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BULLETIN DU MUSÉUM NATIONAL D'HISTOIRE NATURELLE

3 e série, n° 403, septembre-octobre 1976, Zoologie 280

Two new species of Echiura

from Madagascar with a note on

the male of Metabonellia hasuelli

by Alan J. Dartnall *

Résumé. Deux nouvelles espèces d 'Echiura sont décrites du Grand Récif de Tuléar,

Madagascar. Une des espèces est signalée aussi en Australie. Le mâle de Metabonellia haswelli

(Johnston et Tiegs, 1920) est décrit pour la première fois.

Abstract. — Two new species of Echiura are described from the Great Reef of Tuléar,

Madagascar. One of the species is also recorded from Australia. A description is given of the

male of Metabonellia haswelli (Johnston and Tiegs, 1920).

Introduction

The echiuran material described in this paper results, in the main, from the studies

of Dr B. Thomassin upon the reef communities of the Great Reef of Tuléar, Madagascar.

The opportunity has been taken to include an extra honellid record from Australia and

a description of the male of Metabonellia haswelli (Johnston and Tiegs, 1920).

The taxonomie scheme followed is that of Stephen and Edmonds (1972). Type

and voucher material has been deposited in varions Muséums referred to in the lists by

initial abhreviations viz. : Australian Muséum, Sydney = AM ; British Muséum (Natural

History), London = BMNIJ ; Muséum national d’Histoire naturelle, Paris - MNTIN ;

Tasmanian Muséum, Hobart = TM.

BONELLIIDAE

Genus BONELLIA Rolando

Bonellia sabulosa sp. nov.

(Fig. 1-2)

Description of female

Body sac-like with a long proboscis, bifurcate at the tip. The specimens available

were in various stages of expansion and contraction. The relative proportions of body

* Muséums and Art Galleries oj the Northern Territory, P. O. Box 4616, Darwin, 5794, Australia.

403 , 1

1040

ALAN J. DARTNALL

and proboscis form may be seen in figure 1. The specimens were white in colour (preserved

in formalin). The epidermis is thin and transparent on the dorsal surface and thicker

and verrucose on the ventral surface particularly towards the extremities of the bodv.

In three specimens froni station 843A (see locality details) a longitudinal band of

small white papillae arranged roughly in two rows is found on the ventral surface. The

band of papillae extends along the body from the anus to the bifurcation of the proboscis.

On the proboscis the papillae are visible from both sides under low power magnification.

O '—I—I— 1 —l—l—I—I—I—I—I 10 mm

Fig. 1-2. — Bonellia sabulosa sp. nov. : 1, Body form of five specimens ; 2, Nephridium.

A pair of setae is présent, similar in form to those described from Bonellia pilosa Dart-

nall, 1973. Each seta is about 1.5 mm long and lias a reflexed tip.

The general anatomy of the female is similar to tliat of B. pilosa. An intestinal siphon

is présent and a pair of anal vesicles arise from the intestine near the anus. The anal

vesicles are about 2.5 mm long and hâve about forty branches, each branch terminating

in a ciliated funnel.

One nephridium is présent. In sonie specimens the nephridium was surrounded by

the first convolution of oesophagus and foregut. The nephridium is found either on the

right or the left side. Developing eggs were found in the nephridia of four of the females

available. Some nephridia were empty and deflated ; others gravid and swollen. The

nephridial form is illustrated in figure 2. The nephrostome is basal.

The ovary is spread along the ventral blood vessel. A neuro-intestinal linking vessel

is présent.

TWO NEW SPECIES OF ECHIURA

1041

Description of male

The male is apparently nearly identical to that of B. pilosa ( loc . cit.). It is about

O. 8 mm long, has two identical hooked setae (except in two specimens from a female from

station 629 where no setae were apparent), a vestigial gut and a single nephridiurn which

opens subterminally. As in B. pilosa the males were ail found in the pharynx of the females ;

no males were found in any other part of the body or on the proboscis. The reflexed tail

so apparent in males of B. pilosa, was présent in one male of the sériés but not apparent

in the remaining individuals (i.e. one of four).

Type locality and holotype :

The Great Reef of Tuléar, ofï S.W. Madagascar is designated the type locality. The holo¬

type is one of nine specimens collected from a 50 dm 3 shovelled sample of a gravelly, coral bank

on the reef fiat at infralittoral level. Collected 12.IV.1972 ; B. Thomassin (station 629), MNHN.

Allotype : $ AF882 (MNHN) ; from Ç from station 843B (see below).

Paratype material : Eight specimens from same sample as holotype (four in AM, four in TM).

Five specimens from 26 dm 3 gravel, Grand Récif de Tuléar, Madagascai at the entrance of an

outer creek (« Grande Crique ») on the reef front at 10 m depth. Collected 8.VII.1972 ; B. Tho¬

massin (station 843A), TM. Seven specimens from 14 dm 3 gravel at station 843B (same locality

details as 843A), MNHN.

Other material : Eleven specimens from 35 dm 3 gravel from gravel ridge at mouth of outer

creek, Great Reef of Tuléar, Madagascar. Collected 6.VII.1972 ; B. Thomassin (station 837).

Three specimens from 40 ft, One Tree Island, Great Barrier Reef, Australia. Collected 4.N.1972 ;

P. Hutchtngs (AM).

Discussion

The new species is the second species of Bonellia reported from Madagascar. It. may

be distinguished from Bonellia pilosa Dartnall by the absence of investing hairs on the

body and by the branching tubules of it.s anal vesicles. It is apparently close to Bonellia

punicea Sluiter, 1891, for which he type locality is Krakatoa and with which it may yet

prove to be conspecific. In that case Sluiter’s name may, eventually, take precedence.

However, the female of B. punicea is supposée! to hâve a body shaped like a bail which

none of the specimens to hand resemble. Also, the. white papillae of B. punicea are denser

at the ends of the body, a condition not apparent in the three specimens (from Madagascar)

which carried white spots on the body.

Metabonellia haswelli (Johnston and Tiegs, 1920)

(Fig. 3-6)

The male of this species has not been described. Two female specimens of M. has¬

welli were made available from the collections of the Australian Muséum, Sydney (from

Fairlight, Manly, New South Wales ; collected 2.VI.1959 ; E. C. Pope and party).

The nephridia of the female specimens were fully expanded and consisted of a proximal,

1042

ALAN J. DARTNALL

heavily-banded area about 6 mm long followed by a médian and a terminal sac eacli about

IG mm long carrying tho nephrostome at their junction.

This cojitrasts with Johnston and Tiegs’ (1920) figure 1 of plate XV and their

account in whicb anv subdivision of the “ utérus ” was not emphasised though the authors

did point out “ that neither specimen was sexuallv mature

Fig. 3-5. — Metabonellia haswelli : 3, Nephridium ;

4, Nephridial tissue mass with eggs and embedded males ; 5, Male.

The nephridium of each specimen contained an elongated, yellowish mass of tissue

with eggs embedded in, and adhering to the surface. In one specimen males were absent ;

in the other males were found embedded in the nephridial tissue mass.

The males removed front the tissue were about 8 mm long. The male possesses a

single nephridium and an ill-defined gut. The coelom contained a few spernt morulae.

Just behind the anterior end the surface of the animal is folded into a ltood shape which

is applied to the tissue mass. I hâve been unable to détermine whether this area of the

male body sécrétés the material in which the eggs are embedded or whether the male uses

this area as an attachment sucker. The males examined had neither setae or an anal

clasper.

Sections of the nephridial mass were eut at 10 y. and stained with Erhlich’s haemat-

oxylin and oesin. The tissue is vacuolar, stains bine with haematoxylin and encloses the

eggs. The eggs are about 0.2 mm in diameter, possess a central nucléus and the cytoplasm

is bordered with large globules. Connections between the tissue and the enclosed eggs were

also observed (see fig. 6).

TWO NEW SPECIES OF ECHIURA

1043

This work was started to test the hypothesis that the echiurans from south-eastern

Australia attributed to genus Bonellia deserved séparation from that genus. Later it was

fourni that generic séparation had been rnade by Stephen and Edmonds (1972) by érection

of genus Metabonellia. This description of the male of M. haswelli supports the action

of those authors. The species placed in the genus ail hâve males without claspers.

The significance of the situation in which the males of M. haswelli are found must

remain for a worker with access to living specimens who can ascertain whether the tissue

mass is an artefact of préservation.

1044

ALAN J. DARTNALL

THALASSEMATIDAE

Genus OCHETOSTOMA Leuchkart and Ruppell

Ochetostoma punicea sp. nov.

(Fig. 7-8)

Description

Body form is as shown in figure 7. The proboscis is firmiy attached to the body and

is short and stumpy with its latéral edges turned inwards so that they meet near the month.

The edges, however, are not fused. The proboscis is about 9 mm long and about 7.5 mm

wide at its widest part (in the holotype) and is pink in colour (formalin preserved).

Fig. 7-8. — Ochetostoma punicea sp. nov. : 7, Body form (holotype) ; 8, a) anterior and posterior nephri-

dia from right-hand side of hody showing nephrostomes, b) rectum, anal vesicles and rectal caecum

with mesentery to nerve cord.

TWO NEW SPECIES OF ECHIURA

1045

The body is about 28 mm long and about 14 mm across its greatest width. The

anterior part of the body carries a pair of ventral setae and the body surface is thickened

and papillate espeeially on the dorsum and the ventro-lateral shoulders. The médian part

of the body surface is thinner than the preceding and the outline of the internai organs

eau be seen through the body wall. The posterior portion is tough and opaque carrying

prominent papillae attaining a millimétré in diameter. The end of the body is drawn

out into an apical projection where the anus opens to the exterior. The body is pinkish-

brown becoming browner where the papillae are more prominent.

The longitudinal muscles are présent in nineteen or (in one specimen) eighteen bundles.

Fasiculation of the oblique muscles is rather weak in the médian and forepart of the body

wall but clearly visible to the rear.

The setae are about 2.25 mm long ; the main stem is about 0.3 mm wide and 1.5 mm

long. The setal apex is pointed and bent at about 45°. T'here is no interbasal muscle.

Two pairs of nephridia are présent, both opening behind the setae. The nephrostomes

are weakly spirallv coiled. In the holotype the anterior nephridium is about 8 mm long

and the posterior nephridium is about 10 mm long with the free end of the nephridium

more tapered. The nephridia are brown in colour . A circumpharyngeal ring vessel is

présent.

A rectal caecum is présent on the ventral surface of the gut. It is attached to the

nerve cord liy a mesentery. The anal vesicles are long (c. 19 mm), brown and unbranched.

Ciliated funnels are scattered over the surface of each vesicle. The left anal vesicle, in ail

specimens, was found to follow the convolutions of the gut closely whilst the right anal

vesicle ran along the body wall for about 2/3 of its length after which it bends back to

attach to the intestine with a fine mesentery.

One specimen (from station 755) difïers only in colour from the other specimens and

this may be due to différences in préservation. Most of the body of the specimen is translu-

cent and white except at the posterior end where it is pink..

Type locality and holotype : The Great Reef of Tuléar, off S. W. Madagascar is designated the

type locality. The hclotype is one of 27 specimens collected from 50 dm 3 sample of gravel and

silty sand from a boulder tract on the reef fiat of the N. Grand Récif. Collected 1.VI. 1972 ;

B. Thomassin (station 757), MNHN.

Paralypes : Twenty-six specimens from the same sample as holotype ; AM, BMNH, MNHN

and TM. Twentv-five specimens from a further station (station 755) ; data identical to holotype

details.

Discussion

Ochetostoma capense Jones and Stephen, 1955, has already been reported from the

Tuléar région (Thomassin, 1973). Ochetostoma punicea sp. nov. shares a posterior apical

projection with that species and appears similar in the structure of the anal vesicles and

in the presence of a rectal caecum. The new species difïers markedly iri the number of

longitudinal muscle bands — 7 in O. capense, 18/19 in O. punicea.

The other species of Ochetostoma with two pairs of nephridia and 18/19 longitudinal

muscle bands are :

1. Ochetostoma baronii (Greeff, 1879) which has, however, branched outgrowths from

the anal vesicles.

1046

ALAN J. DARTNALL

2. Ochetostoma kefersteini (ten Broeke, 1925) whose first pair of nephridia open anterior

to the setae and which Stephen and Edmonds (1972) point ont may bo a Listriolobus

because it is not known whether the oblique muscle layer of this animal is fasiculated.

3. Ochetostoma myersae Edmonds, 1963, which lias a “ strong, interbasal muscle ” and

whose body papillae are “ soft, white and wartlike ”, quite unlike the tough papillae of

O. punicea,

These observations confirm the need, pointed out by Stephen and Edmonds (1972 :

427) for further study to define this genus. Also among the specimens sent to me for

study by Dr Tiiomassin was an extensive sériés of animais from the Great Reef of Tuléar

that I hâve determined as Ochetostoma erythrogrammon Leuckart and Ruppell, 1828. Varia¬

tion among these specimens emphasises the need for samples from widespread géographie

localities and the extensive synonymy provided by Stephen and Edmonds (loc. cit.) demon-

strates the variety of opinion amongst authors concerning the species of Ochetostoma.

Among the Tuléar specimens are individuals with nephridia varying from three to two

pairs, with one specimen having two left-hand nephridia and three right-hand nephridia.

Three specimens had the first pair of setae opening anterior to the setae and longitudinal

muscle bands varied between 16 and 18. The significance of variation such as this dépends

on further samples.

Acknowledgments

This work commenced at the Tasmanian Muséum, Hobart and was completed in the Muséums

and Art Galleries of the Northern Territorv. 1 am grateful to both institutions for facilities pro¬

vided. My thanks also to Dr Bernard Thomassin who allowed me to work on bis material and

was helpful in many other ways.

REFERENCES

Dartnall, A. J., 1973. — A new species of Bonellia (Echiuroidea, Bonelliidae) from Madagascar.

Téthys, suppl., 5 : 3-8.

Johnston, T. H., and O. W. Tiegs, 1920. ■— A new species of Bonellia from Port Jackson. Bec.

Aust. Mus. Sydney, 13 : 73-76.

Stephen, A. C., and S. J. Edmonds, 1972. — The phyla Sipuncula and Echiura. British Muséum

(Natural Ilistory), London, vm -)- 528 p.

Thomassin, B. A., 1973. — Peuplements des sables fins sui les pentes internes des récifs coralliens

de Tuléar (S.-W. de Madagascar) etc. Téthys, suppl., 5 : 157-220.

Manuscrit déposé le 17 février 1976.

Bull. Mus. natn. Hist. nat., Paris, 3 e sér., n° 403, sept.-oct. 1976,

Zoologie 280 : 1039-1046.

Achevé d’imprimer le 28 février 1977.

IMPRIMERIE NATIONALE

6 504 004 5

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