks like

Remarks: This specimen seems close to

The 4 cm specimens were attached to the reef by their ventral side. The

The body wall thick is and opaque with 4 distinct layers: the external one is

the gut seems to have a variable but accentuated curve. Kott (1985) said the

of this specimen, with the this of rectumspecimen, continuity in with the distinctive.The intestine is reddish brown tunic is shaded by filamentous sea weeds (Pl. 9H). The quadrate has two equal crests. tissues, lobed testis folliclesand a simple dorsal tubercle, when, at a same size, down to slit dorsalthe lamina The dorsal (Fig. 57B). protruding,tubercle, slightly that of in the gut loop instead of 2 or 3. lighter and slightly nacreous.and lighter hard, Theslightly is consistency mammillated and wrinkled. sac is light-brown is formalin.in sac

thin and contains spherical tan to black granules. The next layer contains brown follicles. The oviduct and follicles. sperm duct open separately by non-protruding papillae. Material Examined: MNHN: SI Pol B 315-Papua New Guinea, Eastern Fields The muscular middle fibers. thick, layer is whitish and contains the gonads; while gonads oval, were but as the gut, they can vary. Our morespecimen is similar to Atoll(CoralSea), 140kmsouthwestofPortMoresby, 145°39.9'E,2 m. 10°00.65'S; the internal layer thin is with granules. About tentacles 30 are irregularly distrib her fig. 28j for the gonads, and to her 98k fig. for the gut. The shape of the gut uted in 4 or differs by a more transparent body wall, the absence of black granules in the color in is life not known while that of 260 extended siphons have 4 lighter patches. The tunic reflex of the siphons is milk additional to specimens the fix status of ours. the chocolatebrown, tuniccolor. When fixed,becomes the in of siphons the inside has multiple openings (Fig. The 57B). height of the dorsal lamina progressively increases down the of middle to esophagustheincreases of the level opening. The branchial does not the overly intestine and covered is with small papillae. The gonads (184 The siphons are protruding. slightly on the right and on 99 the inleft this are specimen) totally embedded in the body Australia. The shape of the dorsal endocarp, tubercle, a single a lobed large testis, formula is: wall thickness, at variable depths. The ovary centralis in a sphere of lobed testis in theclear vesicles body wall and the gut structure are common characters. But

In the cloacal siphon a is circular crest covered with small cloacal tentacles.

The lobed anus is gaping. There is one small endocarp smallone The gaping.There lobed anus is the in is intestinal which loop,

half of the body, with 5 to 7 stigmata in a mesh; afterwards it parts from it with with it from parts it afterwards mesh; a in stigmata 7 to 5 with body, the of half

The first sinus on the right of the dorsal lamina is close to it in the anterior anterior the in it is to close lamina dorsal the of right the on sinus first The

iue 57 Figure

C C

a a

carpa carpa

polycarp

__:

. .

; ;

.. ..

·::.· ·::.·

sp. C: C, specimen specimen C, C: sp.

Polycarpa Polycarpa

-..:::.>

; ;

, ,

transverse transverse

, ,

sp. B: B: sp.

.. ..

Monniot Monniot

( (

.. ..

/· /·

'· '·

.. ..

A, A,

opened ventrally; D, dorsal area; E, internal internal E, area; dorsal D, ventrally; opened

section of a polycarp. polycarp. a of section

_:;::Y _:;::Y

'.a '.a

& &

specimen opened ventrally; B, dorsal dorsal B, ventrally; opened specimen

Monniot: New New Monniot: Ascidians

E E

2 2

mm mm

area. area.

side of of side

Pofy 261 261

The The

P. P. nigri

Heller, Heller, 1878

P. P. nigricans

Polycarpa Polycarpa pedunculata.

(Herdman, (Herdman, and 1906)

sp. sp. C

F, F, Pl. 9B

P. P. pigmentata

The The internal layer is thin and light. About 20

. .

Polycarpa Polycarpa

Micronesica Micronesica 29(2), 1996

Figs. Figs. 57C

which which has a light colored internal tunic in the siphons, a less

Kott, Kott, 1985 has a hard, entirely black tunic, and protruding

P. pigmentata P. pigmentata

The The branchial sac is slightly pigmented, thin and brittle. It has 4 folds. An

The The body wall is opaque black 0.5 to 2 mm thick. The black muscles form

Heller, 1878, sensu Kott, Kott, sensu but Heller, 1878, does not It 1985, belong differs to these species.

This large species (12 x 4 cm; 10 cm really occupied by the body wall) is black black the occupied by is really body cm wall) cm; 4 10 x (12 species This large

P. P. nigricans:

Without Without a complete tract digestive a identification species cannot be given.

This species is closely allied allied closely is to This species

The gut was totally missing in one specimen and and in specimen incomplete one totally the in The other gut was missing (Fig.

teriorly teriorly counted: we

rather short tentacles in 2 or 3 orders of size are more more are tight dorsally rather orders than size or of short 3 2 in tentacles ventrally.

the the external layer. The brown, soft middle layer contains whitish the vesicles di

wall. wall. Its axis has a dark brown color.

unperforated unperforated high, attached The right on is its endostyle area side. lies to the body

does does not all the fill V surface (Fig. 57D). The dorsal lamina is long and low; an

prolongated prolongated by a groove. The dorsal protruding, tubercle, slightly complex and is

ameter ameter of which reaches 0.6 mm

in in thickness, nacreous slightly is interiorly.

kled kled and has epibionts. some It is tough but The remains flexible. tunic, mm 1.5

body body curved with slightly is siphons 4 cm apart. The tunic longitudinally is wrin

The The prepharyngeal groove has only one rounded crest. It forms a large dorsal V

ered ered as a peduncle. This externally looks species like

also also black. The posterior part of the tunic thick very and is can almost consid be

6°50.12'N; 6°50.12'N; m. 25 152°40.85'E,

Chuuk Chuuk State, Mortlock Islands, Losap Atoll, oceanside reef, southwest side,

Material Material Examined: MNHN : SI Pol B 327-Federated States of Micronesia,

except except a thin light brown line around the apertures (Pl. The 9B). siphon lining is

fold fold on the right side.

from from

is always always is smaller. It has a gastric caecum which missing is here.

sinuses between the first right right first the fold and between the dorsal sinuses lamina.

siphons siphons even in contracted zooids. It has fewer sinuses on the folds and more

massive shape, shape, massive a dark body wall with light granules, and an additional branchial

cans cans

region. This species is probably probably is able The to gonads species This eviscerate. region. are totally included

57 A). A). The stomach grooves 57 are clearly through visible the wall. There no is cae

in the the gut in loop. cloacal The tentacles. aperture thick with a by small crest circled is

studied, the gonads are inactive. A large endocarp endocarp located are ones large A and smaller gonads two inactive. are studied, the

inside inside the body The wall. ducts are almost In superficially. the invisible specimen

cum. cum. The gut is attached to the body wall by a large trabeculum in the pyloric

opening opening and do the vessels not produce papillae.

The The a protrusion body shows wall anterior A). to the cloacal siphon (Fig. 57

more more than 20 stigmata in a mesh. The folds lower regularly at the esophagus 262 262

5°15.47'S; 145°49 5°15.47'S;

Material Examined: MNHN: S2 Pyu 350-Papua New Guinea, Madang, Madang, Guinea, New 350-Papua Pyu S2 MNHN: Examined: Material

Synonymy: C. Monniot, 1989: 1989: 490; B. 492 fig.8, pl.11 A, Monniot, C. Synonymy:

nado, 1 1 124°32.41 nado, °23.74'N; 10 m. 'E,

Material Examined: MNHN: S2 Her 6-Indonesia, north Sulawesi, Sulawesi, Ma of west north 6-Indonesia, S2 Her MNHN: Examined: Material

threads joining the branchial sac to the body wall, and from from wall, and body the sac to branchial the joining threads

in the body wall, and particularly from from particularly and wall, body the in

the very peculiar characters of this individual. individual. this of characters peculiar very the

in which testes are branched. branched. are testes which in

multiple openings, a resistant branchial sac with an additional fold on the right right the on fold additional an with sac branchial resistant a openings, multiple

At the same size, size, same the At

and dark brown eggs brown dark and

side, a voluminous and opaque gut, gonads with protruding papillae in buttons buttons in papillae protruding with gonads gut, opaque and voluminous a side,

curve, the other spread between the esophagus and the rectum and extending extending and rectum the and esophagus the between spread other the curve,

allied allied to

a line of small tentacles on its rim. rim. its on tentacles small line of a

along the rectum (Fig. 57C). The cloacal siphon has an internal fleshy fleshy with crest internal an has siphon cloacal 57C). The (Fig. rectum the along

combination of a white ovary deeply sunken into the body wall, as a hollow hollow a as wall, body the into sunken deeply ovary white a of combination

The gonads are embedded in the body wall. wall. diverse body The in the elements embedded are the of polycarps gonads The

protruding. More than 100 gonads are scattered on the whole body wall (Fig. (Fig. wall body whole the on scattered are 100 gonads than More protruding.

gonads make small swellings at the body wall surface, the genital papillae are not not are genital papillae wall the small surface, swellings body the make gonads at

vesicle, circled by a half sphere of pyriform testis follicles (Figs. 57E, F). The The F). 57E, (Figs. follicles testis pyriform of sphere half a by vesicle, circled

57C). There are 2 large endocarps, one occupying all the center of the intestinal intestinal the of center the all occupying one endocarps, large 2 are 57C). There do not touch each other but are embedded in a brown tissue. Each gonad is is gonad Each tissue. brown a in embedded are but other each touch not do

esophagus opening. There are 8 to 10 stigmata in a mesh between the folds and and folds the between mesh a in 10 stigmata to 8 are There opening. esophagus

rectum has a translucent wall; wall; wall; the body to is the anus a it translucent is scalloped. has rectum attached

grooves are not visible visible outside not from are grooves

The space between the first fold and the dorsal lamina on the right increases increases right the on lamina dorsal the and fold first the between space The

close and parallel to the dorsal lamina, an uncommon location for a a for location uncommon an lamina, dorsal the to parallel and close

posteriorly with 11 longitudinal sinuses instead of 8 anteriorly, while on the fold fold the while on anteriorly, 8 of instead sinuses 11 longitudinal with posteriorly

5 to 8 on the folds. There are no parastigmatic vessels. vessels. parastigmatic no are There folds. the on 8 to 5

15 sinuses remain instead of 17. The folds become progressively lower to join the the 17. become progressively folds to lower join of The 15 instead sinuses remain

This very common species is distributed in all warm seas around the world. world. the seas around warm all in species is distributed common very This

This sample differs from other large black black large other differs sample from This

Remarks: This large size species seems to belong to a new species closely closely species new a to belong to seems species size large This Remarks:

The gut is small (Fig. 57C) is small (Fig. gut The

hr sn adtoa fl n h rgt side right the on fold additional no is There

RE. 8 8 RE.

. .

cryptocarpa cryptocarpa

15 15

P. cryptocarpa cryptocarpa P.

12'E, 10 10 m 12'E,

15 15

. .

18 18

Pyura albanyensis albanyensis Pyura

(Sluiter, 1885) 1885) characters. (Sluiter, differs by it numerous from but

Herdmania momus Savigny, 1816 1816 Savigny, momus Herdmania

10 10

Monniot & Monniot & Monniot

21 21

. .

It It

14 14

. .

Family Pyuridae Pyuridae Family

would be necessary to verify on other specimens specimens other verify on necessary to be would

The stomach is slightly enlarged and the internal internal the is slightly and stomach enlarged The

possesses a spongy-looking dorsal tubercle with with tubercle dorsal spongy-looking possesses a

17 17

. .

Fig. 58 58 Fig.

A caecum was not detected. The rather long long rather detected. The caecum was A not

8 8

R. R.

P. obscura obscura P.

Michaelsen, 1927 1927 Michaelsen,

3 3

: :

New Ascidians Ascidians New

16 16

Polycarpa Polycarpa

10 10

. .

2110 15 15 2110

The first sinus on the right is is right the on sinus first The

Heller, 1878 which has white white 1878 has which Heller,

with gonads embedded embedded gonads with

15 15

. .

15 15

ovata ovata

10 10 E.L.

Pizon, 1908 1908 Pizon,

Polycarpa. Polycarpa. 263 263

forms forms irregular curves at the of level

1927 1927

It It

29(2), 1996 1996 29(2),

ica ica

Michaelsen, Michaelsen,

Micrones

Pyura a/banyensis a/banyensis Pyura

. .

is is brown in all its mass.

It It

Figure Figure 58

The The oral tentacles are not numerous (16), without order and they are only

A whitish dense musculature musculature dense whitish A makes the body opaque. wall The external layer

The The specimen 6 cm long was attached on the coral reef by its ventral side.

pharyngeal pharyngeal groove has two equal sheets.

irregularly irregularly digitated. They are planted above a strong muscular ring. The pre

has has transverse muscular bundles joined on the dorsal and lateral sides, more

hidden hidden in the bottom of a tunic depression. The tunic is 4 to 5 mm thick, tough

the the retracted animal, the oral siphon cannot be seen and the cloaca! siphon is

has has been torn on each when side in fixed formalin (Fig. 58).

gitudinal gitudinal bundles, regularly spaced. The are muscles so strong that the body wall

spaced spaced on the ventral side. The internal layer has close, but separated, well lon

and and rigid.

uniformly uniformly purple with only large iridescent on spinules the rim of the siphons. In

have have 8 ochre bars separated by black bars. In formalin, the internal tunic turns

substrate substrate cover. Between the epibionts, the tunic is reddish. The short siphons

The The tunic is for the most part coated with small algae in continuity with the 264 264 t oeig n w sias unn i te aewy h sot osl aia is lamina dorsal short The way. same the in turning spirals two in opening its made of sharp languets of medium size, placed on the left side of an unperforatedleft side the an of size, of on medium placed languets sharp of made

counted on the right side: right the on counted

ia esl r rlnae b aile Te rnha mse r qae with square are meshes branchial The papillae. by prolongated vessels are dinal

h rnha flsad epdra V h osl uecei rtuig with protruding is tubercle dorsal The V. dorsal deep a and folds branchial the culated mass in the shape of a cauliflower (Fig. 58). The cylindrical intestine ends 58). cylindricalintestine cauliflower a (Fig. ofThe shape the in mass culated ad Te hn rnha schs ih od ta my oe ec ohr We other. each cover may that folds high 6 has sac branchial thin The band.

t 8simt bten h fls ad t 6 tgaa n h fls te are they folds; the on stigmata 6 to 4 and folds, the between stigmata 8 to 6 eual rse ylreprsimtc vessels. parastigmatic large by crossed regularly

Palau. pedun large a of and cavity) side cloacal right the in ( located lobes them of one nasot etmwt nau u naltrl slit. lateral a in cut anus an with rectum short a in

regular diverticula in the apex of the right side gonad. The left gonad occupies all occupies left gonad The side gonad. right the of apex in the diverticula regular the space between the two legs of the gut (Fig. 58). The sperm ducts were not seen were58). ducts not sperm (Fig.legstwo between the space The the the gut of on the gonad. The terminal gonoducts are almost as wide as the rectum and open and rectum as the wide as almost are gonoducts terminal The gonad. the on ylreppla coet teau. h noap cvrtegnd oe n a and lobes gonad the cover endocarps The anus. the to close papillae large by ag at rud h dgsietat O te ih sd,te ae ie o both on lined are they side, right the On tract. digestive the around part large ie faln n ouios heart. voluminous and long a sides of

ventrally. When fixed, the color turns uniformly dark purple in its purpleWhen ventrally. fixed, whole dark uniformly the thickness, turnscolor

ircsu manaarensis Microcosmus embedded in the internal ventral half of the external tunic. The internal tunic was tunic internal The tunic. external the of half ventral internal the in embedded aeil Examined: Material sad eftp l°56'; 123°59.11 IO l0°15.62'N; m. 'E, top, reef Island,

network of brown pigment granules, sometimes black. The internal siphons have siphons internal The sometimesblack. granules, pigment brown of network and cm 2 about in protrude siphons divergent The high. cm 3 and long cm 5 is their apertures are l cm in diameter. In life, the tunic is purple dorsally and yellow life, In is and dorsally tuniccm in the diameter. purple l are apertures their

reaching 5 mm near the siphons. The epibionts are mostly hydrozoans. mostly are epibionts The siphons. the near 5 mm reaching

tunic, with siphons which were not linked to the external siphons. Sediment was Sediment siphons. external the to linked not were which siphons with tunic,

nylne o h xenloeb ml at ftevnrl ieweeae the are side where ventral the of part small a by one external the to linked only ui esl.Teitra tnc Im hc,i ih rw wt e ihn. Its siphons. red with brown light is thick, Imm tunic, internal vessels. The tunic surface is reticulated with small spines. The reticulated aspect is accentuated by a by is accentuated aspect reticulated spines. small The with is reticulatedsurface

large acicular spinules largely pointing exteriorly. We did not observe spinules on observe spinules not did We exteriorly. spinules largely pointing acicular large h etra spos Te nlso o sdmn isd te nenl ae o the of layer internal the inside sediment of inclusion The siphons. external the

95 6 l ,fg -,Mda;Mla, 1975: 1915: Millar, 16, fig. 309, Madras; l, 6-7, pl. fig. 87, Sea Singapore. and Java

h flsae etyitrutd t h spau oeig n te longitu the and opening esophagus the at interrupted neatly are folds The

The stomach is not enlarged. The hepatic gland is made of smallis 4 finger-like made gland hepatic The enlarged. is not stomach The

hr i n oa o ah iemd o eta tb ad aea rather lateral and tube central a of made side each on gonad one is There

h nml a togy tahd o h ok yisvnrl ie h body The side. ventral its by rock the to attached strongly was animal The Sea, Arafua Caledonia, New Australia, Around Distribution: Geographic

nctigtetncw aebe upie ofn niei nte complete find inside another it to we been tunicsurprised have the cutting In

ircsu manaarensis Microcosmus

. 15 E.

MNHN:

ono &Mnit Nw Ascidians New Monniot: & Monniot

eda, 96 1 l ,fg 33, r ak; Oka, Lanka; Sri fig. 1906:23-31, 11, 2, pl; Herdman,

S2 Mic 131-Philippines, Cebu, south east Mactan east south Cebu, 131-Philippines, S2 Mic

Fig. 59 Fig.

eda, 1906 ? Herdman,

6 R. 6

265 22

1996

Herdman, A, 1906: muscles; B, dorsal area;

Micronesica 29(2),

ends at the esophagus opening but does not join

Microcosmus manaarensis Microcosmus

59.

C, specimen opened ventrally; D detail of the liver.

is undulated is around the top of the branchial folds and cut in a dorsal

is not is possibleto know if this phenomenon is normal or an artifact.

Figure

The musculature is very strong, with a strong retractor muscle in the oral

266

animal.

siphon (Fig. The 59A). siphonshave only radial Strong fibers. ribbons circular of

to make a muscular scarf several mm thick, protruding. The radial fibers of the

external tunic proves that both tunics were separated before the collection of the tentacles are arranged in 3 or 4 orders; they are more developed dorsally. The largest measure 5 to 6 mm and are slightly branched with some buttons on the

fibers cover fibers the radial fibers at the base of the oral siphon. The circular muscles starting at the base of the cloacal siphon join the radial fibers of the oral siphon

made of a ring of non muscular body wall, resistant and divided in two lobes

primary branches (Fig. The 59B). prepharyngeal area is covered with papillae.

veloped. brane progressively higher. cloacal siphon run under this scarf and reappear on the ventral of side the body. The musculature thickand is continuous on body above the the exceptgut.whole The siphonalare linings red, doubled by a almost thelarge velum siphonsclosing

The prepharyngeal groove has 2 separated well crests, the anterior one more de V. The V. dorsal tubercule protrudes (Fig. The 57B). dorsal lamina a is plain mem

bilobate diaphragm closing the siphons. siphons. the closing diaphragm bilobate

near Madras, specimens entirely covered with sediment: "crust of sand covering covering sand of "crust sediment: with covered entirely specimens Madras, near

not seem related to the openings of the external tunic. Oka (1915) has recorded, recorded, has (1915) Oka tunic. external the of openings the to related seem not

In the western Atlantic and off Senegal where the species is recorded (Monniot & & (Monniot recorded is species the where Senegal off and Atlantic western the In

uis s a t gv h apaac o oe et yn wti aohr (. 312) (p. another" within lying test one of appearance the give to as "so tunics

the species as entirely covered by a thick layer of sediment, having two superposed superposed two having sediment, of layer thick a by covered entirely as species the

pillae. There are about 8 long stigmata in a mesh between the folds and 3 to 5 on on 5 3 to and folds the between mesh a in stigmata long 8 about are There pillae.

the gonads. gonads. the both conditions occur at the same location, and some individuals even have a a have even individuals some and location, same the at occur conditions both

sesses a ring of small tentacles. tentacles. small of ring a sesses ih h rtohrnel roe Te ogtdnl esl r hg, ihu pa without high, are vessels longitudinal The groove. retropharyngeal the with the tunic structure and the shape of the lappet closing the siphons. siphons. the closing lappet the of shape the and structure tunic the

from the Banda Sea) for the close similarities of the branchial sac, the gut, and and gut, the sac, branchial the of similarities close the for Sea) Banda the from of sand, appears to be a response to the habitat of the individual. Specimens with with Specimens individual. the of habitat the to response a be to appears sand, of

sltd atro, t h tp fte u lo (i. 9) Te laa spo pos siphon cloacal The 59C). (Fig. loop gut the of top the at anterior, isolated, (synonymy suggested by Van Name, 1921 as that of of that as 1921 Name, Van by suggested (synonymy

are short but distinct. On the left side, endocarps are linked to the gut and one is is one and gut the to linked are endocarps side, left the On distinct. but short are

One year later, Hartmeyer (1919) synonymized the two species species two the synonymized (1919) Hartmeyer later, year One

inactive. Two lobes are on the right side and 3 lobes on the left side. The gonoducts gonoducts side. The left the on 3 lobes and side right the on are lobes Two inactive. scriptions mention a strong musculature. Herdman figured (pl. 2, fig. 27) a strong strong a 27) fig. 2, (pl. figured Herdman musculature. strong a mention scriptions

both species closely allied because of the similarity of the rest of their anatomy. anatomy. their of rest the of similarity the of because allied closely species both

right side: side: right

siphons. siphons.

truding in the descending part. The anus has 2 plain lobes. The gonads were were gonads The lobes. plain 2 has anus The part. descending the in truding h ts" p 1) ilr 17) lo ecie a obe ui (i.8d. l de All 87d). (fig. tunic double a described also (1975) Millar 16). (p. test" the

described under the name name the under described

any fold. The branchial tissue has 6 high overlapping folds. We counted on the the on counted We folds. overlapping high 6 has tissue branchial The fold. any

oral siphon at the tentacle level, and the absence of spinules or scales inside the the inside scales or spinules of absence the and level, tentacle the at siphon oral the name name the netn i hn altl icue i h bd wl nis sedn lm, n pro and limb, ascending its in wall body the in included little a thin, is intestine

the species is characterized by the presence of 4 prolongations which close the the close which prolongations 4 of presence the by characterized is species the and varied debris with a weak musculature, described by Herdman (1882) under under (1882) Herdman by described musculature, weak a with debris varied and glandular ridges, more or less parallel, with small papillae (Figs. 59C, D). The The D). 59C, (Figs. papillae small with parallel, less or more ridges, glandular

Monniot 1994) the only form covered with sediment is known. After Kott (1985) (1985) Kott After known. is sediment with covered form only 1994) the Monniot

presents two very different aspects: specimens with a thin tunic including sediment sediment including tunic thin a with specimens aspects: different very two presents His figure 26 represents these two layers and the siphons of the internal tunic do do tunic internal the of siphons the and layers two these represents 26 figure His

the folds. Parastigmatic vessels are present. present. are vessels Parastigmatic folds. the

cosmus cosmus

hc sn ahrn t at o h ts, hl lehr te r nkd (. 350) (p. naked" are they elsewhere while test, the of parts to adhering sand thick

The gut forms a closed flat loop. The hepatic gland is voluminous, made of of made voluminous, is gland hepatic The loop. flat closed a forms gut The

Posteriorly the folds are lower and the vessels join in a single crest and fuse fuse and crest single a in vessels join the and lower are folds the Posteriorly

It It

Kott (1985) synonymized synonymized (1985) Kott

Remarks: Kott (1985) in the Indo-Pacific region only recognizes one one recognizes only region Indo-Pacific the in (1985) Kott Remarks:

Kott (1985) said: "the condition of the surface test, with or without its coat coat its without or with test, surface the of condition "the said: (1985) Kott

is certain that that certain is

pce wt 6 rnha folds branchial 6 with species

M. helleri helleri M.

M. manaarensis manaarensis M.

and specimens with a thick hard tunic and strong musculature musculature strong and tunic hard thick a with specimens and

M. helleri helleri M.

E. I I E.

M. goanus goanus M.

Monniot Monniot

23 23

a ecie tre times three described was

4 4

24 24

and and

M. helleri helleri M.

& &

4 4

Monniot: New Ascidians Ascidians New Monniot:

M. manaarensis manaarensis M.

21424 21424

by Michaelsen (1918). This author estimated estimated author This (1918). Michaelsen by

: :

M. helleri helleri M.

and and

4 4

25 25

3 3

. .

Herdman, 1882. This species species This 1882. Herdman,

20 20

manaarensis manaarensis

are closely allied, distinct by by distinct allied, closely are

. .

Herdman (1906) described described (1906) Herdman

3 R. R. 3

. .

g/eba g/eba

Traustedt, 1885 1885 Traustedt,

Herdman, 1906 1906 Herdman,

It It

is possible possible is

Micro

267 267

. . . . 268 Micronesica 29(2), 1996 that the two species developed two ecological forms, one living in contact with the sediment with the tunic including varied particles, the other living in rocky areas, with a hard and naked tunic. In this hypothesis, the double tunic of the Philippines sample would not represent a monster, but a particular anatomical character. This specimen possesses large spinules absent in M. helleri. Geographic Distribution: India, Sri Lanka, Singapore, and Java Sea.

Acknowledgments Funding for a part of the collections for this work was from the Coral Reef Research Foundation through a contact with the U.S. National Cancer Institute, National Institutes of Health. Many of the color photographs were provided by the Coral Reef Research Foundation. We are much indebted to numerous mu seums for lending types for examination in our study.

References Berrill, N. J. & D. P. Abbott. 1949. The morphology and larva of the ascidian Pycnoclavella stanleyi n. sp. Canadian Journal of Research (D) 27: 43--49. Brewin, B. I. 1957. Ascidians of New Zealand. Part X. Ascidians from the North Auckland. Transactions of the Royal Society of New Zealand 84: 577- 580. Brien, P. 1970. Regeneration du thorax par un segment abdominal chez Clavelina lepadiformis Muller. Archives de Biologie 81: 441--490. Brien, P. & E. Brien-Gavage. 1927. Contribution a l'etude de la blastogenese des tuniciers. Ill. Bourgeonnement chez Clavelina /epadiformis Muller. Recueil de l'Institut Zoologique Torley Rousseau 1: 31-81. Brunetti, R. 1987. Species of Clavelina in the Mediterranean Sea. Annales de l'Institut Oceanographique (n. s.) 63: 101-118. Cloney, R. A. 1977. Larval adhesive organs and metamorphosis in ascidians. Cell Tissue Res. 183: 423--444. Cloney, R. A. 1978. Ascidian metamorphosis: review and analysis. In F. S. Chia & M. E. Rice (eds), Settlement and Metamorphosis of Marine Invertebrate Larvae, pp. 255-282. Elsevier/North-Holland Biomedical Press, Amsterdam. Cloney, R. A. 1990. 14. Urochordata-Ascidiacea. In K. G. Adiyodi & R. G. Adiyodi (eds), Reproductive Biology of Invertebrates. 4 (B), pp. 391--451. Oxford & IBH Publishing Co. New Delhi, Bombay, Calcutta. Garstang, W. 1891. Report on the tunicata of Plymouth. Part. 1. Clavelinidae, Perophoridae, Diazonidae. Journal of the Marine Biological Association, U. K. 2: 47-67. Garstang, W. 1895. Outlines of a new clasification of the Tunicata. Reports of the British Association for the Advancement of Science 1895: 718-719. Goodbody, I. 1984. The ascidian fauna of two contrasting lagoons in the Neth erland Antilles: Pescadera Baai, Cura~ao, and the lac of Bonaire. Studies on the Fauna of Cura~ao and other Caribbean Islands 48 (202): 21-61. Gottschaldt, R. 1898. Synascidien von Ternate. Abhandlungen herausgegeben von der Senckenbergischen naturforschen Gesellschaft 24: 641-660.

ot P 15b h esl uiaa Sinii eot o h on ury Ex Murray John the of Scientific Reports sessile 1957b. Tunicata. The P. Kott,

Kott, P. 1954. Tunicata. Reports of B.A.N.Z. Antarctic Research Expedition Expedition Research Antarctic B.A.N.Z. of Reports 1954. Tunicata. P. Kott,

Kott, P. 1957a. The ascidians of Australia. II. Aplousobranchiata Lahille: Clav Lahille: Aplousobranchiata II. Australia. of ascidians 1957a. The P. Kott,

Herdman, W. A. 1906. Report on the Tunicata. Ceylon Pearl Oyster Fisheries Fisheries Oyster Pearl Ceylon Tunicata. the on 1906. Report A. W. Herdman,

Herdman, Herdman,

Herdman, W. A. 1891 b. On the genus genus 1891 the On A. b. W. Herdman,

Herdman, W. A. 1891a. A revised classification of the Tunicata, with definitions definitions with Tunicata, the of revised 1891a. classification A A. W. Herdman,

eda, . . 1886 A. W. Herdman,

Herdman, Herdman,

Heller, C. 1878. Beitrage zur naheren Kenntnis der Tunicaten. Sitzungsberichte Sitzungsberichte Tunicaten. der Kenntnis naheren zur 1878. Beitrage C. Heller,

Hastings, A. B. 1931. Tunica ta. Great Barrier Reef Expedition 1928-29. 1928-29. Scientific 1931. Reef Expedition B. A. Barrier Hastings, Tunica Great ta.

Hartmeyer, R. R. Hartmeyer,

Hartmeyer, R. 1909-11. Tunicata (Manteltiere). (Manteltiere). Tunicata 1909-11. R. Hartmeyer, atee, . 91 iclae Acdooia Mtelne as e zoolo dem aus Mitteilungen Ascidiologica. Miscellanea 1921. R. Hartmeyer,

Hartmeyer, R. 1919. 1919. Ascidien. R. Hartmeyer,

Gould, A. 1852. Mollusca and shells. shells. and Mollusca 1852. A. Gould,

pedition 1933-34. 10 (4): 129-150. 10 129-150. (4): 1933-34. pedition

elinidae Forbes Forbes elinidae

Marine and Freshwater Research 8: 64-110. 8: 64-110. Research Freshwater and Marine

Proceedings and Transactions of the Liverpool Biological Society 5 Society Biological Liverpool the of Transactions and Proceedings Supplementary Report 39: 39: 295-348. Report Supplementary

scriptions of two new species, and a classification of the family Clavelinidae. Clavelinidae. family the of classification a new species, two and of scriptions

Museum. Australian Museum, Sydney, Catalogue 17: 17: 1-139. Catalogue Sydney, Museum, Australian Museum.

keys to the species. Journal of the Linnean Society of London, Zoolology 23: 23: Zoolology Society London, of Linnean the of species. the keys to Journal

of the orders, sub-orders, families, sub-families and genera, and analytical analytical and genera, and sub-families families, sub-orders, orders, the of

Report of the Scientific Results of the Voyage of of Voyage the of Scientific Results the of Report

the Years 1873-76. 14 14 1-399. (38): 1873-76. Years the

558-562. 558-562.

1929-1931. 1 1 1929-1931. (4) H.M

der Wissenschaft Wien, 77 (1): 83 (1): 77 Wien, Wissenschaft der

der mathematischnaturwissenschaftlichen Classe der kaiserlichen Akademie Akademie kaiserlichen der Classe mathematischnaturwissenschaftlichen der the Years 1873-76. 6 (17): 1-296. 1-296. (17): 6 1873-76. Years the

Report of the Scientific Results of the Voyage of of Voyage the of Scientific Results the of Report

Reports 4 (3): (3): 69-109. 4 Reports

163 miens Aarsberetning 60 (4): 1-150. 1-150. (4): 60 Aarsberetning miens

ranchiae. Fauna Siidwest-Australiens 5: 251-460. 5: 251-460. Siidwest-Australiens Fauna ranchiae.

gischen Museum Berlin 10: 10: 299-323. Berlin Museum gischen

Wilkes. 12 12 (4): Wilkes. 495-498.

xeiin t Asrla 901. ogia vnk Vetenskaps-Akade Svenska Kongliga 1910-13. Australia to expeditions

nungen des Tier-Reichs. Leipzig Tier-Reichs. des nungen

during the years 1838, 1838, the years 1839, during 1840, 1841, of Charles 1842, the command under

M.S. M.S.

. .

W. A. A. W.

. .

'Challenger' during the years 1873-76. Part. II. Ascidiae Ascidiae compositae. II. 1873-76. Part. years the during 'Challenger'

Calne' uig h yas 1873 years the during 'Challenger'

& &

.Mcale. 97 siieDkybacie n Ptychob und 1927. Diktyobranchiae Ascidiae Michaelsen. W.

1899. Descriptive catalogue of the Tunicata in the Australian Australian the in Tunicata the of catalogue 1899. Descriptive

1882. Report on the Tunicata collected during the voyage of of voyage the during collected Tunicata the on 1882. Report

& &

: :

121-182. 121-182.

. .

Hanley and Polyclinidae Verrill. Australian Journal of of Journal Australian Verrill. Polyclinidae and Hanley

Report on the Tunicata collected during the voyage of of voyage the during collected Tunicata the on Report

Monniot Monniot

In In

& &

Results of Dr of Results

Monniot: New Ascidians Ascidians New Monniot:

. .

spl) 1281 (suppl.): 3

In In

Ecteinascidia Ecteinascidia

109. 109.

United States Exploring Expedition, Expedition, Exploring States United

. .

E. Mjoberg's Swedish scientific scientific Swedish Mjoberg's E.

In In

. .

Part. I. Ascidiae simplices. simplices. Ascidiae I. Part.

rn' Kasn n Ord und Klassen Bronn's

n is eain; ih de with relations; its and

H.M.S. H.M.S.

1773. 1773.

Challenger during during Challenger

: :

144- 269 269

& &

(7): (7):

5 5

B. B. S. Morton

du du N. E. du canal

In In

coralliens coralliens

13 (23): (23): 13 1-18.

recifs recifs

Zoologi Zoologi

for for

Micronesica Micronesica 29(2), 1996

19. 19.

59-66. 59-66.

49: 49:

Vasseur. Vasseur. Ascidies des 1989.

P. P.

Goodbody. Goodbody. The 1982. ascidians of Hong Kong.

Mesogee Mesogee

I. I.

& &

1992a. The The 1992a. Australian Ascidiacea part Aplousobranchia 3, Memoirs (2).

P. P.

463-558. 463-558.

1977). 1977).

Zoologischer Zoologischer Anzeiger 71: 193-203.

stalten stalten 37: 1-74.

Ozeans: Ozeans: Didemniden. Jahrbuch der hamburgischen wissenschaftlichen An

museum museum zu Stockholm. Arkiv

tlichen tlichen Anstalten 35(2): 1-71.

de de Mozambique (Campagne BENTHEDI du 'Suroit', 17 mars-14 avril

westlichen westlichen Indischen Ozeans. Jahrbuch der hamburgischen wissenschaf

C. C. K. Tseng The (eds). Marine Flora and Fauna of Hong Kong and Southern

China, China, pp. 503-554. Hong Kong University Press.

of of the Queensland Museum 32: 375-620.

Queensland Queensland Museum 32: 621-655.

supplement. supplement. Memoirs of the Queensland Museum 29: 267-298.

of of the Queensland Museum 29: 1-266.

ranchia. ranchia. Memoirs of the Queensland Museum 23: 1-440.

notes notes on the symbiosis. Micronesica 85-127. 18:

waters. waters. Beagle (2): 1 13

Society Society of New South Wales 147-212. 105:

oirs oirs of the Queensland Museum 20: 1-47.

of of South Australia 96: 165-196.

ings ings of the Linnean Society of New South Wales 258-261. 97:

Australian Australian Bight and Investigator Strait. Transactions of the Royal Society

coast. coast. University of Queensland Papers, Department of Zoology 2: 127-152.

Research Research 70-118. 14:

clinidae clinidae Verrill (continued). Australian Journal of Marine and Freshwater

Michaelsen, Michaelsen, Ascidiae W. Krikobranchiae. 1930. Fauna Sudwest-Australiens

Michaelsen, Michaelsen, W. 1927. Einige neue westaustralische ptychobranchiate Ascidien.

Michaelsen, Michaelsen, W. Ascidien vom westlichen 1921. Indischen Ozeans aus dem Reichs

Michaelsen, Michaelsen, W. 1920. Die Krikobranchien Ascidien des westlichen Indischen

Michaelsen, Michaelsen, W. 1918. Die Ptychobranchien und Diktyobranchien Ascidien des

Lafargue, Lafargue, F. &

Kott, Kott,

Kott, Kott, P. 1992b. The Australian Ascidiacea, Supplement 2. Memoirs of the

Kott, Kott, P. The 1990b. Australian Ascidiacea, Phlebobranchia and Stolidobranchia,

Kott, Kott,

Kott, Kott, P. The 1990a. Australian Ascidiacea part Aplousobranchia 2, Memoirs (1).

Kott, Kott, P. Two 1983. new genera of didemnid ascidians from tropical Australian

Kott, Kott, P. The 1985. Australian Ascidiacea part Phlebobranchia 1, and Stolidob

Kott, Kott, P. The 1981. ascidians of the reef flats of Fiji. Proceedings of the Linnean

Kott, Kott, P. Didemnid-algal 1982. symbioses: host in species the western with Pacific

Kott, Kott, P. Algal-bearing 1980. didemnid ascidians in the Indo-West Pacific. Mem

Kott, Kott, P. The 1972. ascidians of South Australia. II. Eastern sector of the Great

Kott, Kott, P. Plurellidae 1973. a new phleboranchiate family of Ascidiacea. Proceed

Kott, Kott, P. 1964. Stolidobranch and phlebobranch ascidians of the Queensland

Kott, Kott, P. The 1963. ascidians of Australia. IV. Aplousobranchiata Lahille: Poly 270 270

Monniot, C. C. Monniot,

Monniot, C. 1991b. Ascidies de Nouvelle-Caledonie. X. Stolidobranches (suite). (suite). Stolidobranches X. Nouvelle-Caledonie. 1991b. de Ascidies C. Monniot,

Monniot, Monniot,

Monniot, C. 1989. Ascidies de Nouvelle-Caledonie. VI. Pyuridae et Molgulidae. Molgulidae. et Pyuridae VI. 1989. Nouvelle-Caledonie. de Ascidies C. Monniot,

Monniot, C. 1988. Ascidies de Nouvelle-Caledonie. IV. Styelidae (suite). Bulletin Bulletin (suite). Styelidae IV. 1988. Ascidies Nouvelle-Caledonie. de C. Monniot,

Monniot, C. 1987b. Ascidies de Nouvelle-Caledonie. II. Les genres genres Les II. Nouvelle-Caledonie. 1987b. de Ascidies C. Monniot,

Monniot, C. 1987a. Ascidies de Nouvelle-Caledonie. I. Phlebobranches du lagon du Phlebobranches I. 1987a. Ascidies de Nouvelle-Caledonie. C. Monniot,

Monniot, C. 1983. Ascidies littorales de Guadeloupe. II. Phlebobranches. Bulletin Bulletin Phlebobranches. II. 1983. C. de Guadeloupe. Ascidies littorales Monniot,

Monniot, C. 1972. Ascidies stolidobranches des Bermudes. Bulletin du Museum Museum du Bulletin Bermudes. des 1972. stolidobranches Ascidies C. Monniot,

Millar, Millar,

Millar, R.H. 1967. 1967. genus of the A new ascidian R.H. Millar,

Millar, Millar,

93. 93.

Atlantic; arrival of cosmopolitan species. Bulletin of Marine Science Science 71- 54: Marine of species. Bulletin cosmopolitan of arrival Atlantic;

Nouvelle-Caledonie. Diversite des strategies adaptatives. In A. Crosnier (ed), (ed), Crosnier A. In adaptatives. des strategies Diversite Nouvelle-Caledonie.

National d'Histoire naturelle, Paris Paris naturelle, d'Histoire National

cidies apropos d'especes nouvelles en Mediterranee. Ann. Inst. Inst. Ann. Mediterranee. en nouvelles d'especes apropos cidies

d'Histoire naturelle, Paris Paris naturelle, d'Histoire

Resultats des campagnes MUSORSTOM. 8. Memoires du Museum National National Museum 8. du Memoires MUSORSTOM. des campagnes Resultats

e) Rslas e cmans UOSO. 4 MUSORSTOM. campagnes des Resultats (ed),

63 63

du du

Bulletin du Museum National d'Histoire naturelle, Paris (4), 12A 12A (4), Paris (3--4): naturelle, 491- d'Histoire National Museum du Bulletin

du Museum National d'Histoire naturelle, Paris (4), l0A (2): 163 (2): l0A (4), Paris naturelle, d'Histoire National Museum du

Bulletin du Museum National d'Histoire naturelle, Paris (4), 13 (1-2): 3-37. 3-37. 13 (1-2): (4), Paris naturelle, d'Histoire National Museum du Bulletin

Polyandrocarpa. Polyandrocarpa.

du Museum National d'Histoire naturelle, Paris (4), 5A (1): 51-71. 51-71. (1): 5A (4), Paris naturelle, d'Histoire National Museum du

507. 507.

Bulletin du Museum National d'Histoire naturelle, Paris (4), 1 la (3): 475- (3): 1 (4), la Paris naturelle, d'Histoire National Museum du Bulletin

National d'Histoire naturelle, Paris (3), 57 (Zool. 43): 617-643. 617-643. 43): (Zool. 57 (3), Paris naturelle, d'Histoire National

9A (2): (2): 275-310. 9A

515. 515.

Proceedings of the Zoological Society of London 141 141 (4): 689-746. London of Society Zoological the of Proceedings

Expedition. Journal of Natural History 22: 823-848. 823-848. 22: History Natural of Journal Expedition.

Museum, Copenhagen (Tunicata, Ascidiacea). Steenstrupia 3: 205 3: Steenstrupia Ascidiacea). (Tunicata, Copenhagen Museum,

Journal of Natural History 1: 1: 359-361. History Natural of Journal

Bulletin du Museum National d'Histoire naturelle, Paris (4), 9A 9A (4), Paris naturelle, d'Histoire National Museum du Bulletin

azine of Natural History (12) 9: 913-932. 913-932. (12) 9: History Natural of azine

of the Zoological Society of London 123: 123: 277-325. London of Society Zoological the of

R.H. R.H.

(2): 119-130. 119-130. (2):

R.H. R.H.

R. H. H. R.

Museum National d'Histoire naturelle, Paris Paris naturelle, d'Histoire National Museum

C. C.

& &

91.Acde eNuel-aeoi.VI.Plbbace (suite) 1991a. Phlebobranches Ascidies VIII. de Nouvelle-Caledonie.

& &

1963. Australian ascidians in the British Museum (Natural History). History). (Natural Museum British the in ascidians 1963. Australian

1953. On a collection of ascidians from the Gold Coast. Proceedings Proceedings Coast. Gold the 1953. from ascidians of collection a On

1975. Ascidians from the Indo-west-Pacific region in the Zoological Zoological the in region Indo-west-Pacific the 1975. from Ascidians

96 siin rm oabqe Es fia nas n Mag and Annals Africa. East Mozambique, 1956. from Ascidians

1988. Ascidians collected during the International Indian Ocean Ocean Indian International the during collected 1988. Ascidians

F. Monniot. 1987a. 1987a. Les ascidies Polynesie de Monniot. F.

F. Monniot. 1991. Tunicata: peuplements d'ascidies profondes en en profondes d'ascidies peuplements 1991. Tunicata: Monniot. F.

F. Monniot. 1987b. Discussion sur la valeur de deux genres d'as genres deux de valeur la sur 1987b. Discussion Monniot. F.

F. Monniot. 1994. Additions to the inventory of eastern tropical tropical eastern of inventory the to 1994. Additions Monniot. F.

F. Monniot. 1989. 1989. Ascidies Monniot. F.

Bulletin du Museum National d'Histoire naturelle, Paris (4), (4), Paris naturelle, d'Histoire National Museum Bulletin du

Monniot & Monniot: New Ascidians Ascidians New Monniot: & Monniot

(A), (A),

151: 151: 357--448.

(A), (A),

(MUSORSTOM (MUSORSTOM

143: 143: 229-245.

Pseudodistoma Pseudodistoma

(A), (A),

. .

Memoires du Museum Museum du Memoires

136: 136: 1

fran~aise. fran~aise.

1 1

from Madagascar. Madagascar. from

& &

155. 155.

2). In In 2).

Polycarpa Polycarpa

(1): (1):

Oceanogr. Oceanogr.

Memoires Memoires

196. 196.

J. J.

336. 336.

Forest Forest

31. 31.

271 271

et et

. . . .

. .

et et

Bul

Atriolum Atriolum

Lissoc/inum Lissoc/inum

XXXIII. XXXIII.

Trididemnum

. .

Didemnum. Didemnum.

Publications Publications of the Seto

I. I.

589. 589.

691. 691.

avec avec le milieu naturel et d'autres ports. Bulletin

Micronesica Micronesica 29(2), 1996

P. P. Laboute. Ascidies du 1985. port de Papeete (Po

& &

relations relations

140. 140.

Collection Collection Faunes Tropicales. 20. Editions de l'ORSTOM,

dans dans le lagon sud. Bulletin du Museum National d'Histoire

T. T. Tokioka. Contribution 1976. to the Japanese ascidian fauna.

Ascidiacea). Ascidiacea). Proceedings of the Japanese Society of Systematic

In In

& &

1990. Ascidies de de Ascidies Nouvelle-Caledonie. 1990. VIII. Le genre

1983. Ascidies littorales littorales Ascidies 1983. de Guadeloupe. I. Didemnidae. Bulletin du

Note Note on some clavelinids from the Japanese waters. Publications of

frarn;aise): frarn;aise):

. .

Perophoridae, Perophoridae, Ascidiidae and Corellidae. Proceedings of the Japanese So

tions tions of the Zoological of Society New Zealand 24: 305-334.

the the Seto Marine Biological Laboratory 23: 63-82.

111. 111.

Marine Marine Biological Laboratory 25-170. 35:

Marine Marine Biological Laboratory 73-148. 34:

XXIX. XXIX.

(Tunicata

ciety ciety of Systematic Zoology 32: 30-78.

I. I.

Zoology Zoology 27: 107

two two new species. Memoirs of the National Museum, Science Tokyo. 97- 13:

cidians cidians from the coast of Kii Peninsula, Middle Japan, with descriptions of

letin du du letin Museum National d'Histoire naturelle, (3-4): Paris 299-344. (4) 16A

(Didemnidae) (Didemnidae) dans le lagon sud. Bulletin du Museum National d'Histoire

529. 529.

Bulletin du du Bulletin Museum National d'Histoire naturelle, Paris (3-4): (4), 517- 12A

naturelle, naturelle, Paris (4) (3-4): 14A 565

naturelle, naturelle, Paris (4), IA 673 I (4):

Leptoc/inides Leptoc/inides 535. 535.

Bulletin Bulletin du Museum National d'Histoire naturelle, Paris (4), l0A (2): 197-

Bulletin Bulletin du Museum National d'Histoire naturelle, Paris (4), 9A (3): 499-

235. 235.

Museum Museum National d'Histoire naturelle, Paris (4), 5A (1): 5-49.

Museum Museum National d'Histoire naturelle, Paris 41 426-457. (2), (2):

Caledonia. Caledonia.

du du Museum National d'Histoire naturelle, Paris (4), 7A 481-495. (3):

lynesie lynesie

Paris. Paris. pp. 247

Nott, Nott, J. T. On 1892. the composite ascidians of the North Shore Reef. Transac

Nishikawa, Nishikawa, T.

Nishikawa, Nishikawa, T. The 1991. ascidians of the Japan Sea. II. Publications of the Seto

Nishikawa, Nishikawa, T. The 1990. ascidians of the Japan Sea.

Nishikawa, Nishikawa, T. Ascidians from the 1986. Gilbert and Solomon Islands and Nauru.

Nishikawa, Nishikawa, Ascidians Truk from T. Island, 1984. Ponape Island and Majuro Atoll

Monniot, Monniot, Nouvelle-Caledonie. de F. Ascidies XV genre Le 1995.

Monniot, Monniot, F. 1992. Ascidies de Nouvelle-Caledonie. XII. Le genre Nishikawa, Nishikawa, T. 1980. Contribution to the Japanese ascidian fauna

Monniot, Monniot, F. Ascidies de 1989. Nouvelle-Caledonie. VII. Les genres

Monniot, Monniot, F

Monniot, Monniot, F. 1988. Ascidies de Nouvelle-Caledonie. V. Polycitoridae du lagon.

Monniot, Monniot, F. 1987. Ascidies de Nouvelle-Caledonie. III. Polyclinidae du lagon.

Monniot, Monniot, F

Monniot, Monniot, F. Sur une 1969. collection d'ascidies composees de Dakar. Bulletin du

Monniot, Monniot, C., F. Monniot & P. Laboute. 1991. Coral Reef Ascidians of New

Monniot, Monniot, C., F. Monniot 272 272 k, . 95 eot pn h tnct i h cleto o h Ida Museum. Indian the of collection the in tunicata the upon Report 1915. A. Oka,

k, . 96 n nw eu o cmon ascidians compound of genus new a On 1926. A. Oka,

k, . 97. Uber 1927a. A. Oka,

k, . 97. u Knti dr aaice Btylde Poedns f the of Proceedings Botryllidae. japanischen der Kenntnis Zur 1927b. A. Oka,

k, .&A ily 19.O e eu fsnsiin rm aa. Quarterly Japan. from synascidian of genus new a 1892. On Willey. & A. A. Oka,

Peres,

Quoy,

egnta, . . 1984. K. T. Renganathan,

egnta, T. Renganathan,

h, .J 17.O te lsiiain n te itiuin f h mrn benthic marine the of distribution the and classification the On 1975. J. B. Rho,

h, . . & J. B. Rho,

Rho,

itr W . 90 oeacdas rm ue Sud cletos f 1896. Annals of collections Sound, Pujet from ascidians 1900. Some E. W. Ritter,

itr WE &RA Fryh 11.Acdas fte itrl fSuhr Califor Southern of littoral the of 1917. Ascidians Forsyth. R.A. & W.E. Ritter,

Romanov, V. N. 1989. Colonial ascidians of the family Didemnidae from the seas the from Didemnidae family the of ascidians 1989. Colonial N. V. Romanov,

at, . H Mukai H. Y., Saito,

at, Y. Saito,

at,Y,H ua & Mukai H. Y., Saito,

eor o h Ida Msu 6 1-33. 6: Museum Indian the of Memoirs

rceig o h Ipra Aaey Jpn : 133-135. 2: Japan Academy, Imperial the of Proceedings

Proceedings of the Imperial Academy, Japan 3: 555-557. 3: Japan Academy, Imperial the of Proceedings

Imperial Academy, Japan 3: 607-609. 3: Japan Academy, Imperial

Journal of Microscopical Science (n. ser.) 33: 313-323. 33: ser.) (n. Science Microscopical of Journal

iin o htet be rcre fo Ida Gois e Rpr 3 54- 3: Report New Geobios India. from recorded been hitherto not cidian

lvlnde Acdae: posbacit) el ein eMrele Re Marseille. de region la de Aplousobranchiata) (Ascidiacea: Clavelinidae cueil des Travaux de la Station marine d'Endoume 9: 67-71. 9: d'Endoume marine Station la de Travaux des cueil

ogie. Paris 1830-1934 3: 559-626. 3: 1830-1934 Paris ogie.

55.

ultn u uem ainl 'itie auel, ai () 6 () 257 (2): 6A (4), Paris naturelle, d'Histoire National Museum du Bulletin 262.

nml i Kra 3 Acdas Junl f oen eerh nttt for Institute Research Korean of Journal Ascidians. 3. Korea. in animals

Journal of Korean Research Institute for Better Living. 33: 99 33: Living. Better for Institute Research Korean of Journal

Island, Korea. Korean Journal of Systematic Zoology 5: 59-76. 5: Zoology Systematic of Journal Korean Korea. Island,

fte e Yr Aaey fSine 12: 589-616. Sciences of Academy York New the of

i. nvriy fClfri Pbiain i olg 16: 439-512. Zoology in Publications California of University nia.

Better Living. 15: 121-169. 15: Living. Better

ascidians

he nw pce o te genus the of species new Three (1):1 I s.) (n. SSSR. Fauna waters. adjacent and USSR the of sian).

lications of the Seto Marine Biological Laboratory 26: 347 26: Laboratory Biological Marine Seto the of lications

styelid ascidians II. A new species of the genus the ofspecies new A II. ascidians styelid of 26: 357 26:

Publications of the Seto Marine Biological Laboratory 30: 227 30: Laboratory Biological Marine Seto the of Publications

J.-M.

. violaceus B.

J-. e. 99 sseai suy n h acdas rm Cheju from ascidians the on study systematic A 1989. Lee. J.-E. &

J.-P.

93 eaqe sseaius t ilgqe sr ex see de especes deux sur biologiques et systematiques Remarques 1953.

. aaae 18. tde o Jpns cmon acin IV. ascdians compound Japanese on Studies 1985. Watanabe. H.

368.

M.-K.

Two new species of species new Two

amr. 84 oae e eovre d !Atoae; Zool !"Astrolabe'; de Decouvertes de Voyage 1834. Gaimard.

k. ulctos fte eo aie ilgcl Laboratory Biological Marine Seto the of Publications Oka.

.Mnit 18.Adtos oteacda fua oflndia. fauna ascidian the to 1984. Additions Monniot. & F.

u. 94 sseai suy n h acdas n Korea. in ascidians the on study systematic A 1984. Huh.

Dendroclavel/a,

aaae 18a tde fJpns cmon styelid compound of Japanese 1981a. Studies Watanabe.

Monniot & Monniot: New Ascidians New Monniot: & Monniot

censii garstangi Ecteinascidia

aaae 18b Suis n aaee compound Japanese on Studies 1981b. Watanabe.

Botryllus

Botrylloides

ie ee atn vn oiln Ascidien. sozialen von Gattung neue eine

rm h iiiy fSioa Pub Shimoda. of vicinity the from

rm h vcnt o Shimoda. of vicinity the from

Botrylloides

lie, 1898 Sluiter,

(Syndiazona

n redescription and

355.

clna as colonial A

225. (in Rus (in 225.

136.

240.

o. gen.). nov.

273

- -

zoologique

Societe

de la

Memoires

(3): 115-150.

R. Semon, Zoologische Forschungsreisen in

Micronesica 29(2), 1996

fran9aise).

98.

1895. Tunicaten. 1895.

P. 1909. Die Tunicaten der Siboga-Expedition. II. Die merosomen

P. Tunicaten 1900. aus dem StillenOcean. Zoologischer Jahrbucher.

P. Die Evertebraten 1890. aus der Sammlung des Koniglichen Natur

P. Die Tunicaten 1904. der Siboga-Expedition. Pt. I. Die und socialen

Skizze der Skizze Fauna des Java-Meeres mit Beschreibung der neuen Arten. As

kunding Tijdschrift Nederlandsch-Indie 45: 160-232.

cidiae simplices. Naturkunding simplices. cidiae Tijdschrift Nederlandsch-Indie 329-348. 50:

Australien und den malagischen Archipel. Denkschriften der medecinisch naturwissenschaftlichen Gesellschaft zu Jena 8: 163-186. Nachtrag zu den Tunicaten: 325-326.

einer Reise von Prof. Max Weber in Jahre 1894. II. Tunicaten von Siid Africa. II. Tunicaten von Siid Jahre in WeberProf. von Max 1894. Reise einer Zoologischer Jahrbucher. Abtheilung fiir Systematik, Geographie, und Biol ogie der ogie Thiere 1-64. 11: wissenschaftlichen Veriens in Niederlandish Veriens wissenschaftlichen Indien in Batavia. eine Zugleich

Abtheilung fiir Systematik, Geographie, und der Biologie Thiere 1-35. 13:

holosomen Ascidien. Siboga Expeditie 56A: 1-139.

de Tadjourah (Somalie France 5-21. 18:

Museum von Amsterdam. Bijdragen tot de Dierkunde 21: 1-21.

Marine LaboratoryBiological

from the pearl-oysterfrom thethe ArafuraPublications in bed theof Seto in 1940. Sea Marine Laboratory Biological 4: 75-98.

239-264.

ascidians in Osaka Bay. Publications of the Seto Marine Biological Labo ratory 4: 75

Marine LaboratoryBiological 43-57 5:

oranda (2). Publications of the Seto Marine Biological Laboratory 4: 205 Ascidien. Siboga. Expeditie 56A: 1-112. 218. von der naturforschen Senckenbergischen Gesellschaft 63-78. 35:

VII. Ascidians. Publications of the Seto Marine Biological Laboratory 3:

Sluiter, C.

Savigny, J.C. Savigny, Memoires sur 1816.Animaux les Sans Vertebres. Paris. pp. 239 Sluiter, C. P. Uber einfachen Ascidian voneinige 1885. der NaturBilliton. Insel

Sluiter,

274

Sluiter, C. P. Beitrage 1897. zur Kenntnis der Fauna von Siidafrica Ergebnisse

Sluiter,

Sluiter, C. P. 1905. Tuniciers recueillis en 1904 par en C. Sluiter, TuniciersP. Mr. recueillis 1904 1905. Ch. Gravier dans golfe le

Sluiter,

Sluiter, C. P. Ascidien von 1913. der Aru Inseln. Abhandlungen herausgegeben

Sluiter, C. P. Ober 1919. einige alte und neue Ascidien aus dem Zoologischen

Tokioka, T. Ascidians from 1950. the Palao Islands. I. Publications of the Seto

Tokioka, T. Ascidians collected 1952. by Messrs. Renzi Wada with Wada Seizi Tokioka, T. Ascidians from 1955a. the Palao Islands II. Publications of the Seto Tokioka, Contributions T. 1955b. to Japanese ascidian fauna. XI. mem Sporadic Tokioka, T. Ascidians of Sagami 1953. Bay. Iwanami Shoten, Tokyo. pp. 313 Tokioka, Invertebrate Tokara T.the faunaintertidal 1954a.the of zone of Islands.

Tokioka, T. Contributions 1954b. to Japanese ascidian fauna. X. Notes on some Monniot & Monniot: New Ascidians 275

Tokioka, T. 1958. Contributions to Japanese ascidian fauna. XII. Sporadic mem oranda (3). Publications of the Seto Marine Biological Laboratory 6: 331- 325. Tokioka, T. 1967. Pacific Tunicata of the United States National Museum. Bul letin of the U.S. National Museum 251: 1-242. Tokioka, T. 1970. Ascidians from Mindoro Island, the Philippines. Publications of the Seto Marine Biological Laboratory 18: 75-107. Tokioka, T. & T. Nishikawa. 1975. Contributions to the Japanese ascidian fauna. XXVII. Some ascidians from Okinawa, with notes on a small collection from Hong Kong. Publications of the Seto Marine Biological Laboratory 22: 323- 341. Tokioka, T. & T. Nishikawa. 1976. Contributions to Japanese ascidian fauna. XXX. Further notes on Japanese clavelinids. Publications of the Seto Marine Biological Laboratory 23: 341-350. Trason, W. B. 1957. Larval structure and development of the oozooid in the ascidian Euhermania claviformis. Journal of Morphology 100:509- 546. Trason, W. B. 1963. The life cycle and affinities of the colonial ascidian Pycno clavella stanleyi. University of California Publications in Zoology 65: 283- 326. Traustedt, M. P. A. 1882. Vestindiske Ascidiae simplices I abt. Phallusiidae. Vi denskabelige Meddelelser fra den naturhistoriske Forening i Kjobenhavn 1881: 1-32. Traustedt, M. P.A. 1885. Ascidiae simplices fra det Stille Ocean. Videnskabelige Meddelelser fra den naturhistoriske Forening i Kjobenhavn 1884: 1-60. Van Name, W.G. 1902. The ascidians of the Bermuda Islands. Transactions of the Connecticut Academy of Arts and Sciences 11: 325--412. Van Name, W.G. 1918. Ascidians from the Philippines and adjacent waters. Bul letin of the U.S. National Museum 100: 49- 174. Van Name, W.G. 1921. Ascidians of the West Indian region and southeastern United States. Bulletin of the American Museum of Natural History 44: 283- 494. Van Name, W. G. 1930. The ascidians of Porto Rico and the Virgin Islands. Transactions of the New York Academy of Sciences I 0: 401-512. Vasseur, P. 1969. Deuxieme contribution a l'etude des ascidies de Madagascar region de Tulear. Bulletin du Museum national d'Histoire naturelle, Paris (2) 40 (5): 912-923. von Drasche, R. 1882. Oxycorynia, eine neue Synascidien-Gattung. Verhandlun gen der kaiserlichen zoologische-botanische Gesellschaft in Wien 32: 175- 178. von Drasche, R. 1884. Ueber einige neue und weniger gekannte aussereuropaische einfache Ascidien. Denkschriften der kaiserlichen Akademie der Wissen schaften zu Wien 48: 369-386 Watanabe, H. & T. Tokioka. Two new species and one possibly new race of social styelids from Sagami Bay, with remarks on their life history especially the mode of budding. Publications of the Seto Marine Biological Laboratory 19: 327- 345. 142

135 137

139

183

152

153 155

231

157 158

238

205

210 212

217 219

222

224 226

......

from Japan,

Clavelina

Central and Western Pacific.

......

...... 140

...... 157

......

...

1987

....

1976 ...... 1976 208

...... 1987 ...... 1987 159

......

Index

On a new species of

1891}

1988 ...... 1988 199

1995 ...... 1995 152

1987 ...... 1987 139

1886}

1892 ...... 1892 199

Monniot,

1952 ...... 1952 205

Tokioka,

......

1957) ...... 1957) 203

107.

...... &

1955) ...... 1955) 220

1886) ...... 1886)

Micronesica 29(2), 1996 ...... 1988 200

1973.

1909) ...... 1909) 153

1985) ...... 1985) 241

sp ...... sp

1992

sp ...... sp

Nott,

1990

......

(Kott,

(Herdman,

Monniot,

......

Monniot,

21: 99

......

sp ...... sp

Tokioka,

...... 142

Coral Reefs of the World,

n. sp ...... n. sp n. sp ...... n. sp

n. sp ...... n. sp

sp ...... sp

n. sp ...... n. sp

(Herdman,

(Kott,

sp ...... sp

Kott,

Monniot

sp ...... sp

n. sp

(Sluiter,

Monniot,

sp_

(Tokioka,

Kott,

....

Monniot & Monniot,

n. sp

(Herdman,

Nishikawa

......

n. sp ...... n. sp

1988.

B .. B

sp ...... sp

sp. sp.

logical Laboratory

IUCN.

with remarks on its mode of budding. Publications of the Seto Marine Bio

Clavelina arafurensis Clavelina meridionalis

Cystodytes auck/andicus? Cystodytes punctatus? Cystodytes so/itus

Clavelina obesa? Clavelina robusta Clavelina

Amphicarpa agnata Aplidiopsis ocellatus Aplidium controversum Aplidium lineatum Aplidium /ongithorax Aplidium tabascum Aplidium Aplidium Archidistoma diminuta Ascidia azurea Atrio/um quadratum Bo try I/us firmus I/us try Bo

Didemnum apuroto Didemnum big/utinum

Diazona angulata Diazona carnosa Diazona chinensis Diazona formosa Diazona Diazona labyrinthea Diazona tenera

Didemnumfragi/is Didemnum guttatum Didemnum linguiferum Didemnum mo/le Didemnum mose/eyi Didemnum nigrum Didemnumparau

Received Sep 19 1995, revised April 10 1996.

276

Watanabe, H. & T. Tokioka.

Wells, S.W. (ed.)

Polycarpa cryptocarpa cryptocarpa Polycarpa

Polycarpa captiosa captiosa Polycarpa

Polycarpa aurata aurata Polycarpa

Polycarpa argentata argentata Polycarpa

Nephtheis fascicularis? fascicularis? Nephtheis

Microcosmus manaarensis? manaarensis? Microcosmus

Lissoc/inum voeltzkowi? voeltzkowi? Lissoc/inum

Lissoc/inum patella patella Lissoc/inum

Plurella kottae kottae Plurella

Phal/usiajulinea Phal/usiajulinea

Phallusia arabica arabica Phallusia

Leptoc/inides dubius dubius Leptoc/inides

Lissoc/inum tunicatum tunicatum Lissoc/inum

Lissoc/inum nebulosum nebulosum Lissoc/inum

Lissoc/inum badium badium Lissoc/inum

Leptoclinides uniorbis uniorbis Leptoclinides

Leptoc/inides sulawesii sulawesii Leptoc/inides

Leptoc/inides reticulatus reticulatus Leptoc/inides

Leptoc/inides oscitans oscitans Leptoc/inides

Hypodistoma deerratum deerratum Hypodistoma

Herdmania momus momus Herdmania

Exostoma ianthinum ianthinum Exostoma

Eusynstyela latericius latericius Eusynstyela

Eudistoma Eudistoma

Eudistoma toea/ensis toea/ensis Eudistoma

Eudistoma rubiginosum rubiginosum Eudistoma

Eudistoma gilboviride gilboviride Eudistoma

Eudistomafascicu/um Eudistomafascicu/um

Eudistoma amp/um amp/um Eudistoma

Ecteinascidia styeloides styeloides Ecteinascidia

Eudistoma reginum reginum Eudistoma

Eudistoma laysani? laysani? Eudistoma

Eudistoma incrustatum incrustatum Eudistoma Didemnum Didemnum

Ecteinascidia diaphanis diaphanis Ecteinascidia

Distaplia regina? regina? Distaplia

Diplosoma virens virens Diplosoma

Diplosoma similis similis Diplosoma Diplosoma multitestis multitestis Diplosoma

Didemnum Didemnum

Didemnum stercoratum stercoratum Didemnum

Didemnum rubeum rubeum Didemnum

Didemnum psammathodes psammathodes Didemnum

Didemnum poecilomorpha poecilomorpha Didemnum

Didemnum per/ucidum per/ucidum Didemnum

sp ...... 196 196 sp ......

sp. A ...... 167 167 A ...... sp.

sp. sp.

R R

n. sp ...... 235 235 sp ...... n.

Sluiter, 1919...... 235 235 1919. Sluiter, ......

......

Kott, 1990 ...... 216 216 1990 ...... Kott,

(Hartmeyer, 1909) ...... 170 170 1909) ...... (Hartmeyer,

Savigny, 1816 ...... 233 233 1816 ...... Savigny,

(Quoy & Gaimard, 1834) ...... 249 249 1834) ...... Gaimard, & (Quoy

(Sluiter, 1909) ...... 170 170 1909) ...... (Sluiter,

Savigny, 1816 ...... 263 263 1816 ...... Savigny,

(Sluiter, 1885) ...... 249 249 1885) ...... (Sluiter,

n. n.

(Gottschaldt, 1898) ...... 174 174 1898) ...... (Gottschaldt,

(Sluiter, 1900) ...... 190 190 1900) ...... (Sluiter,

(Sluiter, 1909) ...... 185 185 1909) ...... (Sluiter,

Kott, 1990 ...... 191 191 1990 ...... Kott,

n. sp ...... 170 170 sp ...... n.

(Sluiter, 1909) 1909) (Sluiter,

(Sluiter, 1890) ...... 247 247 1890) ...... (Sluiter,

(Sluiter, 1909) ...... 197 197 1909) ...... (Sluiter,

Millar, 1975...... 194 194 1975...... Millar,

s.str. (Sluiter, 1904) ...... 241 241 1904) ...... (Sluiter, s.str.

n. sp ...... 182 182 sp ...... n.

n. n. n. sp ...... 177 177 sp ...... n.

(Sluiter, 1909) ...... 188 188 1909) (Sluiter, ......

sp sp

n. sp ...... 187 187 sp ...... n.

n. sp ...... 174 174 sp ...... n.

(Drasche, l 882) ...... 201 201 l 882) ...... (Drasche,

n. sp sp n.

Monniot, 1981 ...... 160 160 1981 ...... Monniot,

(Sluiter, 1885) ...... 251 251 1885) ...... (Sluiter,

n. sp ...... 173 173 sp ...... n.

n. sp ...... 189 189 sp ...... n.

Monniot Monniot

Sluiter, 1885 ...... 227 227 1885 Sluiter, ......

n. sp sp n.

n. sp ...... 165 165 sp ...... n.

(Traustedt, 1882) ...... 229 229 1882) ...... (Traustedt,

(Michaelsen, 1920) ...... 175 175 1920) ...... (Michaelsen,

......

(Sluiter, 1895) ...... 198 198 1895) ...... (Sluiter,

(Sluiter, 1909) ...... 178 178 1909) ...... (Sluiter,

sp ...... 168 168 sp ......

(Sluiter, 1895) ...... 163 163 1895) ...... (Sluiter,

n. sp ...... 160 160 sp ...... n.

Herdman, 1906. 1906. Herdman,

......

& &

Monniot: New Ascidians Ascidians New Monniot:

......

265 265

180 180

176 176

193 193

167 167

164 164 277 277

F. F.

H, H,

Lisso

214 214

213 213

260 260

252 252

. .

......

n. n. sp. C,

Atriolum Atriolum

Monniot Monniot

Ap/idium Ap/idium

Aplidium Aplidium

Didemnum Didemnum

H, H,

(Sluiter, (Sluiter, 1909).

Pseudodistoma Pseudodistoma

Nott, Nott, 1892.

......

n. n. sp. F,

Clavelina Clavelina obesa

n. n. sp. C and D,

sp. sp. E,

Eudistomafasci

Leptoc/inides Leptoc/inides du

n. n.

n. n. sp. C,

H, H,

Millar, Millar, 1975. F,

Kott, Kott, E, 1992.

n.sp. n.sp. H,

Didemnum Didemnum biglutinum

Clavelina Clavelina arafurensis

n. n. sp.

Didemnum Didemnum parau

......

Monniot, Monniot, 1983.

. .

......

...... 245

...... 147

...

...... 258

...... 145

Pseudodistoma Pseudodistoma poculum

...... 254

Lissoc/inum Lissoc/inum badium

...

(Sluiter, (Sluiter, 1909). G,

n. n. sp. F,

....

Ap/idium Ap/idium /ineatum

(Herdman, (Herdman, 1891). D,

(Sluiter, (Sluiter, 1909).

....

Leptoc/inides Leptoc/inides reticulatus

Eudistoma Eudistoma rubiginosum

Cystodytes Cystodytes aucklandicus

B, B,

Leptoc/inides Leptoc/inides uniorbis

Eudistoma Eudistoma toealensis

...

Didemnum Didemnum stercoratum

Aplidium Aplidium tabascum

n. n. sp. B,

D, D, ..

......

Monniot, Monniot, 1987. D,

n. n. sp. E,

B, B,

Aplidiopsis Aplidiopsis ocel/atus

......

n. n. sp.

(Drasche, (Drasche, 1882). B,

Color Color Plates

& &

sp. sp.

n. n. sp. B,

Lissoc/inum Lissoc/inum tunicatum

......

1885} 1885}

Micronesica Micronesica 29(2), 1996

n. n.

n. n. sp.

sp ...... sp 148

1955} 1955}

1909} 1909}

(Sluiter, (Sluiter, 1895).

n. n...... sp

G, G,

B. B. G,

n. n...... sp

n. n...... sp 150

n. n.

Didemnumperlucidum Didemnumperlucidum

Eudistoma Eudistoma amp/um

sp. sp.

(Herdman, (Herdman, ...... 1906) 252

Kott, Kott, 1990. E,

Monniot Monniot

Michaelsen, Michaelsen, ...... 1921 149

Didemnum Didemnum guttatum

(Sluiter, (Sluiter, 1909). G,

n; n; sp.

(Sluiter, (Sluiter,

n. n...... sp 255

Clave/ina Clave/ina meridionalis

Michaelsen, Michaelsen, ...... 1927 263

......

(Millar, (Millar,

Eudistoma Eudistoma gilboviride

n. n...... sp

(Sluiter, (Sluiter,

n. n. sp.

Monniot Monniot & Monniot, ...... 1987 144

Michaelsen, Michaelsen, ...... 1930 184

n. sp ...... n. sp

B ...... B

F. F. Monniot, D, 1987.

n. n. sp. F,

n.sp. n.sp.

Aplidium Aplidium

Diplosoma Diplosoma multitestis

sp. sp.

sp. sp...... C 262

sp. sp. .. A

Nephtheis Nephtheis fascicularis

Pseudodistoma Pseudodistoma mega/arva

SP-······················································································· SP-·······················································································

Aplidium Aplidium controversum

Eudistoma Eudistoma incrustans

n. n. sp. H,

Leptoclinides Leptoclinides oscitans

Didemnum Didemnum rubeum

(Sluiter, (Sluiter, 1909).

A. A. F,

Leptoc/inides Leptoc/inides sulawesii

kioka, kioka, 1952. C,

clinum clinum nebulosum

sp. sp. B. D,

Hypodistoma Hypodistoma deerratum

ostoma ostoma ianthinum

Eudistoma Eudistoma reginum

Monniot, Monniot, 1995. E,

culum culum

quadratum quadratum

C, C,

coronatum coronatum

sp. sp.

bius bius

/ongithorax /ongithorax

& & Monniot, G, 1987.

Didemnum Didemnum apuroto

poecilomorpha poecilomorpha

Trididemnum Trididemnum polyorchis

Trididemnum Trididemnum cyclops

Stolonica Stolonica limbata

Sigil/ina Sigil/ina

Sigillina Sigillina signifera

Stomozoa Stomozoa roseola

Pyura Pyura albanyensis

Pseudodistoma Pseudodistoma poculum

Pseudodistoma Pseudodistoma megalarva

Polycarpa Polycarpa

Po/year pa pa Po/year

Po/year pa pa Po/year tokiokai

Po/year pa pa rima Po/year

Pseudodistoma Pseudodistoma tum corona

Polyc/inum Polyc/inum pute

Polycitor Polycitor circes

Polycarpa Polycarpa

Po/ycarpa Po/ycarpa pigmentata

Polycarpa Polycarpa papillata

Plate Plate 5. A,

Plate Plate 4. A,

Plate Plate 6. A,

Plate Plate A, 3.

Plate Plate 2. A,

Plate Plate A, 1. 278 278 Plate 7. A, 7. Plate

Plate 8. A, 8. Plate

Plate 9. A, 9. Plate

Nishkawa & Tokioka, 1976. E, 1976. Tokioka, & Nishkawa

and

sp. G, sp.

carnosa sp. F. sp. n.sp.

Sutr 1904). G, (Sluiter,

cryptocarpa carpa pigmentata carpa

tokiokai

1890).

izn labyrinthea Diazona

Sigillina signifera Sigillina itpi regina? Distap/ia

hlui arabica Phallusia

oyap captiosa Polycarpa

Botryl/us firmus

n. sp. D, sp. n.

n. sp., with sp., n.

Sutr 1885). D, (Sluiter,

izn chinensis Diazona

Hrmn 1906). F, (Herdman,

tlnc limbata Stolonica

ienm mo/le. Didemnum

Monniot & Monniot: New Ascidians New Monniot: & Monniot

n. sp. n. E,

Sutr 1909). H, (Sluiter,

ain, 1816. Savigny,

ot 1990. B, Kott,

Sutr 1885). B, (Sluiter,

n. sp. G, sp. n.

oyap papillata Po/ycarpa

Plurella kottae

/vln robusta C/avelina

Tkoa 1955). E, (Tokioka,

izn tenera Diazona

. sp. n.

izn angulata Diazona

oyap rima Polycarpa

Polycarpa

Phallusiajulinea

Sigil/ina

Polycarpa

n. sp. n. F,

oyap argentata Polycarpa

Kott, 1990. F, 1990. Kott,

Sutr 1885). E, (Sluiter,

n. sp. n.

sp.

sp. B. sp.

uysyl latericius Eusynstyela

p C C, C. sp.

Diazonaformosa

.p G, n.sp.

n. sp. C, sp. n.

lie, 1919. C Sluiter,

sii azurea Ascidia

Polycarpa

Po/ycarpa

Clavelina

Diazona

(Sluiter,

Poly

279

Monniot & Monniot Monniot & Monniot Plate 1 1 Plate

Monniot Monniot

Plate 2 2 Plate

& & Monniot Monniot

Monniot Monniot

Plate 3 3 Plate

& & Monniot Monniot

Monniot Monniot

Plate 4 4 Plate

& & Monniot Monniot Monniot & Monniot & Monniot

lt 5 Plate Monniot

Plate 6 Plate

Monniot Monniot & Monniot & Monniot

Plate 7 Plate Monniot

Plate 8

Monniot Monniot & Monniot & Monniot

Plate 9 Plate