Bull. nat. Hist. Mus. (Zool.) 59(1): 1-9

Issued 24 June 1993

GARTH UNDERWOOD

; LJ P se INE PE

Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 sap, FREOCN I!

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VW AR’ LIMP

Synopsis. The Clelia clelia reported from Dominica and St Lucia are reinvestigated. The specimens concerned are

recognised as a new species, Clelia errabunda. It is derived in relation to the mainland species C. bicolor and C.

rustica and is the primitive sister species of the mainland C. clelia, C. equatoriana and C. scytalina. As such it is

interpreted as an island relict. Records from Dominica and Guyana are rejected, the St Lucia records are confirmed.

INTRODUCTION

_ Nearly a hundred years ago Boulenger (1896) reported the

South American snake Oxyrhopus clelia, now known as

Clelia clelia, from the Windward Islands of Dominica and St

_ Lucia, as well as from mainland localities. Ever since then

this species has been listed for these two islands, either in the

genus Clelia (Barbour, 1930; Schwartz & Henderson, 1988)

or Pseudoboa (Barbour, 1935, 1937). There are other records

of this snake for St Lucia but this remains the only record for

Dominica.

Boulenger (1896) distinguished two varieties of ‘Oxyrho-

pus clelia’: A, with 19 scale-rows at mid-body, and B, with 17

scale-rows. Under variety ‘B’ Boulenger lists: two specimens

from ‘The city of Mexico’, three specimens from “W. Ecua-

dor’ and one specimen each from ‘Demerara’, ‘Dominica’

— and ‘St Lucia’.

Bailey (1970, in Peters & Orejas-Miranda) gives a key to

the mainland species of Clelia. The city of Mexico specimens

key out as Clelia scytalina, the Ecuador specimens as C.

equatoriana. The remaining three specimens reach the

_ scytalina-equatoriana couplet of the key but are not clearly

assignable to either mainland species. Boulenger’s variety ‘A’

_ specimens key out as C.c.clelia and C.c.plumbea, the two

subspecies recognised by Bailey. Clelia was later reported

_ from the island of Grenada. It was described by Greer (1965)

as Clelia clelia groomei; Bailey places this in the synonymy of

~ C.c.clelia.

Further examination of these remaining three specimens

shows that they represent an unrecognised species. In the

hope of determining the affinities of the new form the

specimens of Clelia in the collection of The Natural History

_ Museum, London were examined. A Paris Museum specimen

_ from St Lucia was also examined. The seven species recogn-

_ised by Bailey are represented. It should be noted that

_ Scrocchi and Vinas (1990) have now sunk occipitolutea in the

synonymy of clelia.

MATERIALS AND METHODS

Each specimen was examined in respect of external features,

anterior viscera, maxillary teeth and, in the case of males,

hemipenis. For a representative of each species dissections

were made to show the superficial jaw muscles, ligaments and

labial glands and the upper and lower jaws.

The ventral scale count is according to Dowling (1951).

The notation for the scale-rows on the trunk allows a detailed

record in a single line of type using characters on a word

processor keyboard. An oblique transverse scale-row is iden-

tified by the ventral scale from which it passes backwards, as

shown by M.A. Smith (1943, fig. 10). Where there is a

scale-row fusion the upper of the two merged rows is indi-

cated. For example male specimen BMNH 89.4.8.2 is

recorded as:

V 210 >>5> 17 (36;5</33;5<) 19 (145;4>/142;4>) 17

Ventral scale count 210. Three scale-row fusions from the

back of the head onto the neck, the third recognisable as due

to the fusion of row 5 with the row below. Seventeen rows on

the neck through oblique row 36 left/33 right; row 5 splits

giving rise to 19 rows through oblique row 145 left/142 right;

row 4 merges with row below giving 17 rows. Most specimens

were recorded down the left-hand side only (Table 1, see

Appendix).

The subcaudal count starts with the first scale on the left

side which makes full contact with a scale obliquely forward

and opposite. The dorsal scale-rows on the tail are somewhat

irregular around the base but settle down to even row

stretches: 8, 6, 4, 2. The fusions are rarely symmetrical so that

there are short odd number transitions between the even

stretches. Each transition is included in the preceding,

higher, even row stretch. The transverse rows are identified

by the subcaudal scale-row pairs from which they arise

obliquely. The tail of the above specimen was recorded as:

79 prs, >8 (10) 8 (25) 6 (45) 4 (73) 2

More than eight longitudinal rows through oblique row ten,

eight (and seven) through row 25, six (and five) through row

45, four (and three) through 73 and two rows to the terminal

scute. From this record the lengths of the five stretches are

calculated as the basis of comparison of the individual speci-

mens. The above becomes:

C79 >8:10 8:15 6:20 4:28 2:6

This feature shows marked sexual dimorphism.

For selected specimens scales were mounted on slides to be

examined for the presence of pits and tubercles (Underwood,

1963). The wet scales are laid on a dried film of polyvinyl

alcohol lactophenol mounting medium. When dry the slide is

then covered using Canada balsam. For each specimen the

scales mounted are the frontal, a parietal and a vertical series

of scales at about mid-trunk from row one to the vertebral.

The immunological studies of Cadle (1984) suggest that

species of Pseudoboa are sister to Clelia; his sample included

the type species of each genus. He also found that Oxyrhopus

fitzingeri is closer to Clelia plus Pseudoboa than it is to other

species of Oxyrhopus. ‘Oxyrhopus’ fitzingeri is therefore

included as part of the outgroup. In the absence of an

analysis, the majority outgroup condition is taken to be

primitive for the ingroup.

The four species of Pseudoboa recognised by Bailey share

the special feature of single subcaudals. No evident special

feature unites the species of Clelia. On further study the

boundary between the two genera may be redrawn, or

abolished so that the assignment of the species discussed here

may be changed.

The loreal is rather variable. It may be about as large as the

preocular, in which case it meets supralabials two and three.

It may be smaller and may meet only supralabial two. It may

be absent allowing the prefrontals to meet the supralabials, or

the nasal to meet the preocular (as a unilateral variant, C.

clelia BMNH 89.4.8.2). A large loreal is assumed to be

primitive.

Boulenger (1896) distinguishes between those Oxyrhopus

(s.1.) in which the preocular reaches the upper surface of the

head and those in which it does not. the distinction appears to

be real but it can be influenced by the condition of the

specimen and by the angle of view. The supraocular scale

meets the preocular and often, also, the prefrontal scale. The

lengths of the supraocular-preocular and the supraocular-

prefrontal sutures are compared. The prefrontal suture may

range from two or three times the length of the preocular

suture, as in Clelia rustica, to absent, as in Oxyrhopus petola

in which the preocular meets the frontal.

There are one preocular and two postoculars. The tempo-

rals are 2:3. The upper anterior temporal always meets

postoculars; the lower may meet them (represented by

+/+2:3) or may not meet them (+/—2:3). In every species

represented by more than two specimens this contact is

variable. Where the temporal does not make contact the

labial scale is higher than its neighbours. However, rustica is

nearly constant, on only one side of one of nine specimens

does the lower temporal make contact (BMNH 95.9.17.21).

The outgroup is also variable so polarity is uncertain. In one

rustica there is no suture between prefrontal and preocular

(BMNH 86.1.19.21).

There may be three or two anterior supralabials. As three

is the commonest number in the outgroup it is assumed to be

primitive. There are two supralabials meeting the eye and

three postocular supralabials. Anterior and posterior infrala-

bials are distinguished. The last anterior infralabial is pentag-

onal; from it starts the posterior row and a mesial row. The

anterior infralabials are usually five. Four and six occur as

unilateral variants; one rustica (BMNH 1909.11.2.16) has

four symmetrically. The posterior infralabials range from

four to two. As four is the commonest number in the

outgroup this is assumed to be primitive.

The genials range from posterior about as large as the

anterior to somewhat smaller. This does not appear to be a

taxonomically useful feature.

There are always two, sometimes three, scale row fusions

from the back of the head onto the neck. The rows on the

neck may be 19 or 17. As a majority of the outgroup have 19

this is inferred primitive. In those with 19 rows this number

G. UNDERWOOD

continues through mid-body and then reduces by fusion of

row four or five with the row below to 17 rows. The vertebral

or paravertebral rows were not seen to be involved in

scale-row changes. Those with 17 neck rows may continue

without change to the end of the trunk. In some specimens,

however, a lateral row splits on the posterior neck to give rise

to 19 rows through mid-body, then reducing to 17 rows. Such

a minimum on the neck rising to a maximum around mid-

body and falling towards the vent is widespread in henophid-

ian grade snakes (Underwood and Stimson, 1990). However,

this condition is found in none of the outgroup so it is here

assumed to be a derived, pseudoprimitive feature. The scale-

row pattern is scored as 19:19:17 (inferred primitive),

IVES EI Core ATR ibzei ly

In some specimens the vertebral scale-row is undifferenti-

ated, but in many it is wider than the paravertebral rows. This

modification is a little more pronounced towards the poste-

rior trunk. As most of the outgroup have undifferentiated

vertebral scales this is assumed to be primitive.

These snakes usually have both scale pits and tubercles

(Underwood, 1963). The pits are confined to the apices of the

scales of the trunk and tail, where they usually occur in

pairs. Tubercles may be numerous on the head scales. On the

trunk they are rather irregularly distributed around the centre

of each scale; numbers range from five to zero (Fig. 2). Both

pits and tubercles tend to be better developed on the upper

scale-rows. Where the vertebral scale-row is enlarged

they may be reduced or missing. Pits and tubercles are

found in the outgroup and their presence is inferred primi-

tive.

The anterior viscera were examined and their positions

recorded in ventral scale units. This has the advantage that

juveniles and adults can be compared. Organ positions deter-

mined by measurement are subject to allometric change

(Thorpe, 1975). The features recorded are: tip of the hyoid,

tip of the ventricle, anterior end of the liver and the end of

the tracheal cartilages. In all the entry of the trachea into the

right lung is terminal. This is a derived condition which

Wallach (personal communication) reports to be general in

Xenodontine snakes. In all there is some extension of the

vascularisation of the lung into the roof of the trachea, but it

does not extend far forwards of the heart. The left lung may

be present but small, up to about one or two scale-units long

and vascular; it may be a non-vascular vestige or it may be

absent. Presence is inferred primitive.

The trachea may terminate no more than three or four

scale-units beyond the tip of the heart. It may extend to

overlap the liver, in some cases extending the full length of

the vascular portion of the lung to reach the terminal air-sac.

A short trachea is inferred primitive.

Counts are made of the teeth, and empty tooth places, of

the left maxilla. The solid teeth show a moderate increase in

size from front to rear. They are followed by an interval and

two obliquely placed teeth with anterior grooves (Fig. 3).

Polarity is not inferred. In one juvenile specimen (clelia,

BMNH 1933.6.24.102) no grooves could be seen, even when

the maxilla was removed and dried. The anterior tooth count

is recorded. For sample specimens counts were made of the

teeth on the dentary, palatine and pterygoid bones. They all

have a full length choanal process of the palatine bone with a

broad base, about half the length of the palatine, which

sweeps backwards into a process which overlaps the ptery-

goid bone by two to three teeth (Fig. 3). The maxillary

process of the palatine is turned backwards; it bears a

A NEW SNAKE FROM ST LUCIA

foramen for the maxillary nerve which emerges on the

underside anteriorly.

For sample specimens the skin on the side of the neck was

turned forwards to expose the superficial jaw muscles and

ligaments and the labial glands. The most superficial muscle,

which is easily damaged, is the constrictor colli (Haas, 1973).

In the outgroup this is a thin sheet of muscle which passes

from about the level of the head of the quadrate backwards

and downwards over the jaw articulation to insert on the skin

of the throat. In the species of Clelia examined the muscle

follows a similar course, the anterior portion has a diffuse

origin on the surface of the adductor externus profundus

muscle. The posterior portion arises on the head of the

quadrate; this is inferred derived (Fig. 4).

The cervico-mandibularis muscle arises from the back of

the neck and passes downwards and forwards to insert on the

articular head of the quadrate (Fig. 4). This appears to be a

primitive condition. From the articular head of the quadrate

arises a ligament which passes forwards and divides. The

lateral, labial portion inserts on the posterior supralabial

scales and onto the capsule of Duvernoy’s gland. It peels off

the supralabial scales rather more easily than is usual in

snakes; this is thought to be derived. The mesial, maxillary

ligament passes forwards to insert on the posterior lateral

corner of the maxilla.

Fig. 1.

B, dorsal view of head.

Mucous supralabial glands extend along the margin of the

upper lip from the corner of the mouth to the snout. There

are similar glands along the margin of the lower jaw. Mesial

to the three posterior supralabial scales lies the Duvernoy’s

(venom) gland (Fig. 4).

At the level of the corner of the mouth, mesial to the

maxillary ligament, is the organ termed anterior temporal

gland by Smith & Bellairs (1947) and rictal gland by McDow-

ell (1986). It is found in all of the species examined; it is

usually visible mesial to the posterior end of the Duvernoy’s

gland (Fig. 4).

RESULTS

Clelia errabunda sp. nov.

DIAGNOsIs. A species of Clelia with uniform dark grey adult

colouration of the upperside of the head, trunk and tail,

extending to the lateral margins of the ventral scales, an

undifferentiated vertebral scale row, no left lung and a short

trachea extending no more than five ventral scale units

beyond the tip of the ventricle. Distinguished from rustica

and bicolor by absence of a left lung. Distinguished from

clelia, equatoriana and scytalina by the undifferentiated verte-

bral scale-row and short trachea. Further distinguished from

clelia by 17 scale-rows from neck to vent.

Holotype: BMNH 89.8.14.25, male, St Lucia, West Indies,

collected by G.A. Ramage, presented by West Indies Explo-

ration Committee; snout-vent c.112 cms, tail 32 cms with

extreme tip missing.

Paratype: MNHP 7598, male, St Lucia;

c.116 cms, tail 29+ cms with tip missing.

Referred specimens: BMNH 89.8.14.12, female, ‘Domin-

ica’, West Indies, collected by G.A. Ramage, presented by

West Indies Exploration Committee, snout-vent 138 cms, tail

27.6 cms.

BMNH 1988.717, female, ‘Demerara’, presented by Capt

E. Sabine, R.E., snout-vent 117 cms, much of the tail is

missing.

The other species of Clelia are widely distributed on the

South and Central American mainland and a few offshore

islands (Bailey, 1970). The name is taken from the Latin

errabundus = wandering, in reference to the occurence of

the new form well outside the range of its mainland relatives.

snout-vent

Clelia errabunda sp. nov., type BMNH 89.8.14.25. A, lateral view of head (reversed on account of distortion of right side of head);

The type has two preocular scales on one side, seen in no

other specimen of Clelia. All have two anterior and three

posterior temporals. The lower anterior temporal scale meets

a postocular in the type only. The ‘Demerara’ specimen has

three anterior supralabial scales on the left-hand side, seen

elsewhere only in C. bicolor. All have five anterior infralabial

scales and three posterior. The anterior genials are little, if at

all, larger than the posterior. The four specimens have

tubercles but no pits on the head, as in other Clelia and

Pseudoboa. The number of frontal and parietal tubercles is

high. Most of the trunk scales bear paired apical pits, as is

usual in Clelia, and a moderate number of tubercles (Table 2,

see Appendix).

The island specimens have 14 anterior maxillary teeth (on

the left), which is higher than for clelia and equatoriana

(Table 3, see Appendix); the ‘Demerara’ specimen has 13/12.

The ventral scale counts are high for Clelia, but not extreme.

The subcaudal scales are entirely paired, save that the last

one is single in the ‘Dominica’ specimen. Apart from the

difference of sex the three island specimens are very similar.

G. UNDERWOOD

KIKI

Fig. 2.

Clelia errabunda sp. nov., mounted scales to show distribution of tubercles and pits. a,b, parietal and frontal of BMNH 89.8.14.12;

c,d, frontal and parietal of BMNH 89.8.14.25 (type); e, mid-trunk scales of BMNH 89.8.14.12, from rows: 4, 5, 6, 7, paravertebral and

vertebral. The lower rows lack scale-organs.

DE dt ddite. baehtbnn

Fig. 3. Clelia errabunda, BMNH 85.8.14.12. i, mesial view of lower law: a = angular, d = dentary, s = splenial; ii, lateral view of left

maxilla (reversed); ii, ventral view of left upper jawbones: e = ectopterygoid, f = fangs with grooves, m = maxilla, pa = palatine,

pt = pterygoid; iv, mesial view of left palatine-pterygoid articulation: cp = choanal process, pp = posterior process of palatine.

The ‘Dominica’ female has 234 ventrals and 71 subcaudals:

total 305. The type male has the extreme tip of the tail

missing, judged to be not more than two pairs of subcaudals.

It has 221 ventrals and 82 + (?)2 subcaudals: total

303 + (?)2. The paratype male has 224 ventrals and 75+

subcaudals. The ‘Demerara’ female has 230.5 ventrals and

only 36 remaining pairs of subcaudals.

The ‘Dominica’ specimen has a Duvernoy’s (venom) gland

from behind the eye to the corner of the mouth; it is as high as

the supralabial scales plus the lower temporal scales. The

hemipenes of the two males are 18 subcaudal scale units long,

there are prominent lobes on a shaft 13 units long. The sulcus

spermaticus forks on the shaft at scale six (type) or seven

(paratype). Proximally on the shaft there are very fine spines

and, in the retracted organ, longitudinal folds. From scale

nine to the cleft there are large spines, about 26 in the type

and 38 in the paratype; these are high counts for Clelia. At

the base of each lobe there is a large spine, as is usual in

Clelia. the branch sulcus passes down the middle of an area of

large calyces with a clear margin (a capitulum).

Inspection of The Natural History Museum register raises a

doubt about the provenance of the Dominica specimen. G.A.

Ramage brought back a collection of herpetological speci-

mens from Dominica and St Lucia. These were registered in

1889. They are entered in Boulenger’s hand. The register

starts (with present identifications substituted):

89.8.14. 1-8 Typhlops dominicana Dominica, June 89

A NEW SNAKE FROM ST LUCIA

9-11 Alsophis antillensis Dominica, June ’89

sibonius

12 Clelia errabunda

13. Bothrops caribbaeus

14 Thecadactylus

rapicauda

Dominica, June ’89

St Lucia, April ’89

There follow another eight species attributed to St Lucia,

including the St Lucia endemics Hyla rubra, Sphaerodactylus

microlepis, Anolis luciae and Liophis ornatus. these St Lucia

attributions are not therefore in question. Boulenger’s St

Lucia entries are interrupted by four fish entries in a different

hand.

The Typhlops and Alsophis are forms endemic to Domin-

ica. What, however, attracts attention is the record of Both-

rops from Dominica. We may be sure that if a pit-viper were

living on this island there would have been further reports

since 1889. It is clear that the Bothrops was mistakenly

attributed to Dominica. The specimen of Clelia appears to be

the only documented record of the genus from Dominica. Is

this attribution to Dominica also a mistake? On the other

hand there are several further specimens of Clelia from St

Lucia in the Museum of Comparative Zoology.

In addition to Alsophis antillensis, there is no more than

hearsay evidence of a second species of black snake on

Dominica. Bullock & Evans (1988) list Clelia clelia on

Dominica as “Tete-chyen nwe’. Dr Bullock writes that he has

not seen Clelia but he has reports from informants whom he

regards as reliable. Dr R. Thorpe, Miss A. Malhotra and Mr

M. Day have come across no evidence of Clelia on the island.

On Dominica it would be distinguished from Alsophis by the

uniform black dorsal colouration and by 17 dorsal scale rows

on the anterior trunk. The A/sophis has some irregular yellow

markings and 19 scale rows anteriorly. Unless and until there

is clear evidence of the occurrence of Clelia on Dominica it

should be dropped from the island list. Barbour (1930) says of

‘Clelia clelia’ that ‘“‘This species is surely extinct on St

Lucia. . .”. Long (1974) writes that ‘“‘the cribo no longer

exists in St Lucia. . .”. Dr D. Corke also reports that he has

found no trace of the survival of Clelia on St Lucia.

Even greater doubts arise about the provenance of the

‘Demerara’ specimen of errabunda. The specimen had no

original registration number; the Museum register starts in

1837. A search of the early entries shows no record of

specimens from E. Sabine. The Museum archivist reports

that the trustees’ minutes record donations from Capt Sabine

between 1818 and 1824 with, however, no indication of any

from the Caribbean.

The collection has other snakes from ‘Edw. Sabine’. There

is a male Xenodon merremi, a species widely distributed in

South America and known from Guyana on the basis of other

specimens. There are a male and a female of Oxyrhopus

trigeminus, not otherwise known from this part of South

America.

There are also two lots of Bothrops. A female from ‘Capt

Sabine, Berbice’ and two females and a male from ‘Col

Sabine, Demerara’. These were compared with specimens of

B. atrox and B. brazili from Guyana and with B. caribbaeus

from St Lucia. With ventral and mid-body counts of: M

201:26, F 205:27, F 206:29 and F 210:29 they fall within the

range of counts reported by Lazell (1964) for caribbaeus. The

postocular stripe passes across the last supralabial scale dorsal

to the corner of the mouth as in caribbaeus. The ventral scales

are laterally peppered with dark spots, as in caribbaeus and

Fig. 4. Clelia errabunda sp. nov., BMNH 89.8.14.12. Dissection to

show superficial head muscles and glands (reversed).

Cc = constrictor colli muscle, overlaying other structures;

Cm = cervico-mandibular muscle; Dg = Duvernoy’s gland;

Hg = Harder’s gland; Ig = infralabial gland;

Q1 = quadrato-labial ligament; Sg = supralabial gland;

Rg = rictal gland.

brazili and unlike atrox. In all, the dorsal bands are indistinct.

In two of the Demerara specimens the bands can be seen to

have parallel sides or to converge towards the dorsal midline

as described by Lazell for caribbaeus. The third shows some

diverging bands. The Guyana atrox have dark patches on the

lower flanks which extend onto the ventrals unlike these

Sabine specimens. The brazili have dark bordered bands

which converge towards the midline. These observations

suggest that the ‘Berbice’ and ‘Demerara’ specimens are most

probably caribbaeus, a species known only from St Lucia.

The above considerations raise a doubt that Sabine collected

any specimens in Guyana.

It is evident that Sabine was long enough, supposedly in

Guyana, to achieve promotion from captain to colonel; in

that time he may well have visited St Lucia. The locality of

the ‘Demerara’ Clelia errabunda is therefore discounted. The

uniform 17 scale-rows and undifferentiated vertebrals would

distinguish it from local Clelia clelia.

Dumeril, Bibron & Dumeril (1854) report that the Paris

museum has specimens of ‘Brachyruton cloelia’ from Guy-

ana, Brazil, Mexico and Guadeloupe. The ‘Guadeloupe’

specimen (MNHP 169) is a hatchling with counts V200, C83

and 19:19:19 scale rows. The tip of the heart is at V46, the

trachea extends beyond V90. This is clearly a specimen of

Clelia clelia, with an unusually low ventral count. The locality

is undoubtedly erroneous.

In the Proceedings of the Philadelphia Academy for 1870 it

is reported that Cope “‘called attention to a large specimen of

Trigonocephalus (= Bothrops) from St Lucia, of which some

fourteen inches was enclosed in the oesophagus and stomach

of a larger Oxyrhopus plumbeus (= Clelia clelia).” Later

Cope (1876) wrote that, as he had previously observed, he

had received a specimen of Clelia clelia from Martinique(!)

which had swallowed a large Bothrops. Malnate (personal

communication) examined the specimen(s). It is ANSP 10220

from ‘Santa Cruz’, received from Mrs J.L. Endicott. ‘Santa

Cruz’ presumably means St Croix in the Virgin Islands! It is

unlikely to be a St Lucia locality; most of the place names are

French. This one specimen thus has three different geograph-

ical attributions!

Malnate reports that the Clelia has 17 scale-rows through-

out and an undifferentiated vertebral scale row; it therefore

fits C. errabunda. The half-swallowed Bothrops has 25 scale

rows about midbody, falling to 19 rows. Lazell (1964) gives

mid-body scale-row counts for Bothrops caribbaeus from St

Lucia as 25-29 (mode 27) and for Bothrops lanceolatus from

Martinique counts from 29-33 (mode 31). The Philadelphia

Academy specimen, with 25 rows, is at the lower end of the

range for St Lucia specimens. Beyond reasonable doubt

therefore Cope had a specimen of C. errabunda which had

half swallowed a St Lucia Bothrops caribbaeus.

Relationships of Clelia errabunda

The species currently assigned to Clelia can be arranged at

several levels on the basis of the derived states of the

respiratory system, the vertebral scale row and the ventral

scale counts. This is set out in Table 3.

C. bicolor. The three anterior supralabials, the low ven-

tral scale counts, the undifferentiated vertebral scales, the

presence of a left lung and a short trachea are primitive

features. The high maxillary tooth count appears to be a

derived feature.

C. rustica. This species also has undifferentiated verte-

bral scales, a left lung (a mere vestige in BMNH 81.7.2.9) and

a short trachea. C. rustica and the following species are

derived in relation to bicolor in respect of two anterior

supralabials and higher ventral counts.

C. errabunda, clelia, equatoriana and scytalina share the

derived feature of absent left lung. The latter three species

are further derived than errabunda in respect of the enlarged

vertebral scales and extended trachea.

The type specimen of Clelia clelia Daudin (1803) is not

known to survive; however, the type locality is given as

‘Suriname’. It is therefore assumed that specimens from the

northern coast of South America are typical clelia. Specimens

in The Natural History Museum from this area, from Central

America, from Rio Condoto on the Pacific slope of Colombia

G. UNDERWOOD

(1), from La Paloma nr Santiago R., Ecuador (1) and from

most of the rest of South America show the 19:19:17 scale

row pattern. However, a second specimen from Rio Condoto

and specimens from Ecuador (Guayaquil and east of Loja,

2), Peru (3) and Manacapuro on the Amazon (1) show the

17:19:17 scale-row pattern. These are indistinguishable from

typical clelia in respect of the other characters considered

here. Their occurrence in a north-western area of South

America with near overlap with the 19:19:17 (Rio Condoto)

form on the Pacific slope of Colombia does not look like an

accident of sampling (Fig. 5). The form in eastern Brazil

(plumbea) lacks spines on the hemipenis, some specimens

from the southern part of the range (occipitolutea) are pale in

colour, so clelia is evidently a variable species. Roze (1959)

reports a specimen from Venezuela with counts of:

21:22:19:17 and Chippaux (1986) reports a specimen with

21:19:17 rows from French Guyana.

After the above account was prepared I received from

Zaher (personal communication) a photocopy of a portion of

Bailey’s unpublished PhD thesis. It is evident that at that time

he regarded the island Clelia as sufficiently distinct to merit

subspecific status. He too did not believe the Guyana locali-

ties of the Sabine specimens.

DISCUSSION

The species clelia, equatoriana, scytalina and errabunda share

uniform dark adult colouration and absence of a small left

lung. Most other pseudoboine snakes have a small left lung

and a more varied colour pattern. Within this group of four

species errabunda is primitive to the others in that the

vertebral scale row is not enlarged and the trachea is short.

Fig. 5. Localities of specimens from northern South America and the Lesser Antilles. Solid symbols = precise localities; hollow

symbols = approximate localities; circles = Clelia clelia; triangle = Clelia errabunda; 17, 19 = no of scale rows on neck of C. clelia.

B = Barbados, D = Dominica, Gr = Grenada, Gu = Guadeloupe, L = St Lucia, M = Martinique, V = St Vincent.

A NEW SNAKE FROM ST LUCIA

Because the 19:19:17 scale row pattern is widespread in

pseudoboine snakes and is also found in most clelia we may

infer that it was the condition of the ancestor of this species

group. The 17:17:17 pattern shown by errabunda would thus

be interpreted as a derived feature setting it apart. However,

the occurrence of the 17:17:17 pattern in equatoriana and

scytalina and the 17:19:17 and 21:19:17 patterns within clelia

suggests that little significance can be attached to the scale-

row pattern. The most nearly special feature of the new

species is the high number of large spines on the hemipenis.

Otherwise it is close to the status of what Ackery and

Vane-Wright (1984) call a ‘paraspecies’, without any special

feature setting it apart. The short trachea and unmodified

vertebral scales by which it is distinguished are primitive

features found in hundreds of other species of snakes.

Greer (1965) reports that the Grenada Clelia is diurnal,

unlike its mainland relatives. Clelia clelia from Grenada is

otherwise little different from mainland clelia; this is con-

firmed by Wallach’s report (personal communication) that it

has an extended trachea. It is presumably a relatively recent

immigrant from South America. On the other hand erra-

bunda, on St Lucia, is primitive to the mainland members of

the clelia group. This suggests that it colonised St Lucia at an

early date and that its ancestral stock was later replaced on

the mainland by the more derived c/elia. It is an example of a

primitive form surviving as an island relict.

Boa constrictor occurs on St Lucia and Dominica. The two

island populations and the mainland form are well differenti-

ated from one another and are recognised as separate subspe-

cies (Lazell, 1964). The pit-vipers, Bothrops, on the adjacent

islands of St Lucia and Martinique are sufficiently differenti-

ated that they are recognised as full species by Lazell (1964).

For both Boa and Bothrops this suggests either separate

colonisation of the islands from the mainland or colonisation

of one island and passage to the other long ago. There is at

present no evidence that these are primitive island relicts.

Cope (1870) is reported as saying that the “islands of

Martinique and Guadeloupe had become so infested with the

fer-de-lance” (Bothrops lanceolatus) ‘“‘as to be in parts almost

uninhabitable, and it was chiefly on account of the danger

from this venomous reptile that collecting naturalists of late

years had so seldom visited them’’! ““Some means had been

adopted to check the increase of this pest, but with small

results”. “Prof Cope thought that as the Oxyrhopus

plumbeus (= Clelia clelia) was very numerous in Venezuela

and Brazil, and since it was very harmless and easily pro-

cured, that its introduction in large numbers into Martinique,

etc, would be a simple matter, and one probably to be

attended with good results in the diminution, at least, of this

enemy of agriculture”’.

Lazell (1964) tells us that on both Martinique and St Lucia

the local Bothrops is known as ‘serpent’. We may speculate

that prior to human arrival ‘serpents’ were already estab-

lished on St Lucia before the ‘cribo’ (Clelia) arrived to prey

upon them.

It is said that the mongoose was introduced into St Lucia in

the hope that it would reduce the Bothrops. Today, although

the mongoose may eat Bothrops, it also eats domestic poul-

try. Following human disturbance, it is ironic that the indige-

nous ‘pest’, the ‘serpent’, is supplemented by an introduced

pest, the mongoose, and the indigenous biological control,

the ‘cribo’, is extinct! In the absence of this ‘control’ Lazell

(1964) reports that in some areas of St Lucia the serpent is

‘abundant beyond belief’.

ACKNOWLEDGEMENTS. I am indebted to Van Wallach for a report on

a Grenada specimen of Clelia clelia and for information about other

species of Clelia and other xenodontine snakes. Beat Schatti loaned a

specimen of Clelia equatoriana for examination of the anterior

viscera. Ed Malnate reported on the Philadelphia Academy speci-

men(s). Robert Henderson, David Bullock and Roger Thorpe

replied to a query about the status of Clelia on Dominica. David

Corke commented on the status of Clelia on St Lucia. Hussan Zaher

drew my attention to the work of Scrocchi and Vinas, identified the

Oxyrhopus trigeminus from ‘Guyana’, located specimens in the Paris

Museum and gave me a photocopy of a portion of Bailey’s unpub-

lished PhD thesis. T.E. Pickring, archivist at The Natural History

Museum, traced the early records of Capt Sabine. Colin McCarthy

helped me to find my way through The Natural History Museum

records and read a first draft of this paper. The Museum national

d’Histoire naturelle, Paris, loaned two specimens.

REFERENCES

Ackery, P.R. & Vane-Wright, R.I. 1984. Milkweed butterflies, their Cladistics

and Biology. British Museum (Natural History), London.

Barbour, T. 1930. A list of Antillean reptiles and amphibians. Zoologica, 11:

61-116.

—— 1935. Second list of Antillean reptiles and amphibians. Zoologica, 19:

77-141.

—— 1937. Third list of Antillean reptiles and amphibians. Bulletin of the

Museum of comparative Zoology 82: 1-166.

Bullock, D.J. & Evans, P.G.H. 1988. The distribution, density and biomass of

terrestrial reptiles in Dominica, West Indies. Journal of Zoology, 222:

421-443.

Boulenger, G.A. 1896. Catalogue of the Snakes in the British Museum, Vol. 3,

part A.

Cadle, J.E. 1984. Molecular systematics of Neotropical Xenodontine snakes: I

South American Xenodontines. Herpetologica, 40: 8-20.

Chippaux, J-P. 1986. Les serpents de la Guyane frangaise. Editions de

l’Orstom, Institut frangais de recherche scientifique pour le developpment en

cooperation. Collection fauna tropicale, no XXVII, Paris.

Cope, E.D. 1870. ‘Verbal communication’ Aug 2nd. Jn: Proceedings of the

Academy of Natural Sciences of Philadelphia, p. 90.

1876. On the Batrachia and Reptilia of Costa Rica. Journal of the

Academy of Natural Sciences, Philadelphia, 8(2), p. 131.

Dowling, H.G. 1951. A proposed standard system of counting ventrals in

snakes. British Journal of Herpetology, 1: 97-99.

Dumeril, A.-M.-C., Bibron, G. & Dumeril, A. 1854. Erpétologie générale,

Paris. Vol. VII, part II, p. 1007.

Greer, A. 1965. A new subspecies of Clelia clelia (Serpentes, Colubridae) from

the island of Grenada. Breviora, 223: 1-6.

Haas, G. 1973. Muscles of the jaws and associated structures in the Rhyn-

chocephalia and Squamata. Jn: Gans, C. & Parsons, T.S. (eds) Biology of the

Reptiles. Academic Press, London.

Lazell, J.D. 1964. The Lesser Antillean representatives of Bothrops and

Constrictor. Bulletin of the Museum of comparative Zoology, 132 (3):

245-273.

Long, E. 1974. The serpent’s tale. U.W.1. Extra mural department, P.O. Box

306, The Morne, St Lucia.

McDowell, S.B. 1986. The architecture of the corner of the mouth of colubroid

snakes. Journal of Herpetology, 20: 353-407.

Peters, J.A. & Orejas-Miranda, B. 1970. Catalogue of the Neotropical Squa-

mata: Part I, Snakes. United States national Museum Bulletin, 297, Washing-

ton.

Roze, J.A. 1959. Taxonomic notes on a collection of Venezuelan reptiles in the

American Museum of Natural History. American Museum Novitates, No

1934: 1-14.

Schwartz, A. & Henderson, R.W. 1988. West Indian amphibians and reptiles: a

checklist. Milwaukee Public Museum, Bulletin 74, Wisconsin.

Scrocchi, G. & Vinas, M. 1990. El genero Clelia (Serpentes: Colubridae) en la

Republica Argentina: revision y comentarios. Bolletino del Museo regional

dei Scienze naturale di Torino, 8: 487-499.

Smith, M.A. 1943. Fauna of British India, Reptilia and Amphibia, Vol. 3,

Serpentes, Taylor & Francis, London.

Smith, M.A. & Bellairs, A.d’A. 1947. The head glands of snakes, with remarks

on the evolution of the parotid gland and teeth of the Opisthoglypha. Journal

[oe]

of the Linnaean Society, Zoology, 61: 351-368.

Thorpe, R.S. 1975. Quantitative handling of characters useful in snake

systematics with particular reference to intraspecific variation in the ringed

snake Natrix natrix (L.) Biological Journal of the Linnaean Society, 7: 27-43.

Underwood, G. 1963. A contribution to the classification of snakes. British

Museum (Natural History), London.

G. UNDERWOOD

Underwood, G. & Stimson, A.F. 1990. A classification of pythons. Journal of

Zoology, 221: 565-603.

Vanzolini, P.E. 1986. Addenda and corrigenda to the catalogue of neotropical

Squamata. Smithsonian herpetological information service, No 70, Washing-

ton.

APPENDIX

Table 1 Some representative specimens of Clelia, showing: sex, ventrals, scale row reduction pattern, scale row stretches on tail. * = type

specimen of Oxyrhopus maculatus Boulenger.

Vv Row reductions

scytalina

68.4.7.7 M 212 >>19(12;5>)17

68.4.7.8 F 209 >>>19(7;5>)17

equatoriana

60.6.16.47 M 201 >>17

Geneva 2410.9 M 201 SSy/

60.6.16.48 F 217 >>17

60.6016.49 F 219 >>17

clelia

74.8.4.56 M 216 >>17(44;5<)19(149;5>)17

90.10.6.29 M 214 >>>17(41;4<)19(146;5>)17

1926.4.30.14 F 238 >>17(43;4<)19(159;4>)17

51.7.17.136 M 219 >19(165;4>)17

86.10.4.12 M 213 >>19(165;4>)17

1929.10.19.2 EF 237 >19(209;5>)17

1902.7.29.68 F 231 >>19(142;6>)17

*84.2.23.40 F 213 >>19(179;5>)17

errabunda

89.8.14.25 M 221 >>17

MHNP 7598 M 224 >>17

89.8.14.12 F 234 >>17

1988.717 F 230.5 >>(7;4>)17

rustica

86.1.19.21 M 206 >>19(176;5>)17

81.7.2.9 M 196 >>>19(129;4>)17

95.9.17.21 F 212 >>19(177;4>)17

1933.9.5.7 F 195 >>19(128;4>)17

bicolor

1927.8.1.234 F 178 19(96;5>)17

1980.1651 F 177 19(108;4>)17

Tail row stretches

Cc >8 8 6 4 >

75 15 iB 20 25 4

83 Bl 14 22 30 6

75 5 17 23 30 0

69 4 14 18 28 5

x 3 12 20 = =

- 5 12 21 = =

91 10 19 26 26 10

71 6 20 27 18 0

82 2 8 24 26 12

91 6 20 24 33 8

64 12 13 19 17 3

84 3 14 32 30 5

55 3 7 25 20 0

50 3 11 20 19 0

84 ll 15 22 34 0

75+ 15 14 21 25+ is

71 5 8 a, 31 0

= 4 ul . = =

61 14 22 16 9 0

60 6 23 17 16 0

55 7 17 19 12 0

39 3 9 21 6 0

59 3 15 15 21 5

58 3 12 16 23 4

A NEW SNAKE FROM ST LUCIA

Table 2 Distribution of tubercles on the dorsal, frontal and parietal scales of some selected specimens of Clelia.

Dorsal scale-row nos.

1 2 aie CA ee 16, eT GB eSs. 9 10 » fr par

scytalina 68.4.7.7 M- - 4 4 20° i 3.4 117 98

equatoriana 60.6.16.47 Mi ge = OI eit 2 Slee £2 129 98

clelia 19:19:17 1930.10.10.188 Mi) Sp a =e, es ee eee 3 2 151 104+

clelia 19:19:17 94.3.14.60 Be oe ely, Lt eer 35 4 96 87

clelia 17:19:17 89.4.8.2 M2 3. = 2 8), eS 477 1G 6 150 112

errabunda St L. 89.8.14.25 Mae = = = SS 2 = = 138 92

errabunda St L. MNHP 7598 M = (S - Se ee ly 2-4=5 = 200 104

errabunda ‘Dominica’ 89.8.14.12 Bh ete ee he 186 152

errabunda ‘Demerara’ 1988.719 F=- - - 1 te a 2 141 132

rustica 1909.11.2.16 M- - = = = = = = = = 72 45+

bicolor 1980.1651 Bos Ss = ose see eo oe al 1 45 34

Table 3 Comparison of Clelia species.

Max Vert.

n suplabs Vv cE teeth L.lung Trachea TOW

scytalina F 1 22:3 209 83 13 ae a +

M 1 212 75 14

equatoriana Bae 2A2:3 217-219 - (1G) =F + se

M 2 201 72 12

clelia Bs 252-3 213-228-238 50-73-84 1S 35 + ar

M 13 204-215-226 64-82-91 iW

errabunda Fo 2 2:2:3 231-234 71 14 iF - -

M 2 221-224 84 Sed

rustica | Sogpeai 22-3 195-208-231 39-44-55 11.3 - - -

M 2 195-210-223 61 13.0

bicolor 32 322:3 177-178 58-59 15:5 - - -

V = ventrals, C = subcaudals, min-mean-max; mean no of anterior maxillary teeth; - = primitive, + = derived state of left lung, trachea and vertebral scale-row.

Bull. nat. Hist. Mus. (Zool.) 59(1): 11-31

Issued 24 June 1993

Anatomy of the Melanonidae (Teleostei:

Gadiformes), with comments on its

phylogenetic relationships

GORDON J. HOWES

Department of Zoology, The Natural History Museum, Cromwell Rd, London SW7 5BD

CONTENTS

REE ORL ITE ELOORUB Tae ee aetse aa sae eer aR aerate ee acl rc clea e'ds wales oun vv nitied we ve vis ssa slag taubinn Ce oe cee t ome teen e ER eon tenet tenronm heameedereeere 11

METEENIAScANGUIMETNOUS: jnesavicaeschns deosmpmare Aatescoa ds u8th satu de ddveadse Sona the tects btee tad aocetrok aa tiae centers he ees 12

PRESLE NATIONS ESE CITI MIe a TT OUIT EG acts sete scien qeicic'sstte a visisle selie aisticerss nnrterit ares abioinais's aolneeeiala oer Mem aelcebroeelauie te adeelonelsusinnsse 12

HBTS ITN age = SBME SREB AE 4 once COME RPESE TCO" SEEICHR SREP Soe REC? SOnC HEA EET oMRRCC BERGE Cnr Mac RRCHon SRRRCARR TC Hee RnmOe scree 13

IN CHOI TD ALLEL) .seeemerceras sid vptta rete vin esasidayls Grp catglia'a dein a's u's o's ue 5 asis aap’n Mais sc as fase inte nn eo aalealaae denis adas sdne aang cide 13

STULL SECA S taconite ae pee nee ct aso ances spine te ve siete a sane anepiasioas anideiabinn ee wctecge Tales ap seeasees “ies Relateeeaa See 14

ROE ATE ERP eee rene ee een as Aaiicrs asi sie garnd Wiuts si'cians tieaine anes maapaivamannasitee deiceboetiooess dae teot etic aeaetan inns 16

RADU S meee ech cots. diame ta cacme atdma aaa sang asiela s veincls eeicio oa wmeeeay vadigicles djemantamhiencmansteaceit esp msaiacdees manatee meshes 19

ALA LOI POL RAC ALE Cte tee tate oe Maca ne occid na case ques ntclesiigt cen ares veers same eeeeeh order cartaacodes-Toerecsieteaaennte eee 20

INOIGEAN CMM Minera reccercreortttattte tence cecaietrceescancuccaccescngterscasccacerecoudvetatbeitererescterostcs tometntendheresortasntcs 21

Oster eTUN IF LOGIE ser. donc de yop cOdb Ub OTE NDUEEOEDAEC EODHOAGBSRGRERERED cunehia: pobrnade- oqode oncoctc rode angrocreecdadciica ieanbiscemeionapchee

ESEAIICIMAMALGMES cs de etec atthe ce ch ae tet eons case als od oe one ool er eee eee TUR Ree EAE Eee te PE A Oe 22

BGBLORAMCIKCI Ci rete. iat tete ccatiees sutiee sec vcessdentonvevedecovesnsdceawedens totes soaMR aE a Tee aE E eee Sc eR ER RAR eon cane n aE au 23

BOC PITS Met. theca an tBcterc dete ce cest a reiaues sun sera Wisten sa ne cacbantee none eetedcemeneeeree nee ta dadeRtichactatewctuet oes ae ines 24

EHEC OLAl COLMA ANGNIG CIAL TINS. 4.5375. out 3u LSS ode se won os = SOba de nose MMU te Re ab des SIaeEa eto dasodldeteaenes meee 25

DELERIUM Ree etree Cee. ct aaa e cs Rioseid oie fas toe ea RNC MINS won sun AIS ache Sclvice shh ts SAUMEREDER ES stein caabee sand avai eiaies aaeteeneees 3 PA

Sun Ac denavIScerarana OGY MUSCUlAture, | ......bigr- <0 des. tide sseveeastyeckitlle otbbime dds duis decade smeuk gees Geile. aanee 28

DORIS ELIS SIC EN Pee Mate See hs cick e o EEIAAS ctr wnclpah ovdelsigsiajacion deta dees on « dO RaReS eee MeL ae iaeaeea ay SAPUAUT eb aEM ARG oe o'SSEEIS TNT EEE 28

PRRTLOM LEC OTAERS yeh eapaaty teil tebe «6 cotanddguncdingsen bos «ue + dcbeengde-e Mibteden ecnbebddadteredvads. « hudeehiaeeespe irae» wal. oes 30

FUGLEKCLICES ous saa eae RRR o Tames ee embecb east «oc duousehmdlanwneacemebae ret peteenodtessmtaaren eee: matt ee mere an cette anand 30

Synopsis. The osteology and part of the soft anatomy of the gadiform family Melanonidae, represented by the

genus Melanonus Ginther, 1878, is described. Melanonus has several derived (autapomorphic) sensory features but

only three osteological ones. Although contained within the Gadiformes the family is excluded from both the

Macrouroidei and Gadoidei in lacking a modified palatine and enlarged intercalar and thus represents their

sister-group designated as the Melanonoidei. The Suborder Gadoidei now comprises two families of uncertain

phylogenetic affinity (Bathygadidae and Steindachneriidae) and two Superfamilies, Moridoidea and Gadoidea.

INTRODUCTION

The gadoid genus Melanonus Giinther 1878 contains two

species, M. zugmayeri Giinther, 1878 (Fig. 1), and M. gracilis

Norman, 1930, which together give a broad latitudinal,

circumglobal distribution (Cohen et al., 1990; Howes, 1991a).

Melanonus are relatively small fishes, the largest seen being

230mm total length and, oddly for gadoids, are meso-

bathypelagic (100-3000m). Outwardly, Melanonus resembles

a stomiatoid rather than a gadoid fish with its dark coloration,

large, strongly-toothed jaws and tapering body (Fig. 1).

Until Marshall (1965) recognised (without diagnosis) a

separate family for the genus, Melanonus had been consid-

ered to belong to the Moridae. Marshall (1965) and Marshall

& Cohen (1973) contended that Melanonus was the most

primitive gadiform (anacanthine) fish, a contention based on

the posterior position (at the forebrain) of the olfactory bulbs

and a relatively unmodified caudal fin skeleton. The features

Marshall & Cohen (1973) used to diagnose the Melanonidae

rested on soft anatomical features, viz. an elaborate system of

free ending neuromasts on the head and the corpus cerebelli

extending (forward) to the optic tectum.

Apart from a few observations on the caudal fin skeleton

and gill-arches and a description of its cranial muscles (see

below) the anatomy of Melanonus has never properly been

described. Gosline (1971) complained that ‘‘No account of

the osteology is available. By contrast the family Gadidae has

received more attention from anatomists than almost any

other family of fishes”. Despite these shortcomings several

assertions as to the phylogenetic position of the Melanonidae

have been made.

Rosen & Patterson (1969) cited Marshall (1965; 1966) to

the effect that Melanonus represents a primitive gadoid.

G.J. HOWES

DATTA TTT TTT TTT TTT TTT TTT TTT TL Pepe TT TAGTTATp TNT HenT HTT TATA TTTTATT

Melanonus zugmayeri specimen BMNH 1991.7.9:729-30, 200mm SL, lateral view.

Fig. 1.

Schwarzhans (1980; 1984) combined the Melanonidae and

Moridae (produced as a cladogram in Patterson & Rosen,

1989) and Cohen (1984) and Fahay & Markle (1984) also

suggested a relationship with the Moridae, again based on the

primitive arrangement of the caudal fin skeleton. Markle

(1989) revised his earlier views and placed the Melanonidae

near the base of his cladogram making it (with the exception

of the Ranicipitidae) the sister group of all other gadiforms.

Nolf & Steurbaut (1989) placed Melanonidae as an unre-

solved polychotomy with the Euclichthyidae, Macrouridae,

Moridae and other gadoids. Okamura (1989) omitted the

family from his gadoid classification but implied (p.137) on

the basis of similar anterior rib structure that Melanonus is

closely related to Merluccius. Howes (1989; 1990) also placed

Melanonidae in an unresolved polychotymy, with Stein-

dachneriidae, Bathygadidae and other gadoids. According to

Howes (1990, 1991a & b) the majority of gadoid families

form a monophyletic assemblage termed ‘supragadoids’,

characterized by complete fusion of the upper hypurals into a

single plate. The Macruronidae represent the plesiomorphic

lineage of this assemblage with the Gadidae and Merlucciidae

being the most derived families. The Melanonidae was

assigned with the Bathygadidae, Steindachnertidae, Moridae

and Euclichthyidae to the ‘infragadoids’ and in one scheme

(Howes, 1991b) in alternative positions, one as the sister

group to all gadoids excluding the Bathygadidae and Stein-

dachneriidae, the other as also excluding the Moridae. The

characters on which these phylogenetic positions were based

are, however, ambiguous (see Discussion) and like all previ-

ous studies have suffered from lack of anatomical information

about Melanonus. The following is an account of the osteol-

ogy and other soft anatomical features of Melanonus.

MATERIALS AND METHODS

Specimens used for anatomical descriptions (all from BMNH

collections): Melanonus zugmayeri Uncat. 230mm, ‘Discov-

eny sine 115505" 10°VILA9875™ 20°25%S"Ny 219-39: 5" Ww

T73—8zom; 19919.7°9:729=730; 220mm, “95mm” "SIE;

20°25.8'N-31.4’N, 19° 39.8"W-38.0'W, — 800-875m;

1991.7.9:731—733, cleared & stained, 66, 100, 130mm SL,

17°1.2'N, 19°57.8’W, 400-495m; 1987.1.21:595-596, 215mm

SL, dry skull prepared from 190mm SL, 49° 21.9’N, 11°51'W,

1090-1100m; 1987.1.21:597, 168mm SL, S.W. Bantry,

960-920m; 1981.3.16:377, 173mm SL, West Great Sole

Banks; 1987.1.21:598-601, 175, 187, 193mm SL, one speci-

men, 165mm SL (cleared and stained), 50°02’N, 11°22’W,

910-980m; 1930.1.12:943 (Holotype) 13°58’S, 11°43’E. Mel-

anonus gracilis 1887.12.7:22 (Holotype) 147mm SL, Antarc-

tic; 1930.1.12:934-936, 97, 140, 150mm SL, 46°56’S,

46°03’'W; 1988.11.4:13-20, 45, 49mm SL (cleared and

stained), 35°13’-34°57'S, 17°50’-17° 48’E; 1988.11.4:2,

145mm SL, 50°17.7’S, 18°40.9’E, 300-150m; Percopsis omis-

comayus 1973.3.20:46-8, 52mm (cleared and stained), 62mm

SL, Lac Henry, Quebec, Canada; Bregmaceros sp.

1957.12.2:5-12, 54mm SL (cleared and stained) Senegal;

Gaidropsarus mediterraneus Uncat. 122, 145mm (cleared and

stained), Seaton Point, England.

In addition, material listed in Howes (1988, 1992) and

Howes & Crimmen (1990) was re-examined.

Abbreviations used in the figures

aa anguloarticular

aap premaxillary articular process

ac actinost

afc anterior frontal crest

ap premaxillary ascending process

ar anterior (‘chopstick-like’) ribs

ard anal fin radial

asp autosphenotic

bb basibranchial (numbered)

bh basihyal

bl Baudelot’s ligament

bo basioccipital

boc basioccipital condyle

br branchiostegal ray

cb ceratobranchial (numbered)

cc cerebellar corpus

ccr cerebellar crest

cfc central frontal crest

cl cleithrum

cm coronomeckelian bone

co coracoid

de dentary

dex dorsal section of epaxialis muscle

dhy dorsohyal

dr dorsal fin ray

ANATOMY OF THE MELANONIDAE

drd dorsal fin radial

eb epibranchial (numbered)

ebt epibranchial toothplate

ect ectopterygoid

edd erector and depressor dorsalis muscles

ent entopterygoid

ep epural

epr epineural

esc extrascapular

epo epioccipital

exc exoccipital condyle

exca exoccipital cartilage

exf exoccipital flange

exo exoccipital

f1X foramen for glossopharyngeal nerve

{X foramen for vagus nerve

fl facial lobe

fm foramen magnum

foc foramen for occipital nerves

fr frontal

fv flexor ventralis muscle

fvi flexor ventralis inferioris muscle

ge granular eminence

gg gas-gland

go gonad

gu gut

ha haemal arch

hb hypobranchial (numbered)

hf hyomandibular fossa

hp hypophysis

hy hypural (numbered)

iac interarcual cartilage

ic intercalar

ih interhyal

io infraorbitals (numbered)

iop interopercle

ird interradialis muscle

ki kidney

le lateral ethmoid

li liver

Ilp lateral ethmoid-palatine ligament

Imi mandibular-interopercular ligament

It trigeminal lobe

mc Meckel’s cartilage

md mesonephric duct

mec mesethmoid cartilage

met metapterygoid

mss myoseptal strands supporting ribs

mo medulla oblongata

nI-VII cranial nerves

na nasal

nal first neural arch

naap nerve branch serving adductor arcus palatini muscle

nau acoustic nerve

nbuc buccalis branch of trigeminal trunk

nio infraorbital branch of trigeminal trunk

nil lateral line nerve

nml,2 neuromasts types 1 and 2

nom nerve of supraorbital branch innervating posterior canal

enclosed neuromast

nr nasal rosette

nsab supraorbital branch of trigeminal

ob olfactory bulb

op opercle

ot olfactory tract

pa parietal

pah parhypural

pal palatine

pb pelvic bone

pbb pharyngobranchial (numbered)

pe postcleithrum

pfc posterior (diagonal) frontal crest

phy posterohyal

pmp postmaxillary process of premaxilla

pop preopercle

pp parapophysis

prn prootic notch

ps parasphenoid

psl parasphenoid ascending laminae

pte pterotic

pts pterosphenoid

ptt posttemporal

pu preural vertebra (numbered)

pyc pyloric caeca

qu quadrate

ra retroarticular

Rel ramus canalis lateralis nerve

rd retractor dorsalis muscle

re rostrodermosupraethmoid

sb swimbladder

sbp swimbladder pocket

sc scapular

sca supracarinalis anterior muscle

scl supracleithrum

so supraoccipital

sop subopercle

spt intercalar socket for posttemporal limb

st stomach

sy symplectic

ul ural centrum

7 vertebra (numbered)

vex ventral section of epaxialis muscle

vhy ventrohyal

vo vomer

ANATOMY

Neuromast pattern (Fig. 2).

Melanonus has a unique pattern of free-ending neuromasts

covering the head in addition to those more usual neuromasts

contained in the sensory canals. There are two morphotypes

of the former: 1) the most numerous, are long, flange-like

structures which occur on the skin covering the upper rim of

the infraorbitals, the snout, cheek muscles, preoperculum

and top of the head; 2) button-like structures confined to

specific areas on the lower cheek, snout and frontal.

The flange-like neuromasts are usually arranged longitudi-

nally and more or less in rows. In the snout region, individual

neuromasts may be slightly curved or angled to the general

direction of the others (Fig. 2B). On top of the head, rows are

more definite and those on the snout tend to converge

anteriorly where the organs close to the midline are larger

than the others (exceptional is the star-shaped arrangement

posterior to the medial extrascapular sensory pore in M.

zugmayeri); the neuromasts close to the midline on the

central part of the frontal are also nearly twice the length of

the others (Fig. 2A). In M. zugmayeri the neuromasts along

the anterior part of the supraoccipital have a regular arrange-

ment (Fig.2A) but in M. gracilis they form a pocket or

enclosed area. Distribution on the preoperculum is irregular

and sometimes sparse (the skin is often missing from this

region and it is not possible to make precise counts).

The pitlines of button-like organs are in a double row near

the border of the snout, in a patch above the nares, as an

G.J. HOWES

Fig. 2.

Distribution of neuromasts types 1 and 2 in A, Melanonus zugmayeri on dorsal surface of head and B, M. gracilis on lateral surface

of head. C, innervation pattern of type 1 neuromasts in subnasal region (right side) of M. zugmayeri (dashed lines indicate nerves, arrow

heads indicate termination of nerve branch; large arrow points anteriorly). In this and subsequent figures, scale bars in millimetre divisions.

oblique row across the lower part of the cheek and a double

row across the epioccipital region (Figs 2A,B). The neuro-

masts do not extend backwards on to the body.

There are about 500 flange-like neuromasts covering the

entire head. Innervation is by the ramus canalis lateralis (Re1

sensu Freihofer, 1970) which exits from the posterior frontal

foramen to anastomose through a loose fascia of connective

tissue. The neuromasts are innervated by subranches stem-

ming from a complex nerve network (Fig. 2C).

The Rel nerve branches from the supraorbital trunk of the

trigeminal complex, (Fig. 17), a condition similar to that in

Merluccius (Freihofer, 1970). The branch innervating the

large posterior neuromasts enclosed in the frontal sensory

canal detaches separately from the supraorbital trunk,

whereas in Merluccius the nerves separate off together.

The large, plate-like neuromasts, housed in the sensory

canals number two in the nasal bone, three in the frontal (one

beneath the anterior medial ridge, one beneath the lateral

arch and one posteriorly), one in the anterior part of the

pterotic, one in the parietal, one in each extrascapular, three

in the first infraorbital, one in the second, third and fourth,

two in the fifth and one in the sixth, and five in the

preoperculum.

Infraorbitals (Fig. 3).

There are six infraorbital bones, the first long and relatively

deep with a broadly fretted ventral border, the outer flange

which forms the roof to the sensory canal extends as a shelf

along the anterior half of the bone, but curves laterally along

the posterior half. The ascending process which contacts the

posterior wall of the lateral ethmoid is tall and spine-like. The

second infraorbital is confluent with the first and is as deep

but only a sixth of its length. The medial lamina of the third is

widely separated from that of the second although the

dorsolateral flange is nearly in contact. The third together

with the fourth form the posteroventral corner of the orbit

and the fourth has only a short orbital margin and flange

ANATOMY OF THE MELANONIDAE

iol

Fig. 3. Infraorbital bones of M. zugmayeri in specimens of: A, 66mm SL; B, 100mm SL and C, 130mm SL.

posteriorly, the body of the bone is expanded. The fifth

infraorbital has a long orbital margin, the lower part of which

projects anteroventrally in front of both the third and fourth

to which it is connected by strong connective tissue; it has a

narrow flange along its upper orbital border. The sixth

(dermosphenotic) is as large as the fifth and has a pronounced

orbital curvature which brings its anterior tip to the same

vertical plane as the ascending process of the first infraor-

bital.

In the two smaller specimens of M. zugmayeri examined,

the dorsolateral flange remains undeveloped on the first, fifth

and sixth infraorbitals of the 66mm specimens and the

ascending process of the first infraorbital is inclined anteriorly

in both (Fig. 3A). The anteroventral border of the fifth

infraorbital is less pronounced and in the 100mm SL speci-

men its tip lies medial to the rim of the fourth infraorbital; the

sixth lacks the anterior elongation of the larger (130mm SL)

specimen (Figs 3B, C).

Unlike other gadiforms where the posterior (fifth and

sixth) infraorbitals are shallow, those of Melanonus are as

deep as the anterior ones. The anterior curvature of the

upper infraorbital (dermosphenotic) is more reminiscent of

some macrouroids (see below) than gadoids. The central

position of the ascending process of the first infraorbital is

probably a plesiomorphic gadiform feature (on the basis of

commonality) as is the reduced size of the second infraor-

bital. The extension of the lower part of the fifth infraorbital

and the enlargement of the dermosphenotic are, because of

their restricted distributions, taken to be derived features.

According to Iwamoto (1989) among macrouroids, exclusion

of the third and fourth infraorbitals from the orbit is a derived

condition. In Melanonus similar exclusion has resulted from

ventral extension of the fifth infraorbital whereas in the

macrouroids illustrated by Iwamoto (1989, fig. 5G) it is due

to re-alignment of the fourth infraorbital which covers the

orbital borders of the second and third.

G.J. HOWES

Fig. 4. Neurocranium of M. zugmayeri in A, dorsal and B, ventral views. In A, the right parietal, right nasal and left posttemporal have

been removed. In B, dashed outline circles on the prootic and intercalar indicate the positions of the otoliths.

Cranium (Figs 4-8).

In its overall shape the cranial roof is almost square, the most

noticeable feature being the deep indentation of the lateral

frontal border anterior to the sphenotic, and the prominent

anterolateral projections of the lateral ethmoid wings

(Fig. 4A).

The ethmoid dorsal surface (rostrodermosupraethmoid) is,

in keeping with that of other gadoids (Howes & Crimmen,

1990: 166), being narrow and cruciform with a steep anterior

slope (Fig. 5B). The ossified anterior wall of the ethmoid

forms most of the nasal cavity and a thin, vertical septum of

ethmoid cartilage separates the cavities medially. A shallow

bed of cartilage separates the base of the ethmoid and the

vomer. The vomer has a thick, broadly rounded head bearing

on either side 6 or 7 teeth in smaller specimens and 10-12 in

larger (Figs 4B, 6A). In smaller specimens of both species the

teeth are more or less arranged in a single row but in larger

specimens the posterior teeth tend to be in a patch with one

or two stout and caniniform being almost twice the length of

their neighbouring teeth and three times that of the symphy-

seal teeth (Fig. 6A). The vomerine shaft is relatively short,

extending to just beyond the posterior level of the lateral

ethmoid. The base of the lateral ethmoid is long and broad

and where it meets the vomer bears a deep cavity into which

inserts the palatine ligament. The wall of the lateral ethmoid

is thin and projects forward at an angle of 45°. The postero-

medial wall extends backward to directly contact the

pterosphenoid.

The nasals (Fig. 4A) are large, almost entirely covering the

lateral ethmoid and are narrowly separated from one another

in the midline by the rostrodermosupraethmoid. Each bone

has prominent anterior and posterolateral processes, two

dorsal processes, lateral and medial, are folded inward to

form curved flanges which support the skin roofing the

sensory canal. In large specimens the nasals tend to become

narrow with attrition of the anterior process (Fig. 6B).

The frontals are nearly square except that the posterior half

of the lateral border is deeply indented. Anteriorly, close to

the midline is a high, arch-shaped crest (afc, Fig. 4A), a

similar but longer arch is situated in the centre of the bone

and is sometimes divided into two separate crests (cfc,

Figs 4A, 5B), posteriorly is a low, diagonal crest (pfc,

Fig. 4A). All these crests shelter a neuromast foramen and

serve to support the skin covering the frontal canal system.

Posteriorly, the frontal margin meets the pterotic, is over-

lapped by the parietal and partially overlaps the anterior

border of the supraoccipital. There are no ventral frontal

laminae. The parietals (Figs 4A, SB) are thin, near-diamond

shaped bones each with a single neuromast foramen and

posterolaterally covered by the median extrascapular. There

is no parietal crest.

The autosphenotic (Figs 4A,B, 5B) has a prominent,

bluntly rounded lateral process and is overlapped by the

CO lice

ANATOMY OF THE MELANONIDAE

ptt

exo

Fig. 5. Neurocranium of M. zugmayeri in A, posterior and B, lateral views. In B, the intercalar is unshaded, the margins of the bones

underlying it indicated by dashed lines. C, parasphenoid of 66mm SL specimen in dorsal view.

frontal, parietal and pterotic. The underside of the bone

bears a deep, almost transverse fossa into which articulates

the hyomandibular. The pterotic (Figs 4A,B, 5A,B) accom-

modates the posterior portion of the hyomandibular fossa

along a third of its lateral border. The wall of the pterotic is

somewhat bullate and its cranial surface forms a prominent

lateral shelf.

The pterosphenoid (Figs 4B, 5B) is long and deep forming

most of the dorsomedial wall of the orbit, anteriorly it

contacts the frontal and posteriorly the autosphenotic and

prootic. The parasphenoid (Figs 4B, 5B,C) has a broad keel

with, extending from its centre, a long, low ascending process

which extends laterally at a low angle to the horizontal plane

to meet the prootic; paired, parallel laminae rise from the

central region of the keel to meet the bases of the lateral

ethmoid wing (Fig. 6).

The prootics (Figs 4B, 5B) are large with a deep trigeminal

notch. The posterior border of the bone is rounded and

partially overlapped by a relatively small, ovoid intercalar to

which is attached the inferior limb of _ the

posttemporal(Figs 4B, 5A,B, 6D). The small, pinnacle-like

epioccipitals contact the posterolateral margins of the

supraoccipital and posteriorly the dorsal borders of their

respective exoccipitals; laterally each epioccipital is overlain

by the second extrascapular (Fig. 4A).

The exoccipitals are deeply depressed posteriorly and con-

tain a large, backwardly facing vagus foramen (Figs 5A,B, 7).

Medially, the bones meet across the midline by flange-like

projections. Posteriorly there is an ovate, cartilage-filled

process the base of which meets its antimere in the midline.

Inside each exoccipital a long, ventrally directed process

extends from the medial surface to contact a shallow dorsal

spt

Fig. 6. Melanonus zugmayeri: A, vomer in ventral view; B, nasal

of left side in dorsal view (broken outline indicates anterior nasal

opening; C, extrascapulars of left side in lateral view;

D, intercalar (left, lateral view). All from a specimen of 173mm

SE:

flange rising from the base of the basioccipital (Fig. 7B). The

basioccipital is a trowel-shaped bone the blade of which forms

the posterior basicranium and the handle, the occipital

condyle (Figs 4,5,7). The supraoccipital (Figs 4A, 5A,B, 7) is

well-ossified and lies flush with the frontals, its crest confined

to its posterior margin; laterally, the bone is bevelled where it

meets the parietal. Posteriorly its ventral margin is bordered

by the exoccipital.

The otoliths have been described and figured by Nolf &

Steurbaut (1983; 1989).

Comments on cranial features

Melanonus has a plesiomorphic ethmo-vomerine region,

namely a broadly rounded ethmoid lacking any dorsal eleva-

tion as in macrouroids and with a single, narrow point of

contact with the lateral ethmoid (Howes & Crimmen, 1990; a

more extensive area of contact appears to be a feature of

some supragadoids, Howes, 1990); a laterally expanded lat-

eral ethmoid which contacts the ascending process of the first

infraorbital ligamentously on its posterior face (Howes,

1987); vomer with a relatively short shaft and well-formed

teeth (absence of vomerine teeth in Macrouroidei and some

gadoids is considered independently derived; see Okamura,

1989; Inada, 1989; Howes, 1990). Ophidiiforms have as

broad a variability of the ethmovomerine region as gadiforms

but the lateral ethmoid is characterised by the presence of

basal twin facets which firmly unite with the large palatine

head. Furthermore, the lateral wing of the lateral ethmoid is

usually reduced and feebly developed, but always has a

lateral facet which articulates with the first infraorbital

(Howes, 1992).

The frontals of Melanonus have a plesiomorphic gadiform

morphology; both gadoid and macrouroid taxa bear frontal

crests of varying development as do ophidiiforms and this

G.J. HOWES

may be a ‘paracanthopterygian’ feature. Howes (1990:79)

noted the lack of ventral frontal laminae in Melanonus and

considered this a derived condition associated with the ante-

rior displacement of the frontal area of the brain (see p.27).

Ventral frontal laminae are widely distributed amongst ophi-

diiforms. There is no prominent V-shaped ridge pattern on

the frontals in Melanonus and no ‘mucosal’ cavity, a feature

of supragadoids.

Nasal bones are plesiomorphically separated in the midline

but in macrouroids are joined for most of their lengths, a

feature regarded as synapomorphic for the group (Iwamoto,

1989; Howes & Crimmen, 1990). Among gadiforms the size

of the nasals is variable but they are nearly always large,

trough-like bones containing two neuromasts. Among plesio-

morphic gadoids (e.g. Bathygadidae) the size of the nasals

approaches that of macrouroids but the bones remain sepa-

rated along the midline. The melanonid condition is thus

considered plesiomorphic although the nasal bones have a

distinct apomorphic shape which more closely approaches

that of some macrouroids than gadoids.

The pterotic of Melanonus has a plesiomorphic gadiform

morphology and resembles that of Bathygadidae in being

broad with a rounded posterior margin and short hyomandib-

ular fossa (Howes & Crimmen, 1990, fig. 6).

The pterosphenoid is unusually large for a gadiform; the

widespread condition (and among ophidiiforms) being small,

occupying the dorsoposterior region of the orbit and widely

separated from the lateral ethmoid. The enlarged anteriorly

extended bone is therefore considered autapomorphic for

Melanonus. The parasphenoid displays no particular derived

feature and corresponds with the situation in the majority of

gadiforms, namely a broad flat keel with parallel laminae

(Howes, 1990:81).

The deeply incised trigeminal notch of the prootic resem-

bles most closely that of some Phycidae and the Muraenolepi-

didae, but unlike those taxa the anterior wall of the prootic is

directed medially as in most infragadoids and macrouroids

(Howes, 1990:82). The intercalar is small in comparison with

that in other gadiforms where in gadoids it is exceptionally

large covering the entire posterolateral cranial wall. A large

intercalar is one of the characters diagnostic of paracan-

thopterygians but is secondarily absent in lophiiforms and

batrachoidiiforms. Among ophidiiforms and percopsiforms

the intercalar is generally not as large as that of gadiforms,

and in those two former groups is confined to the upper half

of the posterior cranial wall and does not extend ventral to

the basioccipital anteriorly but is interrupted by the prootic.

The relationship between the epioccipital and supraoccipi-

tal is an unusual one amongst gadiforms in that the posterior

walls of the exoccipitals meet across the midline and so

exclude the supraoccipital from contributing to the upper

margin of the foramen magnum. Elsewhere in paracan-

thopterygians this condition occurs among ophidiiforms

(Howes, 1992) where the exoccipital has enlarged backward

and upward to cover the posterior margin of the supraoccipi-

tal. Even in the largest cleared and stained specimen of M.

zugmayeri examined for this feature the ventral tip of the

supraoccipital does not reach the margin of the foramen

magnum (Fig. 7B). The supraoccipital lacks a dorsal crest,

there being a posterior lamina (Fig. 7). Howes (1990:82)

discussed the variability of the supraoccipital crest amongst

gadoids. An elevated cranial crest is possibly the plesiomor-

phic condition for paracanthopterygians but a low, reduced

crest is widely distributed amongst all groups and in lophii-

ANATOMY OF THE MELANONIDAE

SO nal

boc

Fig. 7. Melanonus zugmayeri posterior part of cranium in lateral oblique view of 130mm SL specimen showing in A, first neural arch and

vertebra attached and in B, removed to expose the posterior features of the basi- and exoccipitals. Note the supraoccipital does not

contribute to the border of the foramen magnum.

forms and ophidiiforms appears to be the common condition.

It is assumed that this feature has been repetitively evolved in

these groups.

Jaws (Fig. 8)

The premaxilla (Fig. 8C) has tall, thin and widely separated

ascending and articular processes, and a tall, spine-like

postmaxillary process. The toothed surface is narrow, bearing

for most of its length two rows of sharp pointed teeth. The

outer row teeth are straight or extend slightly laterad, the

inner row teeth which are about twice the length of the outer

are inwardly curved; posteriorly there are three rows of teeth,

the ones of the centre row being the same size as those of the

inner (Figs 8D,E). Ina 100mm SL specimen of M. zugmayeri

the posterior teeth are so arranged as to form distinct

transverse rows (Fig. 8F) but this is not evident in the 66mm

or 130mm SL specimens. The maxilla has a tall articular head

and a short medial articular process forming a rather acute

angle with the head (Figs 8A,B). The shaft of the bone is

slender and posteriorly bears shallow dorsal and ventral

processes.

The dentary (Fig. 8G) is short and deep with a correspond-

ing shallow mentomeckelian cavity; it has a high steep

coronoid process. The sharp pointed teeth are set in an

irregular single row, numbering 22 in 88mm and 100mm SL

specimens of M. zugmayeri,28 in a 130mm and 34 in a 175mm

SL specimen. The anterior teeth are small followed by four or

five successively larger ones, then four or five relatively large

teeth separated by three or four smaller ones. Posteriorly the

teeth diminish in size. The anguloarticular (Fig. 8G) is tall

with a steep posterior slope and short, vertical anterior

margin; the articular condyle is long and narrow. The coro-

nomeckelian bone (Fig. 8G) is a well-developed, cylindrical

element with slight dorsal and ventral posterior flanges. The

retroarticular (Fig. 8G) is boot-shaped, the leg being curved

forward and the foot long and shallow. A strong labial

ligament is anchored to the rim of the dentary (Howes, 1988,

fig. 12).

The overall jaw morphology of Melanonus is plesiomorphic

for gadiforms, the upper jaw bones, apart from having a

smaller postmaxillary process of the premaxilla, are little

different from those in bathygadids (Howes & Crimmen,

1990). Macrouroids are characterised by having a large

postmaxillary process of the premaxilla situated posteriorly

(Okamura, 1970; Howes & Crimmen, 1990). There is no

‘gadoid notch’ at the base of the postmaxillary process. The

lower jaw more closely resembles that of gadoids or ophidii-

forms than macrouroids in having a relatively shallow angu-

loarticular and boot-shaped or J-shaped retroarticular.

G.J. HOWES

Fig. 8. Melanonus zugmayeri, jaw bones. A and B right maxilla from a 130mm SL specimen in: (A) dorsal and (B) medial and slightly

ventral views; C, premaxilla in lateral view; D-F premaxilla, anterior (D) and posterior (E) regions from 130mm SL specimen and (F)

100mm SL specimen, ventral views; G, lower jaw of 130mm SL specimen in medial view.

Macrouroids tend toward a deeper anguloarticular and

greater variability in the shape of the retroarticular (Oka-

mura, 1970; Howes & Crimmen, 1990). A boot-shaped

retroarticular is lacking in both percopsiforms and lophii-

forms.

Palatopterygoquadrate (Fig. 9)

The palatine (Fig. 9A,B) is long, its posterior tip extending to

nearly halfway along the ectopterygoid, its rostral process is

long and slender and overlies the maxilla, its base bears a

broad facet which articulates with the ethmoid cavity and the

body of the bone rises to a high posterior crest. There are two

rows of sharply pointed teeth.

The anterior part of the ectopterygoid (Fig. 9A) lies along

the medial face of the palatine and its ventral stem reaches

the quadrate joint; laterally it is slightly overlapped by the

entopterygoid (Fig. 9A). The latter is a relatively large bone

with a rounded dorsal profile and is sloped mesad, its

posterior border is well separated from the hyomandibular by

the metapterygoid. The metapterygoid (Fig. 9A) is axe-

shaped its posterior margin rising high up the leading edge of

the hyomandibular shaft.

The melanonid palatine is unique amongst gadiforms, in its

length, nature of contact with the pterygoids, and in bearing

teeth. The common condition, and one which is considered

synapomorphic for gadiforms (p.29) is for the palatine to be

reduced in length with a vertical or slightly angled posterior

border meeting a similar blunt margin of the ectopterygoid

and forming a hinge-type joint (see figures in Okamura, 1970,

1989; Howes, 1990, 1991b; Howes & Crimmen, 1990). This

union differs from that commonly encountered in other

paracanthopterygians where the posterior limb of the pala-

tine is attenuated and articulates firmly with the leading edge

of the entopterygoid and lateral face of the ectopterygoid.

Percopsids resemble gadiforms in having a near vertical

abutment of the palatine with the ento- and ectopterygoids

(Fig. 9D). However, there is a posterior stem which overlaps

the upper lateral margin of the ectopterygoid. Macrouroids

are characterised by the lack of direct contact between the

palatine and ethmovomerine bloc (Howes & Crimmen,

1990).

Norman (1930) noted there were ‘teeth on the pterygoid’,

an error perpetuated by Howes (1991b, caption to fig. 35).

The pterygoid bones of Melanonus display plesiomorphic

morphologies; the large entopterygoid and high posterior

ANATOMY OF THE MELANONIDAE

Fig. 9. A-—C Melanonus zugmayeri: A, palatoquadrate, hyosymplectic and opercular bones of 130mm specimen in medial view, light hatched

area represents ligamentous system connecting opercular bones to hyomandibular; B, palatine of 100mm specimen, right side, lateral view;

C, hyomandibular, left side, of 100mm SL specimen in anterior view showing foramen for hyoid branch of facial nerve (arrowed) and

lateral flange; D, Percopsis omiscomayus, palatine and pterygoids in lateral view (heavy dotting indicates cartilage).

metapterygoid process are present in macrouroids, bathyga-

dids and macruronids (Howes & Crimmen, 1990; Howes,

_1991b). Reduction of the entopterygoid and metapterygoid

appears to be characteristic of supragadoids (Howes, 1990).

Amongst ophidiiforms the metapterygoid abuts against the

lower limb of the extended anterior portion of the hyoman-

dibular.

The quadrate (Fig. 9A) of Melanonus has a wide angle

between its posterior border and the posteroventral spine.

The size of this angle is variable among gadiforms and

appears correlated with the orientation of the suspensorium.

An ‘interosseuos space’ between the symplectic and preoper-

culum (Okamura, 1970; 1989) is also a condition of the

angular separation of the two parts of the quadrate, being

absent where the angle is small (Howes, 1990).

Hyoid arch (Figs. 9-11)

The hyomandibular (Fig. 9A) has, as in all gadiforms, a single

articulatory condyle. The bone is narrow-waisted with the

relatively long shaft oval in section, a foramen for the hyoid

branch of the facial nerve pierces its posterior margin (Fig.

9C). Posteriorly is a long, horizontal process which articulates

with the opercle. The lateroposterior face contacts the border

of the preopercle. Medially a band of ligamentous connective

tissue joins the shaft with the opercular process and a wider

band runs at right angles to it to attach to the subopercle and

interopercular-subopercular ligament (Fig. 9A).

The course of the hyoid branch of the facial nerve is

partially exposed laterally, due to attrition of the outer part of

the hyomandibular, part of which remains as a lateral flange

which is a common feature (synapomorph) for gadiforms,

Fig. 10. Melanonus zugmayeri hyoid bar of 100mm SL specimen:

A, medial view; B and C, urohyal in lateral and dorsal views.

one not shared by ophidiiforms or lophiiforms (Howes,

1992).

Other hyoid arch bones are much like those of the majority

of gadiforms; the posterior half of the anterohyal =ceratohyal

auct. (Figs 10A, 11A) is deep and in this respect resembles

that bone in some macrouroids (eg. Nezumia, Abyssicola,

Coelorhynchus, Coryphaenoides; Okamura, 1970), more

closely than gadoids. However, this feature is variable and a

similar range of morphotypes can be found among ophidii-

forms (Markle & Olney, 1990, fig. 13). As in most gadiforms

and ophidiiforms there are 7 branchiostegal rays which

appears to be the plesiomorphic paracanthopterygian num-

ber, (six occur frequently in lophiiforms). The urohyal

(Figs 10B,C) bears a closer resemblance to that of gadoids

rather than macrouroids in having a shallow dorsal keel and a

long, prominent anterodorsal (basibranchial) process

(Howes, 1990, fig. 16B).

The basihyal (Fig.11A) is a dumbbell-shaped bone lying

between the dorsohyals and crossed by a ligament which

connects them; anteriorly a thick cartilaginous ‘tongue’ pro-

trudes forward, posteriorly, the basihyal is slightly over-

lapped by the first basibranchial(see below). The interhyal

(Fig. 9A) is typically gadiform, being long and slender,

contacting the symplectic cartilage dorsally and the posterior

socket of the posterohyal ventrally. Markle (1989, fig. 6A)

shows a common ligamentous connection between the

interhyal-posterohyal and interopercle. I find this to be one

involving thick connective tissue although a discrete ligament

runs from the medial side of the interhyal to the medial

posterior tip of the posterohyal.

Opercular bones (Fig. 9A)

The opercular bones are relatively generalised except that the

suboperculum has a straight to concave leading edge rather

than the common gadiform condition of a rounded to pointed

margin. The interopercle is shallow and nearly oblong with

rounded dorsoposterior and anteroventral borders; it is

widely separated from, and ligamentously connected to the

subopercle. In general, macrouroids have the interopercle

orientated horizontally (e.g. Okamura, 1970, figs 26; 27)

whereas in gadoids the bone is angled, sometimes steeply as

in Melanonus. Melanonus lacks the interopercular fossa

present in a subgroup of ‘supragadoids’ (Howes, 1990). The

opercle is relatively large for a gadiform and overlaps most of

the subopercle. The preopercle has a short lower, anteriorly

directed limb and a narrow laminate (symplectic) process

G.J. HOWES

which, plesiomorphically, contacts the symplectic cartilage.

In its derived form the symplectic process of the preopercle

contacts the lateral face of the hyomandibular (Howes, 1990).

Branchial arches (Fig. 11)

There are three basibranchials (Fig. 11A), the first and

second ossified, the third cartilaginous. The posterior margin

of the first overlies the the anterior border of the diamond-

shaped second which is separated from the small diamond-

shaped third. The first and second hypobranchials (Fig. 11A)

are long with marked posterior curvature, both contacting the

first basibranchial and bear gill-rakers on their outer and

inner margins; the third is short and lacks gill-rakers. The first

and second ceratobranchials (Fig. 11A) bear five or six

clustered-spinous rakers on their outer and the same number

of slender, triple-spine rakers on their inner margins; the

third has seven outer and inner shorter rakers and the fourth

has four short rakers on its outer margin only. The anterior

tips of the fifth ceratobranchials are apposed but not firmly

united in the midline and are ligamentously connected to the

third basibranchial; a narrow tooth patch bears ca 25 slender

pointed teeth.

The epibranchials (Fig. 11B) are 30% the length of the

ceratobranchials. A strong uncinate process on the first

epibranchial is connected by a chondrified ligament to a large

interarcual cartilage; the third epibranchial bears a long tooth

plate bearing ca 20 sharp pointed teeth. There are four

pharyngobranchials (Fig. 11B), the first being an ossified

element; the second-third pharyngobranchial tooth plates

bear strong, pointed teeth.

Markle (1989) has described and commented on the upper

dhy

bb1-3

Fig. 11. Melanonus zugmayeri branchial arches of 130mm SL

specimen: A, dorsal view of lower arch elements; B, ventral view

of upper arch elements. In A, basihyal is also shown in lateral

view.

ANATOMY OF THE MELANONIDAE

branchial arch of Melanonus which he considers, due to the

presence of a large and chondrified interarcual ligament, to

be plesiomorphic for gadiforms. In his cladogram, however,

he mistakenly ascribes to Melanonus the loss of the second

pharyngobranchial. The lower gill-arch is also plesiomorphic

in that the basibranchials are unexpanded and there is no

forward ventral elongation of the third hypobranchial as in

macrouroids and ophidiiforms; the first hypobranchial is

typically long as in gadiforms but lacks an expansion where

the ligament running to the dorsohyal attaches. Spinous

gill-rakers of both the clustered- and triple-spine type are

widespread amongst ‘infragadoids’ and macrouroids.

Pectoral girdle (Fig. 12)

The vertical and horizontal limbs of the cleithrum (Fig. 12A)

are nearly equal in length; the medial cleithral lamina is thin

and only prominent near the cleithral tip. Markle (1989)

noted that the foramen which notches medial borders of both

the scapula and coracoid is present only in the former

(Fig. 12A). Markle (1989) and Howes & Crimmen (1990)

commented on the variability of this feature; plesiomorphi-

cally the foramen lies entirely within the scapula, a condition

almost entirely confined to ‘infragadoids’ although it is also

recorded in the ‘supragadoid’ Lota. The supracleithrum

(Fig. 12B) is a lanceolate bone with a slightly expanded

dorsal articulatory surface which contacts the posttemporal.

There are four actinosts and 12 or 13 pectoral rays in the M.

zugmayeri specimens examined (Norman, 1930, gives 13 for

M.zugmayeri and 12-14 for M. gracilis; Fahay & Markle,

1984, give a range for the genus of 10-16). The single

postcleithrum (Fig. 12A) has a broad head and slender,

slightly upwardly curved stem. It articulates in a cleft oppo-

site or slightly above the coracoid-scapula junction (see also

Markle, 1989, fig. 10).

The posttemporal (Fig. 12B) is V-shaped, its upper limb

broad proximally and tapering distally; its lower limb, which

is firmly united with the intercalar is thin, rod-like and

completely ossified.

The extrascapulars (Figs 4A, 5A,B, 6C) number four, each

having upturned borders and containing a neuromast. The

lateral extrascapular covers the posterior corner of the

pterotic, two lie in contact with one another along the medial

part of that bone and the innermost lies along the lateral part

of the parietal. In large specimens of M. zugmayeri the

scl

Fig. 12. Melanonus zugmayeri: A, pectoral girdle in medial view; B, posttemporal and supracleithrum in lateral views; C, pelvic girdle in

dorsal view.

medial extrascapular is more closey aligned with the supraoc-

cipital, resting along a lateral ridge of the bone and in a

specimen of 173mm SL (Fig. 6C), it appears that the lateral

and a medial extrascapular have become fused, judging by

the presence of two neuromast foramina in the single large

bone.

Pelvic girdle (Fig. 12C)

The pelvic bone is narrow and tubular, broadening proxi-

mally where its cartilaginous tip contacts its antimere sym-

physially. Distally the pelvic process is narrow and straight

and connected with its antimere by ligamentous tissue. There

is no lateral pelvic process or spine (cf. Bathygadidae, Howes

& Crimmen, 1990). There are usually 7 fin rays; Fahay &

Markle (1984) give a range for the genus of 5-7.

The pelvic girdle lies well forward with the anterior tips of

epr

G.J. HOWES

the pelvic bones lying between the cleithra so that the origin

of the pelvic fin lies beneath or just anterior to that of the

pectoral (Fig. 1). The position of the pelvic girdle in relation

to the pectoral girdle is variable amongst gadiforms. In the

majority of gadoids the pelvic girdle is situated well forward,

particularly so in the more derived ‘supragadoid’ taxa such as

gadids, gaidropsarids and muraenolepidids, so that the origin

of the pelvic fin lies in advance of that of the pectoral fin.

‘Infragadoids’ tend to have the pelvic girdle situated beneath

or behind the pectoral (e.g. bathygadids, steindachneriids).

In morids, however, the pelvic girdle lies well forward. There

is some variability in position among macrouroids but gener-

ally, the pelvic girdle lies posterior to the pectoral so that the

origin of the pelvic fin is situated directly beneath that of the

pectoral fin. With respect to the position of the pelvic fin

relative to that of the pectoral, the Melanonidae appear to

represent an intermediate condition between the derived

forward and plesiomorphic posterior positions.

Fig. 13. Melanonus zugmayeri vertebral column. A, first vertebra and neural arch of 130mm SL specimen in lateral and anterior views; B,

anterior part of vertebral column of 100mm SL specimen showing retractor dorsalis muscle of one side, in ventral view; C and D vertebral

column and fin supports of 130mm SL specimen: C, anterior vertebrae, D, 14th-17th vertebrae showing anterior anal fin supports (lateral

views; ribs shown in black for clarity).

ANATOMY OF THE MELANONIDAE

Vertebral column and median fins (Figs 13-15)

There are 12-14 abdominal and 45 or 46 caudal (those with

closed haemal spines) vertebrae in Melanonus (Fahay &

Markle, 1984 give total counts of 58-62 for the genus). The

first neural arch and spine are well-developed and form an

ankylosed unit with the centrum. The prezygapophyses of the

- first vertebra (Fig. 13A) are oval in section, hollow and

cartilage-filled and firmly in contact with the similarly shaped

paired condyles of the exoccipital. The wall of the neural arch

covers the upper posterior wall of the exoccipital leaving

exposed a notch through which pass the occipital and lateral

line nerves (Fig. 7B). The laminae of the neural arch extend

forward to embrace the posterior extension of the supraoccip-

ital crest (Fig. 7A). The second vertebra is anteroposteriorly

compressed and lacks processes or ribs; the third-fifth verte-

brae support successively shorter chopstick- shaped ribs

which extend almost horizontally, at their tips lie epipleural

(epineural) ribs the heads of which are ligamentously

attached to their respective myosepta (Fig. 13B,C). The

sixth-twelfth centra bear triangular parapophyses to each of

which is attached a posteriorly curved epipleural rib. Accord-

ing to Okamura (1989) there is a total of eleven epipleural

| ribs in Melanonus; ten are counted here in M. zugmayeri.

There is a single dorsal fin comprising 72-78 rays. The first

| dorsal ray is often minute, the second and successive rays are

long and flexible, supported by distally tapered rod-like

radials which tend to occur in pairs within each interneural

space, their proximal tips converging (Fig. 13C,D). The

origin of the dorsal fin occurs between the third and fourth

_ neural spines. There are no supraneurals (predorsals). The

| anal fin has 50-58 rays and lacks a stout anterior spine; the

shape of the radials is similar to those which support the

dorsal fin (Fig. 13D).

| Caudal fin skeleton (Fig. 15A,B). The caudal fin skeleton

_ of Melanonus resembles that of the Moridae in that the first

_ and second hypurals are incompletely fused; each support a

| single fin ray. In morids all the hypurals are fused only

| proximally whereas in Melanonus fusion of hypurals 1 and 2 is

' both proximal and distal leaving a central opening

(Fig. 1SA). Hypurals 3-5 although fused in specimens of M.

zugmayeri of 130mm SL are only partially fused in 66mm and

100mm SL specimens (Fig. 15B). Paulin (1983, fig. SA)

figures a caudal skeleton of M. gracilis in which hypurals 1

and 2 are entirely fused and the fifth is reduced. In a 45mm

SL specimen of M. gracilis, all the hypurals are separated for

their entire lengths whereas in a 49mm SL specimen they are

fused distally but not proximally. There are two elongate

epurals each supporting a fin ray; in a 100mm SL specimen of

| M. zugmayeri they are joined proximally (Fig. 15B). Unlike

/ morids, Melanonus lacks X and Y bones a feature shared with

| Macruronidae, Gadidae and Lotidae. A long parhypural

| articulates basally with the fused hypurals 1 and 2 and

| supports a single fin ray.

Comments on features of the vertebral column and median

_ fins. Chopstick-shaped ribs, similar to those of Melanonus,

have been reported for Macruronus, Lyconus, Steindachneria

-and Merluccius by Okamura (1989) and Inada (1989) who

‘arrive at Opposite conclusions with regard to their character

| polarity. According to Okamura this rib-type suggests a close

| relationship between the taxa in which they occur. Inada, on

the other hand, regards them as a plesiomorphic gadoid

feature. Although Inada’s (1989) reasoning appear to be

based on an a priori assumption of merlucciid plesiomorphy I

would agree with his assumption. In fact, this type of rib is

more widely distributed amongst gadoids than has been

reported and also occurs amongst ‘supragadoids’ other than

Merlucciidae (Fig. 14A).

In Melanonus the ribs occur on vertebrae 3-5 as in Lyco-

nus, but they are on vertebrae 3 and 4 in Macruronus (both

Macruronidae), 3-6 in Merlucciidae, and 34 in Gaidrop-

saridae. In Steindachneriidae the ribs are on vertebrae 3 and

4 but the rib on the fourth has less than half the thickness of

that on the third whereas in the above cited taxa the ribs are

of equal thickness. Furthermore, the epipleurals attach

directly to the distal tips of the chopstick ribs in Steindachne-

ria whereas in the other taxa they are indirectly attached by

ligamentous strands running to the myosepta (as in Mel-

anonus). In morids and ‘supragadoids’ epipleurals are

attached directly to the vertebral ribs. Patterson & Rosen

(1989) interpreted the vertebral ribs Steindachneria and

Gadus as a parapophysis with attached epipleural. The ribs in

Steindachneriidae, however, are like those of Melanonidae,

Macruronidae and Merlucciidae in articulating with the ven-

tral cavity of the centrum. In the Bregmacerotidae the third

and subsequent vertebrae bear parapophyses to which are

attached cartilage-formed ribs (Fig. 14B). The loss of epi-

pleurals from the first and second centra is a gadiform

synapomorphy (Markle, 1989).

Howes (1991b) regarded the first neural arch of Macruro-

epr

Fig. 14. Anterior region of vertebral column in: A, Gaidropsarus

mediterraneus; B, Bregmaceros sp. In A, black shading in the ribs

(ar) indicates zones of cartilage.

G.J. HOWES

hy3-5

fv fvi ird

Fig. 15. Melanonus zugmayeri Caudal fin skeletons of A, 130mm SL and B, 100mm SL specimens. C, caudal fin musculature (although a

superficial layer of connective tissue and some muscle has been removed the vertebrae are exposed in situ as shown).

nus (Macruronidae) as a composite unit incorporating an

accessory neural arch suggesting that the first centrum had

been incorporated in the ‘basioccipital’. Since, however, 1)

ribs are always lacking from the first two vertebrae in

gadiforms, 2) Baudelot’s ligament always occurs on the first

centrum and 3) an accessory neural arch does not occur above

aulopiforms, it seems untenable that incorporation has

occurred in macruronids.

The caudal fin skeleton is lacking in the majority of

gadiform taxa but where it does occur its most significant

features are fusion of the upper hypurals and the presence of

X and Y bones (lost in some taxa, see above), both of which

contribute to the symmetry characteristic of ‘supragadoids’.

The morid caudal fin skeleton is regarded as the plesiomor-

phic gadoid type since it approaches that of most other

teleosts in its asymmetry and in having distally separated

hypurals. In this latter respect, Melanonus demonstrates a

further derived condition in having the hypurals distally fused

(see further discussion on p.30).

Of particular note is the condition of the caudal fin

musculature (Fig. 15SC) which differs from that described in

gadoids (Howes, 1991) where hypochordal longidorsales,

flexores dorsales and inferiores are absent, the interradiales

have a characteristic linkage pattern between the caudal fin

rays and are continuous with the dorsal and anal fin rays.

Howes (1991: 104) pointed out the absence of the latter in

Melanonidae, but overlooked the fact that the caudal fin

musculature more closely resembles that of other paracan-

thopterygians and acanthopterygians in having discrete dorsal

and ventral flexores and an amalgamated segment of interra-

dialis musculature corresponding to the superficial interradia-

lis.

Melanonus, Lyconus and Brosme are the only gadiform

taxa to possess a single dorsal fin, most have two and the

more derived Gadidae have three. Dorsal fin origin is usually

above the second and third neural spines, as in Melanonus,

but the origin of the second dorsal is variable, the radial

supporting the first ray of that fin being between the eight and

ninth, ninth and tenth or tenth and eleventh neural spines.

According to Inada (1989) the single dorsal fin of Lyconus

evolved from amalgamation of two separate fins. Inada’s

evidence relies on a notch being present in the fin at a point

above the proximally curved thirteenth radial which lies

between the eighth and ninth neural spines which as just

noted is the region commonly associated with the origin of

the second dorsal fin. No similar ‘evidence’ occurs in Mel-

anonus.

Among gadoids the first radial of the first dorsal fin usually

lies between the second and third neural spines, but this is

variable being between the first and second in gaidropsarids,

and in Bregmacerotidae the first radial has become directed

forward so that the first dorsal fin ray lies above the supraoc-

cipital. In macrouroids, the first supporting radial is also

usually between the second and third neural spines but

sometimes between the third and fourth. Percopsids, like

melanonids have the first radial between the third and fourth

neural spines. In ophidiiforms the position of the first radial is

variable and can lie between any of the neural spines from the

ANATOMY OF THE MELANONIDAE

first to the tenth. In batrachoidiiforms it is usually between

the third and fourth neural spines and in lophiiforms the

eighth and ninth or more posterior neural spines.

Supraneurals, preceding the first dorsal fin are rarely

present in gadiforms (Patterson & Rosen, 1989).

Baudelot’s ligament (Figs 13A,B) stems from the lateral

cavity of the first vertebra to connect with the supracleithrum.

The retractor dorsalis muscle originates from the fourth

through sixth vertebrae; on the sixth it is attached to the

leading edge of the parapophysis (Fig. 13B).

Brain (Fig. 16).

The brain of Melanonus is situated well forward, the telen-

cephalon and anterior part of the mesencephalon being

anteriorly displaced beyond the cranial cavity so as to lie in

the orbital cavity formed by the enlarged pterosphenoids.

The olfactory and optic lobes are large. The olfactory tracts

are well separated and each tract is short and thick compris-

ing at least twelve separate nerves each of which branches to

=™

nvo gé n¥W nau

feed the individual laminae of the nasal rosette. The olfactory

bulb is large and lies against the lobe which is narrowly

separated by a fissure from the laterally situated optic lobe.

Upon leaving their respective lobes ventrally, the optic tracts

cross and travel directly laterad a short distance to the eyeball

which is only narrowly separated from the telencephalon.

The cerebellar corpus is flat and lies pointing anteriorly

between the optic lobes. This is a unique condition among

gadiforms (noted by Marshall & Cohen, 1973 as diagnostic of

the Melanonidae), normally the corpus is bulbous and ele-

vated (Okamura, 1970) or lies posteriorly along the cerebel-

lar crest. The cerebellar crest is flat and elongate flanked

ventrolaterally by extensive trigeminal lobes. The cerebellar

body extends posteriorly to entirely overlap the vagal lobes

along the basal part of the medulla oblongata, also a unique

gadiform condition. The granular eminence is large but not

laterally extended. Ventrally, the inferior lobes, pineal body,

hypophysis and vascular sac are all well-developed.

The brains of some gadoids and macrouroids have been

described by Svetovidov (1953), Okamura (1970) and Howes

nIX nx nil

| Fig. 16. Melanonus zugmayeri brain in A, dorsal and B, lateral views. In A, the pathways of the optic tracts beneath the lobes are indicated

by dashed lines and the margin of the prootic is indicated by dashed lines lateral to the trigeminal-facial nerve complex.

& Crimmen (1990) and of those published descriptions Breg-

maceros has the most similar overall morphology. Like that

of Melanonus the brain is elongate with extensive trigeminal-

facial lobes, a long cerebellar crest and closely connected

olfactory bulb and lobe. However, there are major differ-

ences in the relatively small size of the olfactory and inferior

lobes and in the cerebellar corpus being orientated posteri-

orly along the crest, having a posterolateral fissure and

leaving the midline of the optic lobes exposed.

Anterior placement of the forebrain was considered a

gadiform character by Svetovidov (1948) and among gadoids

there is a tendency for the brain to be shifted forward. In

those few morids investigated and in macrouroids the fore-

brain is generally confined to the cranial cavity. In some other

paracanthopterygians (ophidiiforms, Howes, 1992 and per-

copsids pers. obs.)the telencephalon lies in the orbital cavity

as in Melanonus.

It is problematic as to which features of gross brain

morphology can be used as phylogenetic markers. The degree

of separation of the olfactory bulb from the lobe is variable in

gadiforms (discussed by Howes & Crimmen, 1990) but the

plesiomorph condition, possessed by Melanonus, is seemingly

for them to be closely associated. The shape of the olfactory

lobe is also a highly variable feature and one that might, at

least, be generically characteristic.

Summarising data from gadoid brain descriptions given by

Svetovidov (1953) it appears that elongate and short cerebral

crests are equally distributed amongst the taxa he studied. A

short, tall cerebral crest, common to gadoid brains, is also the

common condition among paracanthopterygians. However,

the granular eminence, although often large is laterally

extended only in the Gadidae (sensu Dunn, 1989 and Howes,

1991b).

Swimbladder, viscera and body musculature

(Fig. 17).

The swimbladder is an elongate ellipsoidal, thin-walled sac

adhering tightly to the vertebral column apart from where the

long bilobed kidney runs on either side of the midline.

According to Marshall & Cohen (1973) the melanonid swim-

bladder is reduced and there are two retia. In the specimens

of M. zugmayeri examined for this feature, the gas-gland

covers nearly two-thirds of the anterior floor of the sac and

there are four retia supplying separate lobes. Posteriorly the

gas-gland tapers and is deeply pocketed. The oval appears to

be beneath the retial area.

The stomach is siphon-shaped, exceptionally thick-walled

with a deeply and much convoluted mucosal membrane;

there are six or seven caeca lying ventrally; the intestine is

long and double-bended. The bilobed kidney is extensive,

almost enveloping the stomach. The gonads lie posteriorly on

either side of the swimbladder to which they are attached by

thin strands.

The anterior body musculature is similar to that described

for Bathygadidae (Howes & Crimmen, 1990) except that

melanonids lack the same degree of differentiation between

dorsal and ventral sections of the epaxialis musculature (Fig.

17), the dorsal section being apparent only anteriorly (the

general condition) and not extended as far posteriorly as in

bathygadids.

G.J. HOWES

epr

sca

dex

vex

Fig. 17. Melanonus zugmayeri anterior body musculature and

visceral cavity dissected on the right side; the anterior part of the

liver has been cut away to expose the pyloric caeca and the

swimbladder has been dissected.

DISCUSSION

Melanonus has undoubtedly derived sensory features; the

brain has a unique morphology amongst paracanthoptery-

gians, extending well forward into the orbital cavity, the head

is covered with a unique type and pattern of open-ended

neuromasts innervated by the ramus canalis lateralis of the

trigeminal nerve, the RLA nerve being absent. In its cranial

osteological characters three can be considered derived: the

shape of the fifth infraorbital, and exclusion of the supraoc-

cipital from contributing to the foramen magnum. The first

two of these osteological characters are autapomorphic; the

enlarged pterosphenoid is undoubtedly correlated with the

anterior position of the telencephalon. The third is a feature

shared with ophidiiforms, in that group, however, the exoc-

cipital is expanded dorsoposteriorly so as to exclude most of

or the entire supraoccipital from the rear of the cranium and

from contact with the first neural spine. In melanonids the

supraoccipital is excluded from the border of the foramen

magnum by its failure to extend ventrad during development,

but nevertheless it still forms the upper posterior border of

the cranium and contacts the first neural spine.

Aside from these autapomorphies there are no other

apomorphies which are shared with other gadoid taxa. The

diagnosis of “Gadoidei’ has proved difficult since most syn-

apomorphies so far proposed are either not exclusive to, or

their distribution has not been completely documented, in the

taxa currently embraced under this category (see below).

Howes (1990; 1991a; 1991b) proposed a series of gadoid

clades of which ‘supragadoids’ were recognised on the basis

of a fused upper hypural plate of the caudal skeleton. A

sequence of other synapomorphies, including an interopercu-

lar fossa, contact of the posterior face of the lateral ethmoid

wing by the first infraorbital and reduction of pterygoid bones

(Howes, 1990) excluded Melanonidae from this group. Other

ANATOMY OF THE MELANONIDAE

MELANONOIDE! MACROUROIDEI

6-9

1-5

GADOIDE!

(Bathygadidae Steindachneriidae Moridoidea Gadoidea

14;15

12 ;13

10;11

Fig. 18. Proposed relationships of Melanonoidei with other gadiforms. Synapomorphies: 1, absence of pars jugularis, i.e. common aperture

for principal cranial nerves (also occurs in some ophidiiforms); 2, loss of intermusculars from vertebrae 1 and 2; 3, scapular-coracoid

foramen; 4, attrition of lateral face of hyomandibular; 5, levator arcus palatini covers lateral face of jaw musculature; 10, palatine forming a

hinge or butt-joint with pterygoids; 11, enlarged intercalar contributing to posterior wall of cranium; 12, pharyngohyoideus muscle mediated

by sternohyoideus; 13, interradiales muscle connected to dorsal and anal fin rays, loss of various caudal fin muscles and entire caudal skeleton

in some taxa; 14, palatine contacts mesethmoid; 15, X and Y bones in caudal skeleton (lost in some taxa); 16, complete fusion of upper

hypurals and symmetry of hypural plates. Autapomorphies for Melanonoidei; 6, supraoccipital excluded from margin of foramen magnum; 7,

cranial neuromast pattern and innervation; 8, brain position and morphology; 9, enlarged pterosphenoids contacting lateral ethmoids.

Synapomorphies 1-5 and 10-11 from Gosline (1968; 1971); Howes (1988; 1989; 1990; 1991b); Markle (1989); Patterson & Rosen (1989).

Synapomorphies for macrouroids summarized by Iwamoto (1989) and Howes & Crimmen (1990) and for moridoids by Paulin (1983).

taxa so excluded are Moridae, Euclichthyidae, Steindachneri-

idae and Bathygadidae. The two latter lack a caudal fin

skeleton,thus the incomplete fusion patterns of hypural bones

possessed by morids, euclichthyids and melanonids cannot be

extended to these taxa. The cranial and vertebral osteology of

Bathygadidae is plesiomorphic in comparison to other gadoid

taxa whereas that of Steindachneriidae is relatively derived

(pers. obs. see also Fahay’s, 1989, notes on pelvic girdle

morphology).

The Melanonidae lacks a feature common to other

gadiforms (macrouroids + gadoids), namely, a short palatine

forming a butt or hinge joint with the ento- and ectopterygo-

ids. In almost all gadiforms the palatine has a truncated near

vertical margin which forms a mobile (laterally expanding)

joint with the anterior margins of the pterygoid bones (p.20).

Melanonus has a plesiomorphic palatine where the stem

firmly contacts the margin of the ectopterygoid. Moreover,

the palatine extends some distance along the ectopterygoid

and is toothed. Since no other gadiform has palatine teeth it

might be assumed that the melanonid palatine is the primi-

tively composite dermo- and autopalatine whereas other

gadiforms have lost the dermal component. In other paracan-

thopterygians, ophidiiforms and lophiiforms possess the ple-

siomorphic, long posteriorly extended and toothed palatine;

percopsids resemble gadiforms more closely in having an

edentulous bone which abuts the straight anterior margins of

the ecto- and entopterygoids but which still retains a posteri-

orly directed stem (p.20).

The Melanonidae possesses three of those characters iden-

tified by Patterson & Rosen (1989) and Markle (1989) as

gadiform synapomorphies or potential synapomorphies,

namely, absence of epipleural ribs from the first and second

vertebrae; a scapular-coracoid foramen and absence of a

lateral commissure, cranial nerves I-VII exiting through a

common aperture. Two other potential synapomorphies

listed by Patterson & Rosen (1989) are presence of X and Y

bones and liver LDH pattern. X and Y bones are absent in

melanonids and can only be judged as a plesiomorphic state

or, against the congruence of other synapomorphies, as

secondary loss. In the latter case the feature then appears as

synapomorphic for a subgroup of gadoids (Fig. 18). LDH

liver pattern has not been tested for in this taxon.

Two other synapomorphies appear to be: 1) the form of the

hyomandibular, which in the majority of gadoids and mac-

rouroids has attrition of the anterior border and lateral face,

fully or partly exposing the pathway of the hyoid branch of

the facial nerve (Howes, 1989; 1991b; 1992); 2) the levator

arcus palatini covering the adductor mandibulae musculature

laterally (Howes, 1988; 1991b).

Melanonids have a small intercalar, a bone which in other

gadiforms contributes to a substantial part of the lateroposte-

rior cranial wall. In size the melanonid intercalar approaches

that of Percopsis. Whether in Melanonus the bone is plesio-

morphically small or whether there has been reduction sec-

ondarily can only be assessed against the distribution of

other, known derived features (Fig. 18). An intercalar is

absent in lophiiforms and batrachoidiforms, an assumed

secondary loss (Patterson & Rosen, 1989).

The single dorsal fin is probably a plesiomorphic feature

(p. 26). Among paracanthopterygians, an elongate second

dorsal fin is assumed to be synapomorphic for anacanthines

(sensu Patterson & Rosen, 1989). Melanonids share with

ophidiiforms (including carapids and bythitoids), two gadoid

genera (Lyconus and Brosme) and Macrouroididae a single

dorsal fin which must be seen as resulting from either the

‘loss’ of the first dorsal with anterior encroachment of the

second, or the amalgamation of the two fins. It is impossible

to distinguish between such phylogenetic events although

either way the condition is seen as derived. Iwamoto (1989)

considered the single dorsal fin of macrouroidids to be

derived but that of the gadoid Brosme as plesiomorphic

retention. Judging by the incongruent distribution of the

character it is almost certainly homoplastic. The further

partitioning into three fins in Gadidae represents a further

derived state.

In jaw musculature melanonids are little different from

morids and bathygadids (Howes, 1988). Howes (1990; 1991b)

noted a medial shift of adductor muscle Alb which would

suggest a close phylogenetic relationship with supragadoids.

This shift, however, is apparently induced by the presence of

a unique transverse ligament which runs from the palatine to

the inner face of the second infraorbital and which constricts

and turns Alb inwards. This is not the same condition as the

entire medial shift of an unconstricted Alb in ‘supragadoids’.

Melanonids have an unusual condition of the hyoid muscu-

lature whereby the pharyngohyoideus (= rectus communis)

attaches to the third hypobranchial as well as the urohyal

(Howes, 1988). Urohyal attachment of the pharyngohyoideus

is shared with macrouroids, two gadoid families and all other

ctenosquamates (Lauder, 1983; Howes, 1988); in remaining

gadoids the pharyngohyoideus is mediated by the sternohyoi-

deus. It is assumed that the two exceptional gadoid families

(Muraenolepididae and Ranicipitidae) have lost the sternohy-

oideus attachment, the pharyngohyoideus being attached to

the tip rather than the lateral face of the urohyal keel as it is

plesiomorphically in melanonids.

The melanonid caudal fin musculature (p. 26) lacks those

features regarded as synapomorphic for gadoids (since mac-

rouroids lack caudal fin skeletons and associated musculature

it cannot be known whether this derived form of muscle

arrangement was a gadiform feature subsequently lost in

macrouroids). Melanonids have a caudal fin muscle arrange-

ment only slightly modified from that present in other para-

canthopterygians and in acanthopterygians.

Although it cannot be doubted that the Melanonidae

belongs among Gadiformes there is no evidence to suggest

that it be regarded as a member of the Gadoidei. To be

included within the Gadoidei, the elongate toothed palatine,

lack of X and Y bones, reduced intercalar and single dorsal

fin must be regarded as reversal and loss characters. The

caudal fin skeleton demonstrates an advanced condition to

that of the Moridae (the plesiomorphic gadoid taxon) in

G.J. HOWES

having, in adults, almost complete fusion of the upper

hypurals which alone, would signify inclusion within the

‘supragadoids’. Indeed, I have argued elsewhere (Howes,

1991b: caption to fig. 35) that the reported separation of

hypurals in young ranicipitids (which I place amongst the

‘supragadoids’) is a character reversal; a conclusion drawn on

what appears to substantial support from other synapomor-

phies. In the case of melanonids the principal evidence

against the caudal fin skeleton being a character reversal is

that the associated musculature has a plesiomorphic arrange-

ment, lacking those derived elements found in the muscula-

ture of morids and other gadoids, including ranicipitids (as an

adult, Raniceps has the typically symmetrical gadoid caudal

fin skeleton, lacking in Melanonus). Thus the fusion or partial

fusion of the upper hypurals in melanonids is considered to

have occurred independently to that in gadoids above the

morid level.

Taking into account these arguments and the anatomical

evidence presented herein, the Melanonidae is regarded a

basal gadiform taxon, representing, as Markle (1989) had

previously hypothesised, the sister-group to both gadoids and

macrouroids (Fig. 18). Such a phylogenetic arrangement

leads to a higher level re-classification of the Melanonidae.

Following Markle (1989) and recognising the family as being

phylogenetically coordinate with the Macrouroidei and

Gadoidei, it is placed in the suborder Melanonoidei. Those

taxa which I have previously recognised as a monophyletic

group termed ‘supragadoids’ are equivalent to Markle’s

(1989) Superfamily Gadoidea. The Moridae and Euclichthy-

idae are regarded by Markle (1989) as sister taxa on the basis

of asymmetry of procurrent caudal fin rays; I know of no

supporting osteological synapomorphies for this relationship

but provisionally accept it. Together these taxa form the

sister-group to the Gadoidea and as such must be regarded as

the Superfamily Moridoidea (= Moriformes,

part,Schwarzhans, 1984). The ‘infragadoids’, Steindachneri-

idae and Bathygadidae have no such status since they form an

unresolved polychotomy with the Gadoidei + Moroidei and

Macrouroidei.

ACKNOWLEDGEMENTS. My thanks are due to Douglas Markle, Nigel

Merrett and Colin Patterson for their helpful and critical comments

on the manuscript of this paper. Since this is the final research project

carried out while employed at The Natural History Museum I take

this opportunity to thank all members of staff, associates and

students past and present of the Fish Section for their advice,

assistance, patience, generosity and friendship over the past twenty-

five years. I am particularly indebted to my mentors, P. Humphry

Greenwood and Ethelwynn Trewavas.

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Bull. nat. Hist. Mus. (Zool.) 59(1): 33-44

Issued 24 June 1993

A review of the serranochromine cichlid fish

genera Pharyngochromis, Sargochromis,

Serranochromis and Chetia (Teleostei: Labroidei)

PETER HUMPHRY GREENWOOD

Visiting Research Fellow, Zoology Department, The Natural History Museum, Cromwell Road,

London SW7 5BD; Honorary Associate, J.L.B. Smith Institute of Ichthyology, Private Bag 1015,

Grahamstown 6140, South Africa.

CONTENTS

Re CAA CON OL ee Saat a aes eet aot ence a oat can ac carc sn in co MR Salad Ansan sn einala o's oe ss dee niSds v's anaicesaangedsinnaes 33

MVELIO CS ARC MAT CSU AN sect tena nn cabelas aia eB oo 6 cr,o mae gee dal sla nad alas dainty s Hanan gadis dpldelsrnlesia not asinamnsnssig Aba 34

ERE AR OCHO TLIC CN COT Vinee ead oe sistara emis ada ciceclacie iin « 6.0. Seah soln sige asic soa wilocb sie a nape. oids/naaiet< <niadasica sine serene Gaunyc’s ae 36

ITER TCA PL GUN CRE leer Mee iced elo cn Cea odin decid w Snir Sv alindriaslnccuasen sauce nea sinass scnacine 36

The generic or subgeneric status of Serranochromis Regan, 1920, and Sargochromis Regan,

PAVE LE COLSICORCOME he tr Mee tata ca cnaes cam cmacslccnacs pias ac ame se tee eM abiasite ana sida at a Sines waptaue aeinels lass doles elaKoadts eeretomutieiatg 36

PWC RCS CHEM APEC WAV AS DEO OM tna tana ares se sita nie « inv cia con een amee Oca eae opie semaines dakiiwawnele elie - se qniaist sie weit 38

SU CSDeOIISEE MARV ILCERLONMS GLCCHWOOU., LIT 9L s sccnc: ccacsacacmaninds diseh ccna th shiddeadcerecctctessaceceassesessneseascnncers 39

MOG ASIC Ee met eiat antes tee Mtae tet saned este cde tne ston sintewiretss.ctvarcarocterttenuctvetccdcosentasccncstaccerseenc dere 40

Miemhvietic relationships Of the SerranOCHTOMIMES .......tesaeccsseccsvccecssccecetstavccrtaccstasecccetacteccstaccccssaness 40

PRE OEMe OV: AUG GAIABNOSCS re. tet ceeded: Jct oP a AON aE wna vos Renee ra ahs eran tah onnna dt Saccacaceeaeetecdeectersersdestevcete. 41

PREM CMICCOCEICIIIS! Abang te cee crete na eed ects. coaaED Neds a vnwsinsteyeeeee ae Cts aeeraage ton Acheck race tebiiars dec teresunedlavanedenens 43

ENS LREMICOS Met. tea na soice.« Aeriaas teamed detain delineate tadedtns sow scttber dove seddedehasesestes icven tees seencdcnrecdetursedistastesstenl 43

Synopsis. Recent taxonomic changes, newly described taxa and groups of taxa, and the introduction of taxonomic

characters not previously employed require a revision of the informally recognised, essentially fluviatile group of

southern African cichlids, the so-called serranochromines. The genera included in this assemblage are Pharyn-

gochromis, Greenwood, Sargochromis Regan, Serranochromis Regan, and Chetia Trewavas.

Previously, Sargochromis was considered to be a subgenus of Serranochromis, but new evidence indicates that it

should be reinstated as a distinct lineage (i.e. genus). The species originally described as Serranochromis

(Sargochomis) gracilis should now be transferred to the genus Chetia. Formerly the latter taxon was thought to be

monotypic, but it is now expanded to include five species. One of these, Chetia brevis Jubb, had been included

tentatively in the genus Astatotilapia, but is now returned to the genus in which it was described originally.

The monophyletic origin of the serranochromines has still to be established. For that, and other reasons, doubt is

cast on the phylogenetic reality of the ‘Pharyngochromis — Chetia — Serranochromis’ group of endemic species in

Lake Malawi. The suggested interrelationships of the serranochromine genera presented below, and based on

shared derived characters, cannot, for the same reasons, be considered a truly phylogenetic one.

INTRODUCTION

In a paper (Greenwood, 1979) reviewing and reconsidering

the generic classification of several cichlid taxa then referred

to the genus Haplochromis, an informal group of three

genera was recognised on the basis of its constituent species

having particular types of squamation and anal fin markings

(op. cit: 229-316). The group was, and still is considered one

of convenience because no cladistically based hypothesis

could be erected to establish the monophyly of its contained

genera, viz. Serranochromis Regan, 1920 (with which was

incorporated, as a subgenus, Sargochromis Regan, 1920),

Chetia Trewavas, 1961, and Pharyngochromis Greenwood,

1979. A scheme of possible interrelationships of these taxa,

suggested earlier by Trewavas (1964), was also discussed in

my 1979 paper.

Recent studies of the genera call for a revision of the

group’s taxonomy at the generic level, and a reconsideration

of their possible interrelationship. For example, the genus

Chetia, treated as monotypic by Greenwood, (1979) is now

thought to contain five species (Balon & Stewart, 1983;

Greenwood, 1984, 1992, and below, p. 38), certain problems

regarding the generic classification of several Angolan hap-

lochromine species have been clarified (Greenwood, 1979,

1984 & 1992), new ideas on the supposed relationship of

Serranochromis and Sargochromis have been put forward by

Lippitsch (1991: 99-100), and Eccles & Trewavas (1989: 21)

have formally recognised, amongst the endemic genera of

Lake Malawi, a large assemblage of species which they

designate ‘The Pharyngochromis — Chetia — Serranochromis

group’.

The term ‘serranochromine’ will be used in this paper as a

group name for the genera Chetia, Serranochromis, Sar-

gochromis and Pharyngochromis. Its use should not be

construed as an indication or even a presumption of the

group’s ultimate and formal recognition as a Tribe. In the

sense employed here it is comparable with my earlier use of

the informal categories ‘haplochromines’ and ‘pelmatochrom-

ines’ (Greenwood, 1979 & 1987). Such continued use, and

introduction, of informal groupings clashes with the tribal

status given by Poll (1986) to several cichlid assemblages in

Lake Tanganyika, and with the geographically and taxonom-

ically more extensive tribe Haplochromini defined by Eccles

& Trewavas (1989), a tribe which also includes the serrano-

chromines. In my view, these authors’ actions are premature.

Too few critical higher-level taxonomic studies have yet been

made on the Cichlidae to support the phylogenetic relation-

ships that are (or should be) implicit in the award of formal

tribal status. For instance, Eccles & Trewavas (1989: 21)

define the Haplochromini as: ‘Maternal mouth-brooding

cichlid fishes of Africa and the Jordan Valley in which the

basioccipital bone participates with the parasphenoid to form

the apophysis for the upper pharyngeal bones”’. The value of

the apophyseal character has been questioned by several

authors (see review in Greenwood, 1978, also Greenwood

1986) and it may have evolved more than once among

African taxa (Greenwood, 1987); maternal mouthbrooding

has apparently evolved independently in both the Tilapiini

and Haplochromini (the tribes, respectively, sensu Trewavas,

1983, and Eccles & Trewavas, 1989), and in one species of the

genus Chromidotilapia of the pelmatochromines (sensu

Greenwood, 1987: 169) in which paper it is also argued (op

cit: 194-199) that this group should not be included, as it was

by Trewavas (1983), in the tribe Tilapiini.

Thus, the purpose of the present paper is simply to clarify

the taxonomic status of the ‘serranochromine’ genera, and to

establish a basis for further and phylogenetic studies of those

taxa and those of Lake Malawi.

METHODS AND MATERIALS

ANAL FIN SPOTS AND TRUE OCELLI. One of the features used

to define the serranochromines is the presence of maculae on

the anal fin, usually in both sexes (Greenwood, 1979). A

distinction was made there between true ocelli (such as occur,

but almost exclusively in males, in a large number of hap-

lochromine species, e.g. those in Lakes Victoria, Edward and

Kivu), and the spots or maculae found in the serranochrom-

ines and the haplochromines of Lake Malawi (see figure in

Eccles & Trewavas, 1991). Judging by a recent description of

a new Serranochromis species (Winemiller & Kelso-

Winemiller, 1991) it is clear that some confusion still exists

when discriminating between these two kinds of anal fin

markings. Granted, it is often difficult to do so when only

preserved material is available, but in life the difference is

obvious, as colour photographs in aquarium books will show

(e.g. Konings, 1991).

The densely pigmented ovoid or near circular centre of the

true ocellus, usually circumscribed by a narrow, darkly pig-

mented ring, is surrounded, or almost surrounded, by a zone

of virtually transparent, or at least freely translucent, and

P.H. GREENWOOD

apparently unpigmented fin membrane. This clear zone is of

variable width and outline, but is often concentric with that of

the pigmented centre. In life, the clear zone seems to

emphasise the coloured central area, thereby making it stand

out from the rest of the fin membrane, be that membrane

pigmented or hyaline. In colour photographs of live or freshly

dead specimens, the clear zone often appears to be dark or

even black, a result either of the dark background against

which the fish was posed, or the shadow cast by the fish’s

body and the anal fin itself. To the best of my knowledge,

true ocelli do not occur, at least in nature, on any of the other

paired fins, although these fins are often maculate. Anal

ocelli are also of rare occurrence in females, but large

non-ocellate spots are sometimes present on that fin in the

females of species whose males have true ocelli. The spots in

such females occupy the same position as the ocelli in males,

and are often of the same size. As compared with maculae

(i.e. non-ocellate spots) on the anal fin, true ocelli are

generally larger, and are always readily distinguishable from

the maculae on the other unpaired fins.

In contrast, non-ocellate anal spots, besides usually being

smaller than ocelli, are, with few exceptions (see Greenwood,

1992) more numerous and differ little in their overall appear-

ance from those on the other fins, although the central

pigmented portion may differ quite markedly in colour. The

essential difference between maculae and ocelli, however,

lies in the absence of a transparent or freely translucent area

surrounding the pigmented centre of a macula, which, like

that of an ocellus, is bounded by a very thin ring of dark

pigment. Instead, the macula’s pigmented centre is circum-

scribed by a ring, usually narrow, of lighter pigment which

separates it from whatever ground colour the fin membrane

may have.

This outer, lightly pigmented ring is not visible in some of

the preserved serranochromine specimens I have examined,

and the central spot is bounded only by the very narrow ring

of dark pigment separating it from the chromatophores in the

fin membrane. Whether or not this situation is a preservation

artefact cannot be determined at present.

In their account of the anal fin markings in the newly

described species Serranochromis altus, Winemiller & Kelso-

Winemiller (1991: 679) describe, unfortunately without an

illustration, the fin in both sexes as having “. . .30-40 large

pink or pink-orange ovate spots, each ringed with a transpar-

ent, white ocellus. . .”. I would suggest that the use of the

words ‘transparent and white’ in apposition is somewhat

contradictory, and that ‘translucent white’ would describe the

condition more accurately, especially since it is one I have

seen in fresh specimens of Serranochromis (and Sar-

gochromis) species from the Okavango river and swamp

system in Botswana.

PRESHANK LENGTH OF THE MAXILLA; LENGTH OF THE PRE-

MAXILLARY ASCENDING AND ALVEOLAR PROCESSES. The

preshank length of the maxilla, relative to its shank length,

and the length of the alveolar process of the premaxilla

relative to the length of the entire ascending premaxillary

process, are two morphometric characters not used in earlier

papers (Greenwood, 1979, 1989, 1981).

Preshank length of the maxilla is measured, on the bone’s

medial aspect, from the anterior tip of the medial arm of the

maxilla’s saddle process, to the mid-point of the anterior

vertical projection from the upper margin of the bone’s shank

(see Fig. 1). Shank length is measured, also directly and on

THE SERRANOCHROMINE GENERA REVIEWED

the bone’s medial aspect, from the midpoint of the vertical

projection to the posterior point on the maxilla’s posterior

margin (see Fig. 1).

———————————

Fig. 1. Medial aspect of maxilla (from Serranochromis

macrocephalus), viewed somewhat dorsolaterally, to show points

of measurement for: A, preshank length, and B, shank length.

Scale bar = 5 mm.

The length of the premaxilla’s ascending process is mea-

sured, directly, from the dentigerous surface of the bone at its

symphysis with the premaxilla of the opposite side, to the

dorsal tip of the process. The length of the alveolar process

uses the same ventral (i.e. dentigerous surface) reference

point; its dorsal limit being the highest point on the dorsal

margin of the process (see Fig. 2).

STUDY MATERIAL

See Greenwood (1979, 1984, 1992) for specimens used in

previous studies of Serranochromis, Sargochromis, Chetia

and Pharyngochromis. Additional material:

Sargochromis coulteri RUSI 26627. Namibia (4 specimens)

Sargochromis coulteri RUSI 36419. Namibia (1 specimen)

Sargochromis coulteri RUSI 28106. Namibia (1 specimen)

Sargochromis giardi RUSI 35782. Zambia (3 specimens)

Sargochromis carlottae RUSI 31204 Namibia (3 specimens)

Sargochromis carlottae RUSI 31214 Namibia (2 specimens)

Sargochromis coulteri RUSI 31199. Namibia (1 specimen)

Serranochromis robustus RUSI 31169. Botswana; Okavango

river (1 specimen)

Serranochromis thumbergi RUSI 22660. Dam, Empangeni

area, Natal R.S.A. (1 specimen presumably an introduc-

tion)

Serranochromis longimanus RUSI 23877. Botswana; Oka-

vango swamps (6 specimens)

Serranochromis macrocephalus RUSI 24175. Botswana; Boro

river (9 specimens)

Serranochromis angusticeps RUSI 26854. Okavango swamps

(4 specimens)

Chetia brevis Holotype AMG. P. 951. Lomati river, R.S.A.

Chetia brevis Paratypes AMG. P. 952. Lomati river, R.S.A.

(5 specimens)

Chetia brevis AMG. P. 1422. Lomati river, Barbeton district,

R.S.A. (3 specimens, one partly skeletonized; see below)

Chetia flaviventris AMG. P. 1298. Mogel river, Waterberg,

R.S.A. (7 specimens, one partly skeletonized; see below)

Chetia flaviventris AMG. P. 6871. Tweeport, Rustenburg

district, Limpopo system, R.S.A. (10 specimens)

Fig. 2.

Frontal view of left and right premaxillae (from

Serranochromis macrocephalus) to show points of measurement

for: A, height of ascending process, B, height of alveolar process.

Scale bar = 5 mm.

Chetia flaviventris Palala river at Muisvogelkraal (24° 00'S,

28° 24’ 30"E) R.S.A. (5 specimens)

Chetia mola Holotype. ROM 29825. Luonga river (Zaire

drainage), Zambia

Skeletal material

Pharyngochromis acuticeps RUSI 36553. Okovango river

(see also Greenwood, 1992)

Sargochromis carlottae RUSI 31204. Namibia

Sargochromis carlottae RUSI unregistered. 142 mm S.L.

Sargochromis giardi RUSI unregistered 210 mm S.L.

Sargochromis giardi RUSI unregistered 244 mm S.L.

Sargochromis cf Sargo. greenwoodi RUSI unregistered

Serranochromis macrocephalus RUSI 24175. Botswana, Boro

river

Serranochromis macrocephalus RUSI unregistered 107 mm

Sul?

Serranochromis macrocephalus RUSI unregistered 220 mm

Sik:

Serranochromis

220 mm S.L.

Serranochromis angusticeps RUSI unregistered 220 mm S.L.

Serranochromis angusticeps RUSI unregistered 230 mm S.L.

Serranochromis angusticeps RUSI unregistered 410 mm S.L.

Serranochromis longimanus RUSI unregistered 150 mm S.L.

Chetia flaviventris AMG P. 1298. Mogel river, Waterberg,

R.S.A.

Chetia brevis AMG P. 1422. Lomati river, R.S.A.

macrocephalus RUSI unregistered ca

Institutional abbreviations:

AMG: Albany Museum, Grahamstown

ROM: Royal Ontario Museum, Toronto

RUSI: J.L.B. Smith Institute of Ichthyology, Graham-

stown

SERRANOCHROMINE TAXONOMY

Introduction

In their revision of certain haplochromine genera from Lake

Malawi, Eccles & Trewavas (1991: 21) divide the non-

tilapiine taxa of that lake into three groups, one of which they

call the Pharyngochromis — Chetia — Serranochromis group.

That action I consider to be premature, both because there is

little concrete information available about the phyletic inter-

and intrarelationships of the Malawi species, and because, as

Eccles & Trewavas point out, there are differences between

the squamation of those species and that of the serrano-

chromines as construed in this paper (see p. 40). Granted, I

have suggested (Greenwood, 1979: 314) that Serranochromis-

and Chetia-like taxa could have been involved in the origin of

the Malawian cichlid flocks, but that idea was not put forward

on the basis of characters constituting a testable hypothesis.

Rather, it was intended, because of the superficial resem-

blance between the two groups, to promote an awareness that

possible intergroup synapormophies should be looked for in

future research.

As recognised in this paper, the serranochromines are an

assemblage of mainly fluviatile taxa whose geographical

range encompasses the Zambezi, Save-Runde, Limpopo,

Cunene, Quanza, Okavango and Zaire river systems, with

one species (Serranochromis robustus robustus [Ginther]),

occurring in Lake Malawi (see Balon & Stewart, 1983;

Bell-Cross, 1975; Eccles & Trewavas, 1989; Greenwood,

1984, 1992; Jubb, 1967, 1968; Ladiges, 1964; Poll, 1967;

Skelton, 1993, Trewavas, 1961, 1964).

Since the last published inventory of serranochromine

species (Greenwood, 1979), revisional studies (Greenwood,

1984, 1992) and the description of new species (Balon &

Stewart, 1983, Winemiller & Kelso-Winemiller, 1991, and

Greenwood, 1984 [see p. 38 below]) have both increased the

number of included taxa and extended the geographical range

of the group.

Morphologically, the two distinguishing features of the

serranochromines are the following. (i) The presence, often

in both sexes, of non-ocellate maculae (see p. 34) on the anal

fin. Generally these spots are very numerous with, in certain

species, as many as 30-40 covering almost the entire fin.

However, their number and size show considerable intraspe-

cific (and intergeneric) variability, with as few as three or four

large spots occurring in some individuals of a species where

the maximum number is 18-20 (Greenwood, 1992). When

many spots are present their arrangement may give the

impression of an irregular distribution on the fin, but (as was

noted by Oliver [1984], pace Greenwood, 1979: 315) there is

a basically linear regularity in their arrangement. (11) All

scales above the lateral-line series are cycloid, as are the

majority of scales below that level. Some weakly ctenoid

scales may occur anteriorly on the flanks, especially in small

specimens, the ctenii being confined to a narrow arc situated

near the centre of the scale’s free margin.

A third but less trenchant feature of the serranochromines,

as compared with other fluviatile non-tilapiine and non-

pelmatochromine taxa (both sensu Greenwood, 1987:

194-199) is a tendency for there to be a higher modal number

of abdominal vertebrae (modes 16 or 17 in serranochromines,

[but 14 in one taxon] cf 12 or 13 in the other taxa); however,

P.H. GREENWOOD

the ranges for total vertebral counts in the two groups

overlap.

Where information is available on breeding habits, the

serranochromine species are known to be female mouth-

brooders, and in all taxa the neurocranial apophysis for the

upper pharyngeal bones is formed from the parasphenoid and

the basioccipital bones. In those respects, the group would

conform with Eccles & Trewavas’ (1989) definition of the

tribe Haplochromini.

Recently, Lippitsch (1991) drew attention to the possible

value of scale morphology and patterns in resolving certain

problems of intergeneric relationships within the serrano-

chromines. To determine if the features described by Lip-

pitsch could also provide another ‘group’ characteristic, I

examined, at a magnification of 50x, the superfical morphol-

ogy of flank and other scales in several species belonging to

both subgenera of Serranochromis (sensu Greenwood 1979;

but see p. 37), all species of Chetia, and in different popula-

tions of the single Pharyngochromis species. In general I

found a fairly high level of intrageneric variability, as well as

some individual variability in the features noted by Lippitsch

(1991: 99-100; figs D & E) i.e. ornamentation of the caudal

field, and the presence of a soft caudal rim to the scale.

Further studies will be necessary before the value of these

features can be established, both at the group and lower

levels of serranochromine taxonomy. However, another

squamation feature noted and discussed by Lippitsch, namely

the number of scale rows between the posterior orbital

margin and the preoperculum, has proved of considerable

value in reviewing generic level taxonomy within this group.

The generic or subgeneric status of Serranochromis

Regan, 1920, and Sargochromis Regan, 1920,

reconsidered.

In an attempt to revise the Haplochromis generic concept on

phyletic lines, several so-called Haplochromis species were

assigned, as a subgenus, to the genus Serranochromis (Green-

wood, 1979). Since one of these species is the type of Regan’s

genus Sargochromis (S. codringtoni [Blg, 1908]) that name

was resurrected for the new subgenus.

The principal argument for this taxonomic rearrangement

was that the taxa included in the new concept of Serrano-

chromis all share what appeared to be three derived features,

viz: an increased number of abdominal vertebrae, higher

gill-raker counts, and an increased number of branched rays

in the dorsal fin (Greenwood, 1979: 299).

In the light of new data, especially those stemming from an

increased knowledge of the genus Chetia (see p. 38) and,

particularly, Lippitsch’s (1991) observations on the different

type of postorbital squamation patterns present in the two

presumed subgenera, I would now revise my earlier action

and recognise both Serranochromis and Sargochromis as

distinct lineages, and thus accord each generic status. The

reasons for that action are as follows.

Firstly, as Lippitsch (1991) noted, in Serranochromis but

not in Sargochromis there are at least two, and often more,

vertical rows of scales between the posterior orbital margin

and the upper part of the preoperculum’s ascending arm. In

Sargochromis only a single row is present. A double row,

however, also occurs in Chetia (personal observations, see

below). Further investigation of both Serranchromis and

Chetia reveals that the double and multiple scale row condi-

tion is correlated with underlying osteological and myological

THE SERRANOCHROMINE GENERA REVIEWED

characters that are not present in Sargochromis or Pharyn-

gochromis, the two other serranochromines with only a single

postorbital scale row. These correlated features are an

increase in the relative size and bulk of the upper part of the

levator arcus palatini muscle, and an anteriorly directed

lengthening of the postorbital process from the sphenotic

bone, particularly its ventral region associated with the origin

of the muscle (see fig. 3 and fig. 8 in Greenwood, 1992). In

Sargochromis and Pharyngochromis the muscle is relatively

smaller, and the postorbital sphenotic process has the almost

uniformly narrow form (Fig. 3) found in the generalised

haplochromine skull (and, it may be added, the skulls of

Tylochromis and Heterochromis, the genera thought to repre-

sent the least derived lineages of African cichlids; see Oliver,

1984, and Stiassny, 1989 & 1991).

The other reason for reconsidering the subgeneric status of

Sargochromis is the increased information now available,

especially for Chetia (see p. 38), which clearly indicates an

extensive overlap in the gill-raker and dorsal fin-ray charac-

ters previously used (Greenwood, 1979) to define Serrano-

chromis (then including Sargochromis). These characters,

therefore, no longer can be considered synapomorphic for

Serranochromis and Sargochromis alone.

With the elimination of those two characters, the only

derived feature shared uniquely by Serranochromis and Sar-

gochromis is the increased number of abdominal vertebrae.

Against that presumed synapomorphy must be set the

derived postorbital scale-row character shared only by Serra-

nochromis (sensu Regan, 1920) and Chetia, which genera also

share two apomorphic features not previously recognised.

These are: (i) an increase in the range and modal number of

caudal vertebrae (range 15-18, modes 16 and 17 in Serrano-

chromis, range 15-17 modes 16 and 17 in Chetia, compared

with ranges of 12-16 in Sargochromis, and 14-16 in Pharyn-

gochromis, with modes of 14 and 15, and 15 in the genera

respectively); (ii) an increase in the range and modal number

of circumpeduncular scale, viz. 18-20, rarely 16, in Serrano-

chromis and Chetia, neither taxon with a clear-cut modal

number, as compared with 16 or 18 (mode 16) in Sar-

gochromis and 15 or 16 (mode 15) in Pharyngochromis. The

recognition of this feature as an apomorphy is based on the

circumpenduncular counts for Tylochromis and Hetero-

chromis (see above), where the range, and modes, are 15-16.

Thus, taking into account the presumed derived characters

shared only by Serranochromis and Chetia, it would seem that

the higher count of abdominal vertebrae in Serranochromis

and Sargochromis should be treated as a homoplasy and not,

as previously thought, an apomorphy indicating an immedi-

ate common ancestor for the two taxa. For that reason I agree

with, and now formally act upon Lippitsch’s (1991) conclu-

sion that “It seems advisable . . . to recognise Sargochromis

as a distinct genus. . .”’.

The possible relationship of Sargochromis within the serra-

nochromines is discussed on p. 41, and a revised generic

diagnosis is given on p. 42.

A few further comments need to be made about the genera

Serranochromis and Sargochromis. The number of postor-

bital scale rows in certain Serranochromis species can be as

high as four or five, and, as far as I can determine, the

number of rows is relatively constant intraspecifically. At

present insufficient information is available on possible inter-

or intraspecific differences in scale ornamentation, or on such

| variation in the presence or absence of a soft free margin to

| the scales (see Lippitsch, 1991). The small sample of species I

Fig. 3. Posterior portion of the neurocranium, in left lateral view,

to show differences in the size of the sphenotic postorbital process

(Sp.) in Sargochromis, Chetia and Serranochromis. A,

Sargochromis carlottae (RUSI: 31204, 89 mm S.L.; B, Chetia

brevis (AMG: P1422, 90 mm S.L.) and C, Serranochromis

macrocephalus (RUSI: 24175, 112 mm S.L.) Scale bar = 5 mm.

have examined from the Okavango swamp and river system

in Botswana seems to indicate that such variation may occur.

A soft margin, for example, was absent in specimens of

Serranochromis thumbergi and S. longimanus, but is present

in S. macrocephalus. From the same samples some individual

variability was noted both in the extent of the granular area

on the caudal field of flank scales below the upper lateral-

line, and the presence or absence of a granular area in at least

some scales above that line. Apparent interspecific variation

is also seen in the extent and nature of cheek, opercular and

interopercular squamation patterns. The value of these fea-

tures as a basis for intrageneric classification has, however,

yet to be tested on a wider geographical and taxonomic basis.

A feature of Serranochromis, Sargochromis and Chetia

noted in an earlier paper (Greenwood, 1979), where it was

ranked as an apomorphy, is a reduction in the maximum

number of inner tooth rows in both jaws to a single or

irregularly double series. The new species, S. altus, described

by Winemiller & Kelso-Winemiller (1991), is exceptional in

having as many as six rows anteriorly, reducing to a single or

double row posteriorly (the figures adjusted from Winemiller

& Kelso-Winemiller who include the outer tooth row in their

counts). These authors also note that the number of inner

rows increases with growth “. . .in all species of Serrano-

chromis that we have examined. ..”. Their list of study

material indicates, however, that only one other species, S.

angusticeps, was studied, and that the maximum number of

inner or outer rows in that species is three. Since museum

specimens are often not fully representative of a species’ full

size-range, the possibility of growth related changes in the

number of tooth rows should perhaps be treated, for the

moment, as an open question. In that connection, however, it

should be noted that in Sargochromis, one species, Sargo.

thysi, has four inner rows in both jaws at a size when only one

or two rows are present in other congeneric species.

Another dental feature in Serranochromis, one also shared

with Sargochromis, (and probably Chetia), is for the two

median teeth in the outer row of the inner tooth series to be

enlarged and displaced anteriorly (Greenwood, 1984: 216;

also see figs 3 & 8 in Trewavas, 1964). Amongst the sample of

Serranochromis specimens examined, which included all spe-

cies of the genus except S. altus. (see p. 35, and Greenwood,

1979 & 1984 for additional material), such tooth displacement

occurs in most species but not in all individuals of a species.

Its frequency of occurence is lower in Sargochromis than in

Serranochromis, and the condition is known from only one

Chetia species (C. gracilis; see below and p. 39). Because of

this inconstancy in its expression I would now consider the

character more in the nature of a trend, albeit a derived one,

rather than the trenchant synapomorphy recognised earlier

(see Greenwood, 1984; 216).

A revised generic diagnosis for Serranochromis is given on

p. 42, and the possible relationships of the genus are dis-

cussed on p. 40. To the list of included species published in

Greenwood, 1979 (pp. 302-303) must be added S. altus

Winemiller & Kelso-Winemiller (1991) from the Zambezi

system.

Little need be added to previous accounts of the genus

Sargochromis, type species Paratilapia codringtoni Blgr.,

1908 (see Greenwood, 1979; 1984, in both papers the taxon

treated as a subgenus) except to note that the species

described as Serranochromis (Sargochromis) gracilis Green-

wood (1984) from the Cunene river, is now considered to be a

member of the genus Chetia (see p. 39). The transfer of this

P.H. GREENWOOD

species reduces to two the number of Sargochromis species

having, relative to other congeneric taxa, slender lower

pharyngeal bones with few and only partially molariform

teeth, viz. Sargochromis greenwoodi (Bell-Cross) and Sarg.

coulteri (Bell-Cross). The latter species, however, appears to

show considerable variability in these features, with some

individuals having noticeably coarser lower pharyngeal bones

than do others, a feature invariably correlated with an

increased number of molariform teeth (Greenwood, 1984:

217-221). As was discussed in that paper, the species level

taxonomy of Sargochromis is far from satisfactory, so this

seemingly intraspecific variability should only be accepted

with some reservation (see also Greenwood, 1965 and

Hoogerhoud, 1986).

In lacking a pronounced ventral expansion of its sphenotic

postorbital process(see p. 37), the neurocranium in Sar-

gochromis is immediately distinguishable from that of Serra-

nochromis (see Fig 3; also comments in Greenwood, 1979:

303, and figs 13 & 16).

A list of Sargochromis species is given in Greenwood,

(1979, pp. 304-305), and a revised generic diagnosis on p. 42

below.

The genus Chetia Trewavas, 1961

TYPE SPECIES: Chetia flaviventris Trewavas, 1961.

It has not proved possible to formulate a trenchant generic

definition for Chetia since the taxon has yet to yield a single

diagnostic autapomorphy. Its definition, therefore, is based

on, as it were, negative features, namely those which exclude

the five member species from inclusion in any of the three

other serranochromine genera, especially Serranochromis

with which Chetia has the greatest superficial and some

detailed similarity. Thus, although Chetia shares with Serra-

nochromis the apomorphy of two postorbital scale rows (see

p. 36), a high number of caudal vertebrae (15-17, modes 16

or 17) and an increased number of circumpeduncular scales

(18 or 20, rarely 16), it does not share with Serranochromis

apomorphies of an increased number of abdominal verte-

brae, (and as a consequence, an increase in the total number

of vertebrae), nor the increased lateral-line scale count of

that genus (35-41, a character probably correlated with the

increased number of vertebrae). Neither do the two genera

share the autapomorphic dental character of Serranochromis,

namely the development of a totally unicupsid jaw dentition

at a very small size; i.e., at some length, yet to be determined,

less than 29 mm S.L.; see Greenwood, (1979: 300).

On the basis of that analysis, the simplest diagnosis for

Chetia is: a Serranochromis lacking the apomorphic features

of that genus (see also Greenwood, 1992: 49-50 and p. 43

below).

At the time of my earlier generic review (Greenwood,

1979), Chetia comprised two species, namely the type, Chetia

flaviventris, and Chetia brevis, Jubb, 1968; the species coming

from the Limpopo and Incomati river systems of South Africa

respectively.

Based on Jubb’s original description together with an

examination of the holotype and two other Chetia brevis

specimens, I excluded, in my 1979 paper, the species from

Chetia and placed it, as incertae sedis, in the genus Astatotila-

pia (Greenwood, 1979: 284 & 307). That decision was based

on three supposed features of C. brevis. (i) Jubb’s (1968)

description of the anal fin markings as ocelli, together with

THE SERRANOCHROMINE GENERA REVIEWED

his and my personal observations of their large size and

restricted number (3 or 4) as compared with the small and

numerous anal spots in other serranochromines (see p. 34),

especially Chetia flaviventris. (ii) Jubb’s (op. cit) statement

that the scales are ctenoid, and its implication that such scales

are the predominant form. (ili) The presence of bicuspid

teeth in the outer oral tooth rows of some specimens more

than 100mm S.L., contrasting with the situation in Chetia

flaviventris where, on the information then available, few

bicuspids were thought to be present in fishes 71-85 mm

S.L., and only unicuspids occurred in specimens above that

length.

Having now been able to examine colour-transparencies of

live C. brevis, it is clear that the anal spots are not true ocelli

(as defined on p. 34), but are large versions of the maculae

found in other serranochromines, including Chetia flaviven-

tris. The taxonomic significance to be attached to their large

size, and small number, cannot yet be assessed. However, a

wide size-range of anal maculae, their size negatively corre-

lated with their number, occurs in the monotypic serrano-

chromine genus Pharyngochromis, some individuals of which

have spots as large as those in Chetia brevis.

The larger collection of C. brevis specimens now available,

together with a re-examination of the entire type series,

shows that Jubb’s blanket description of the scales as ctenoid

is somewhat misleading. As in Chetia flaviventris, the scales

of C. brevis are cycloid above the lateral-line, and mostly

cycloid below that level. A variable number, usually small, of

ctenoid scales is present anteriorly on the flanks, such scales

being most numerous in fishes less than 80 mm S.L. In other

words, the same pattern as occurs in Chetia flaviventris is also

found in C. brevis.

An examination of larger samples of both C. flaviventris

and C. brevis also revealed that bicuspid teeth can be present

in idividuals of C. flaviventris up to a standard length of

90 mm, and that true unicuspid teeth as well as very weakly

shouldered (almost unicuspid) teeth occur in C. brevis speci-

mens between 80 and 90 mm S.L. This situation severely

weakens my third reason (see above) for excluding C. brevis

from the genus Chetia.

Furthermore, the presence of two vertical rows of postor-

bital scales in both Chetia brevis and C. flaviventris (see

p. 36), a feature previously overlooked by all workers, pro-

vides additional evidence for returning the species ‘brevis’ to

the genus in which it was originally, and I now acknowledge

correctly placed by Jubb (1968).

Since 1979 another two species have been included in the

genus, namely the taxon Chetia mola described by Balon &

Stewart (1983) from the Zaire river system, and Hap-

lochromis welwitschi (Blgr.) from Angola (see Greenwood,

1979 & 1984).

Chetia mola differs from all other congeneric species in

having a greatly enlarged lower pharyngeal bone with a

heavily molarized dentition (see fig. 12 in Balon & Stewart

1983), a type of pharyngeal mill more usually associated with

species of Sargochromis. However, unlike members of that

genus, C. mola has a double or sometimes triple row of

postorbital scales, and a lower number of abdominal verte-

brae.

To the existing four species of Chetia, I would now add a

fifth, a species from the Cutato river, Angola, originally

described (Greenwood, 1984) as Serranochromis (Sar-

gochromis) gracilis. The reasons for this transfer are the

double row of postorbital scales in ‘gracilis’, the persistence

of compressed, bi- and weakly bicuspid teeth in the outer

tooth row of both jaws in specimens over 100 mm S.L. and of

tricuspid teeth in the inner tooth rows in such invididuals (see

Greenwood, 1984: 227, and above), and the presence of only

15 abdominal vertebrae, a combination of derived and plesio-

morphic characters not occurring in any known Sargochromis

or Serranochromis species.

On the basis of comparisons with the few specimens

available, Chetia gracilis (n=2) differs from the only other

Chetia species recorded from southwestern Africa, C. wel-

witschi (n=5), in having a narrower interorbital width

(18.2-18.6 cf 21.0-23.3% of head length in C. welwitschi), a

larger eye (25.0-25.6, cf 18.6-22.25% head; the effects of

allometry are unlikely to account for this difference since the

two samples overlap in the size range of individuals repre-

sented), and a shallower cheek (22.7-23.3 cf 32.0-33.3%

head). The lower pharyngeal bone in C. gracilis is somewhat

stouter, and its median teeth coarser than in C. welwitschi (cf

figs 19 & 20 in Greenwood, 1984).

In its morphometric and meristic features, C. gracilis

closely resembles C. flaviventris and C. brevis, species respec-

tively from the Limpopo and Incomati river systems in South

Africa. From Chetia brevis, C. gracilis is distinguished by its

narrower interorbital width (18.2-18.6 cf 23.0-26.0% head

length), and from C. flaviventris, especially when specimens

of approximately the same size are compared, by its shal-

lower cheek (22.7—23.3 cf 25.5—-33.6% head). It also seems

likely that, when the effects of allometric growth are taken

into account, the eye diameter is greater in C. gracilis than in

C. flaviventris. The teeth in the median row of the lower

pharyngeal bone of C. gracilis are coarser and stouter than

those of C. flaviventris, in that respect being more like the

teeth in C. brevis (cf fig. 19 in Greenwood, 1984, fig. 19 in

Greenwood, 1974, and fig. 4B in Jubb, 1968).

Regrettably, apart from Chetia flaviventris (see du Plessis

& Groenewald, 1953; Trewavas, 1961) and C. mola (see

Balon & Stewart, 1983; fig. 10b) little or nothing is known

about the live coloration of the other Chetia species.

A revised generic diagnosis for Chetia is given on p. 43.

Included species:

Chetia flaviventris Trewavas, 1961. Limpopo river system.

Chetia brevis Jubb, 1968. Incomati river system

Chetia mola Balon & Stewart, 1983. Luongo river, Zaire

system.

Chetia welwitschi (Boulenger), 1898. Cunene and Zaire river

drainage systems, Angola (see Greenwood, 1979).

Chetia gracilis (Greenwood), 1979. Cutato river (Cubango

drainage system), Angola.

The genus Pharyngochromis Greenwood, 1979.

TYPE SPECIES: Pelmatochromis darlingi Boulenger, 1911.

A detailed revision of this monotypic genus, together with

an annotated synonymy, has been published recently (Green-

wood, 1992). Three nominal species all classified in the genus

Haplochromis since Regan’s revision of 1922 (or in one case,

Pharyngochromis), viz. Pelmatochromis darlingi Boulenger,

1911; Chromis jallae Boulenger, 1896, and Pelmatochromis

multiocellatus Boulenger, 1913, are now treated as junior

synonyms of the single Pharyngochromis species, P. acuticeps

(Steindachner) 1866.

There are some indications that P. acuticeps could be

considered either as a superspecies composed of several

topospecies or as an assemblage of evolutionary species, but

no clear-cut features allowing a formal taxonomic division of

the taxon can be identified (see Discussion in Greenwood,

1992).

Anatomically and morphologically, Pharyngochromis is

the least derived member of the serranochromines. Its sole

autapomorphic feature is the higher position occupied by the

posterior scales of the upper lateral-line series relative to the

base of the dorsal fin (see Greenwood, 1992). The last five to

seven (rarely four or eight) pored scales in that series are

separated from the dorsal fin base by only one large and one

much smaller scale. In the other serranochromine genera

only the last one or two pored scales are separated from the

fin base in this way, the other posterior scales having at least

two large scales of equal size interposed between them and

the fin base (see Greenwood, 1979 & 1992).

In most P. acuticeps specimens (from all localities) the

lower pharyngeal bone is slightly enlarged, and some of its

median row teeth, which are always coarser than their lateral

congeners, have molariform or submolariform crowns

(Greenwood, 1992, fig. 7). There is, however, considerable

individual variability in the degree to which this bone is

enlarged and its teeth are molarized. In these respects, P.

acuticeps resembles certain Sargochromis species, particularly

S. coulteri (Bell-Cross) and S. greenwoodi (Bell-Cross). But,

since the two genera differ in several other features, I would

consider this resemblance to be homoplastic and not, as

Trewavas (1961: 9) suggested, one of phylogenetic signifi-

cance.

If the serranochromines are a monophyletic lineage, (see

p. 41) their recent common ancestor could well have resem-

bled the extant Pharyngochromis acuticeps, except for the

incipient hypertrophy of the pharyngeal jaws in the latter.

Pharyngochromis acuticeps has a very wide distribution

which includes the Zambezi and Save-Runde river systems,

the Okavango river and its delta swamps, Lake Calundo, the

Lucala river (Quanza drainage) and some unidentifiable

localities in Angola. Records of the species (as Haplochromis

darlingi) from the Limpopo river system (Jubb, 1967,

repeated in Greenwood, 1979) are now known to be errone-

ous and probably based on the misidentification of small

Chetia flaviventris specimens.

A revised generic diagnosis of Pharyngochromis is given in

Greenwood (1992:48) and on p. 42 below.

CONCLUSION

The phyletic relationships of the

serranochromines

The difficulties encountered in determining both the inter-

and intrarelationships of these fishes were discussed in two

previous papers (Greenwood, 1979 & 1992), as was Trewa-

vas’ earlier (1964) tentative scheme of their relationships.

Basically the problem lay, and still lies, in establishing

whether or not the group is of monophyletic origin.

One of the two features unifying the serranochromines, the

non-ocellate anal fin markings, is probably a plesiomorphic

character, perhaps representative of a stage in the evolution

of true ocelli and one in which the markings, unlike true

ocelli, are not confined to males (Greenwood, 1979; Oliver,

P.H. GREENWOOD

1984). However, the possible function of anal maculae as

egg-dummies (sensu Wickler, 1962; 1963), or even if they

play any part in reproductive behaviour (Hert, 1989), have

yet to be determined. There is also the possibility that

non-ocellar anal markings have evolved more than once

within the haplochromine cichlids (sensu lato), as their pres-

ence in species of Thoracochromis (Greenwood, 1979, 1984)

could suggest. (Alternatively, Thoracochromis and the serra-

nochromines may be more closely related than other morpho-

logical and anatomical evidence would indicate.) In this

context, Eccles & Trewavas’, (1989: 27) obervations on three

species of the endemic Malawi genus Aulonocara are petti-

nent. One of these species has simple spots, a second has

spots surrounded by a contrasting border, and the third no

markings at all, although the fin has a pale border. Since

Aulonocara has a number of distinctive anatomical autapo-

morphies, this situation certainly suggests that not only have

different kinds of anal fin markings evolved more than once

but have done so within a single genus.

Another aspect of the problem involving phylogeny and

anal markings is Oliver’s (1984: 108) suggestion that hap-

lochromines with multiple non-ocellar spots, together with

those having true ocelli, comprise a monophyletic assemblage

whose members are more closely related to one another than

to any species with what he considers to be the pleisiomorphic

condition of anal maculae, namely spots indistinguishable

from those on the other unpaired fins, especially the dorsal

fin. That hypothesis has yet to be tested by the identification

of suitable congruent and derived features characterizing the

two supposed lineages, and raises questions about the signifi-

cance of the seemingly unique anal marking of Pseudo-

crenilabrus species (Greenwood, 1989).

All in all, the current evidence for anal fin markings being

of value in reconstructing phylogenies is not encouraging,

particularly at the taxonomic levels under consideration here.

For that reason I cannot agree with Eccles & Trewavas’

(1989) use of the feature as grounds for suggesting a close

relationship between the fluviatile serranochromines and

most of the endemic haplochromines of Lake Malawi. While

agreeing that non-ocellar anal spots are plesiomorphic fea-

tures, these authors believe that the absence of true ocelli in

the two groups “. . .combined with the geomorphological

history of the region . . . may be accepted as evidence for the

relationship of the Malawian group of species with the

haplochromines of the Zambezi area” (i.e., with Serrano-

chromis, Sargochromis and Pharyngochromis; Chetia has not

been recorded from the Zambesi system). Certainly the

similarity in anal fin markings would seem to support the

intuitive feeling that the two groups could be related (see

p. 34), and thus encourage a search for other characters to

confirm or refute that impression, but in itself I would not

rate it as ‘evidence’.

The second morphological character used to define the

serranochromines, i.e. cycloid scales above the lateral line

and a preponderance of such scales below that level, is, I

would argue, a derived condition (Greenwood, 1979; see also

Oliver, 1984; Lippitsch, 1991). Nevertheless, it is not clearly

an autapomorphic feature of the serranochromines. For

example, in the Lake Malawi haplochromines mentioned

above, there is also a marked reduction in scale ctenoidy, but

here, although scales above the lateral-line, like those in the

serranochromines, are cycloid, the ctenoid scales occurring

below that level are confined to the posterior part of the body

and not the anterior part as in serranochromines. In a

THE SERRANOCHROMINE GENERA REVIEWED

phylogenetic context how are these similarities and differ-

ences, to be evaluated?

Again, a pattern of reduced ctenoidy like that in the

serranochromines occurs in some but not all species of

Thoracochromis (Greenwood, 1979; 291; 1984:192 & 200), a

genus in which, apparently, there are both true ocellar and

non-ocellate types of anal fin markings (see above), but with

both kinds occurring only in males. Possibly the ‘genus’

Thoracochromis is polyphyletic and that some of its species

should be included in the serranochromine assemblage.

Here, as is so often the case, one is hampered both by a

paucity of detailed information on live coloration and the

relatively few specimens available for anatomical and mor-

phological studies.

Another difficulty, linked with lack of information, lies in

the possibility that further research could establish that there

really is a close phylogenetic relationship between the serra-

nochromines and certain haplochromines of Lake Malawi. In

that eventuality, it is possible that the nearest relative of one

or more of the fluviatile serranochromine taxa is to be found

in the lake’s fauna, thus rendering the serranochromines, as

currently conceived, either a para- or a polyphyletic group.

Although at present the monophyly of the serranochrom-

ines cannot be established or refuted, it is possible to con-

struct, on the basis of shared derived features, a tentative

intragroup taxonomy.

As compared with Pharyngochromis, the genera Chetia,

Serranochromis and Sargochromis all share two derived fea-

tures (see Liem, 1991) associated with the upper jaw skele-

ton, viz an increase in the shank length of the maxilla relative

to its pre-shank length, and an increase in the length of the

alveolar process of the premaxilla relative to the length of the

entire ascending process of that bone (see Methods). In

_Pharyngochromis the preshank portion of the maxilla is from

1.2-1.3 times longer than its shank length, whereas in Chetia

and Sargochromis the two parts are of equal length (with, in

some Sargochromis species the preshank portion slightly

shorter) and in Serranochromis the shank is noticeably longer

(as much as 1.3 times so). In Pharyngochromis the length of

the alveolar process of the premaxilla’s ascending process is

60-66% of the length of the entire ascending process; in

Sargochromis it is from 69-76%, in Chetia 73-77% and in

Serranochromis 73-83%.

Neither of these ratios, either inter- or intragenerically,

‘appears to be influenced by the size of the 17 specimens

examined, all in the size range 79-410 mm S.L. and repre-

senting ten species.

| On the basis of those two characters, a Pharyngochromis

}and a Chetia — Serranochromis — Sargochromis subgroup can

‘be recognised within the serranochromines. Both these

‘derived features are the only ones shared by the three latter

/genera (the increased number of abdominal vertebrae used

/previously to unite Sargochromis and Serranochromis in a

‘single genus [Greenwood, 1979] is now thought to be a

‘homoplasy; see p. 37).

Chetia and Serranochromis both share three derived fea-

\tures not found in Sargochromis or in Pharyngochromis viz.

(i) an increased modal number of caudal vertebrae (range

15-17, modes 16-17 in Chetia, and 15-18, modes 16 and 17 in

Serranochromis) compared with Sargochromis (range 12-16,

modes 14 and 15) and Pharyngochromis (range 14-16, mode

15). (ii) Two or more vertical rows of postorbital scales (cf a

\single row in Sargochromis and Pharyngochromis, see p. 36).

(iii) An increased number of scale rows around the caudal

peduncle, i.e. 18-20, rarely 16, compared with 16-18 rarely

15, (mode 16), in Sargochromis, and 15 or 16 (mode 15) in

Pharyngochromis.

On the basis of those three derived characters, and using

the term ‘sister taxon’ without any phylogenetic implications,

then, within the serranochromines, Pharyngochromis is the

sister taxon to the group Sargochromis, Chetia and Serrano-

chromis, and within that latter group, Sargochromis is the

sister taxon of Serranochromis and Chetia combined.

That scheme bears, in broad outline, a close resemblance

to Trewavas’s (1964) diagram suggesting the interrelation-

ships of Serranochromis, a scheme based essentially on

lateral—line and dorsal fin ray counts (which may, of course,

be correlated, in part, with the vertebral counts used here)

and certain characteristics of the pharyngeal jaws.

When comparing the two schemes, allowances must be

made for the fact that two of Trewavas’ Haplochromis species

(lucullae and darlingi) are now treated as synonyms of

Pharyngochromis acuticeps (Greenwood, 1992), that the sta-

tus of H. angolensis, H. humilis and H. toddi is uncertain or

unknown (Bell-Cross, 1975, Greenwood, 1979), and that H.

welwitschi is now considered to be a species of Chetia (see

p. 39). Also, the two Haplochromis species, mellandi and

frederici, placed in limbo between the genera Haplochromis

and Sargochromis in Trewavas’ diagram, are now included in

Sargochromis (Bell-Cross, 1975; Greenwood, 1979 and above

p. 37), as is Haplochromis carlottae.

Trewavas (op. cit. 9-10) superimposed on her diagram a tree

indicating the possible phylogenetic relationships of the taxa,

both inter- and intragenerically. It is here that our views would

not coincide, mainly because I do not think there is the evidence,

based on cladistic methodology, to justify the relationships

proposed, even at an intergeneric level (see p. 40). Certainly

there are no features justifying Trewavas’ (op. cit. 10) suggestion

that Serranochromis is a diphyletic and gradal taxon or that a

cladal grouping would recognise Chetia, Serranochromis robus-

tus and S. thumbergi on the one hand, and Chetia welwitschi

(Trewavas’ Haplochromis welwitschi), and the remaining Serra-

nochromis species on the other. Nor can I accept Trewavas’

uncertainty about the separation of Sargochromis codringtoni,

mellandi, carlottae and greenwoodi (as Haplochromis frederici in

her scheme; see Bell-Cross, 1975) from the ‘Haplochromis’ (i.e.

Pharyngochromis,) root of her tree (see Greenwood, 1979, 1992

and above). A truly phylogenetic assessment of the groups’

relationships must, as discussed on p. 36, await the results from

further and preferably multidisciplinary research into the system-

atics of all the serranochromine species, and those many Malawi

species that Eccles & Trewavas (1989: 21) have placed in their

Pharyngochromis — Chetia— Serranochromis group.

Generic key and diagnoses

A single row of scales between the posterior orbital margin

and the vertical limb of the preoperculum ................... A

Two or more scale rows between the posterior orbital margin

and the vertical limb of the preoperculum .................... B

A. Last 2 or 3 pored scales in the upper lateral-line series

separated from the dorsal fin base by not less than two scales

of approximately equal size ................ A(i) Sargochromis

Last 5 to 7 (rarely 4 or 8) pored scales in the upper lateral line

separated from the dorsal fin base by one large and one small

SCAGEEP,, F- bedy. Se Oe Pent ata: Sao A(ii) Pharyngochromis

A(i):

Abdominal vertebrae [15] 16-18 [19], modes 16 and 17;

caudal vertebrae 12-16, modes 14 and 15; total number of

vertebrae 28-32 (mode 31).

Dorsal fin with 13-16, modes 15 and 16, spinous rays and

11-16, modes 12 and 13, branched rays. Anal fin with 3 spines

and 8-11, mode 9, branched rays. Caudal fin truncate,

subtruncate or almost rounded.

Scales in the lateral series 28-34, modes 30 and 31. Cheek

with 3-6 horizontal rows. [15] 16-18, mode 16, scales around

the caudal peduncle.

Gill-rakers in the outer series on the first ceratobranchial

9-15, modes 12 and 13.

Outer series of teeth in both jaws composed mainly of

unequally bicuspid teeth in fishes <150 mm S.L., but pre-

dominantly of unicuspids in larger specimens. Inner series of

jaw teeth, except in one species, arranged in a single or

double series anteriorly and anterolaterally, reducing to a

single row posterolaterally; in the exceptional species there

are 4 rows anteriorly and anterolaterally, and a single row

posterolaterally.

Pre-shank length of the maxilla equal to, or slightly shorter

than the shank-length (see p. 34). Height of the premaxillary

alveolar process 69-76% of the height of the entire ascending

process (see p. 35). For comments on neurocranial morphol-

ogy and other osteological features (including the lower

pharyngeal bone and its dentition, see text and figures in

Bell-Cross (1975) and Greenwood (1979: 303-305, figs.

16-18; and 1984: 216-225, figs. 12-17).

Lower pharyngeal bone hypertrophied in the majority of

species, greatly so in some, but only slightly enlarged in two

species. The extent and degree to which the dentition of this

bone is molarized is positively correlated with the degree of

the bone’s hypertrophy. In species with slightly enlarged

bones only a few molar-like or submolariform teeth are

present, and are confined to the median tooth rows. The

ventral outline of the bone’s anterior keel is almost straight

and rarely extends below a horizontal drawn through the

deepest point on the bone’s ventral surface below the denti-

gerous area. In specimens with a greatly enlarged lower

pharyngeal bone, however, the ventral margin of the keel

extends a little below that level (see figs 12-17 in Greenwood,

1984).

Anal fin spots small and numerous (as many as 40). ..........

TE TASES. UD. LSI Sees EM SS ed Sargochromis

A(ii):

Abdominal vertebrae [13] 14 or 15 [16], mode 14; caudal

vertebrae 14-16, mode 15; total number of vertebrae 28-31,

mode 30.

Dorsal fin with 14-16, mode 15, spinous rays and [10]

11-13 [14] branched rays. Anal fin with 3 spines and 8 or 9

branched rays (no distinct modal number). Caudal fin dis-

tinctly truncate, subtruncate or almost rounded.

Scales in the lateral series [30] 31-36, mode 33, modal

range 32-34. Cheek with [3] 4-6 horizontal rows. 15, rarely

16, scales around the caudal peduncle.

Gill-rakers in the outer row on the first ceratobranchial

7-12, modes 9 and 10.

Outer series of jaw teeth composed of unequally bicuspid

teeth in fishes <80 mm S.L., although some unicuspids can

be found in larger fishes within that length range. Unicuspids

become the predominant form in fishes >90 mm S.L. Inner

series of teeth, in both jaws, arranged in 1-3 rows anteriorly

P.H. GREENWOOD

and anterolaterally, reducing to a single row posterolaterally.

The number of inner rows anteriorly appears to be positively

correlated with an individual’s size.

Pre-shank length of the maxilla clearly greater than the

shank length (see p. 34), i.e. about 1.2-1.3 times longer.

Height of premaxillary alveolar process 60-66% of the height

of the entire ascending process. For comments on neurocra-

nial form and other osteological features, see Greenwood

(1992).

Lower pharyngeal bone in most individuals showing a

slight degree of hypertrophy. In specimens over 50 mm S.L.,

the median rows of lower pharyngeal teeth are composed of

noticeably coarser teeth than those situated laterally, and

some can have submolariform crowns; the degree of molar-

ization is most marked in fishes over 100 mm S.L. Irrespec-

tive of the degree to which the lower pharyngeal bone is

enlarged, its anterior keel is deep, with a curved ventral

outline whose deepest point lies below a horizontal drawn

though the deepest point of the ventral surface underlying the

dentigerous part of the bone (cf Sargochromis above); see fig.

7, Greenwood, 1992.

Anal fin spots of variable size and number, from as few as 3

or 4 large spots to as many as 19 small ones ....................

Sue RTC 2a. Jas Lee SRE Pharyngochromis.

B. 16-18 (rarely 15 or 19) abdominal vertebrae; inner and

outer rows of jaw teeth composed entirely or mostly of

unicuspids in fishes over 30mm S.L. (and posibly in smaller

incividtialstasiwell)"", Sess. secsccnseseree cee B(i) Serranochromis

14 or 15 abdominal vertebrae; many bicuspid (or weakly

bicuspid) teeth present in the outer tooth rows of both jaws in

HISHES ASMAnPe AS GUM Seles certs nee see eee eee B(ii) Chetia

B(i)

Abdominal vertebrae [15] 16-18 [19], modes 16 and 17;

caudal vertebrae [15] 16-18, modes 16 and 17; total number

of vertebrae 31—36 (no distinct modes).

Dorsal fin with 13-18, modes 15 and 16, spinous rays, and

13-16, modes 14,15 and 16, branched rays. Anal fin with 3

spines and 9-13, modes 10 and 11, branched rays. Caudal fin

subtruncate or almost rounded.

Scales in the lateral series [34] 35-41, no distinct modes.

Cheek with 3 (rarely) to 11 horizontal rows (modally 5-9

rows). 18-20 scales around the caudal peduncle (no distinct

mode).

Gill-rakers in the outer series on the first ceratobranchial

[8] 9-13, modes 10,11 and 12.

Outer and inner series of jaw teeth composed of unicuspids

in specimens over 30 mm S.L. Inner series of both jaws, in all

but one species, arranged in a single or double row (rarely 3

rows) anteriorly and anterolaterally, and a single row poster-

olaterally. In the exceptional species there are as many as six

rows anteriorly and anterolaterally, reducing to a single or

double row posterolaterally.

Pre-shank length of the maxilla shorter than its shank-

length (see p. 34), which is ca 1.2-1.3 times longer than the

pre-shank portion. Height of the premaxillary alveolar pro-

cess 73-82% of the height of the entire ascending process (see

p. 34). For comments on the neurocranium and other osteo-

logical features see Greenwood (1979: 299-302; figs. 13-15)

and Trewavas (1964).

Lower pharyngeal bone slender, its dentigerous surface

elongate and narrow (see figures in Trewavas, 1964, and

Greenwood, 1979). No molariform pharyngeal teeth; even

THE SERRANOCHROMINE GENERA REVIEWED

those teeth in the median rows are only a little coarser than

the other and fine teeth on the bone.

Anal fin spots small and numerous (as many as 40)

Rett wits Precast oe bteeeaeuiadecieinnbsGnieeese naa Serranochromis

B(ii):

Abdominal vertebrae 14 or 15 (no distinct mode); caudal

vertebrae 15-17, modes 16 and 17; total number of vertebrae

30-32, mode 31.

Dorsal fin with 14 or 15, mode 15, spinous rays and 10-13,

modal range 11 or 12, branched rays. Anal fin with 3 spines

and 7-10 branched rays (no distinct mode). Caudal fin

truncate to subtruncate.

Scales in the lateral series 32-35, modal range 32-34.

Cheek with 4-6, modes 5 and 6, horizontal rows; 18 or 20

(rarely 16) scales around the caudal peduncle.

Gill rakers in the outer series on the first ceratobranchial

9-11, modes 10 and 11.

Outer series of teeth in both jaws composed mainly of

unequally bicuspids, but with a few unicuspids or weakly

shouldered bicuspids present, in fishes < 80 mm S.L.; how-

ever, in some specimens of Chetia flaviventris, unicuspids

predominate in fishes in the upper part of that size-range. In

specimens >100 mm S.L. the outer teeth are predominantly

unicuspid, with a few very weakly shouldered bicuspids also

present. Inner tooth rows of both jaws arranged in a double

series anteriorly and anterolaterally, reducing to a single row

laterally and posteriorly. In one species (Chetia gracilis) at

least some specimens have the two anterior median teeth in

the outer row of inner teeth enlarged and displaced anteriorly

relative to the other teeth in that row.

Pre-shank length of the maxilla equal to its shank length

(see p. 34). Height of the premaxillary alveolar process

73-77% of the height of the entire ascending process (see

p. 35).

Except in one species, the lower pharyngeal bone is not

enlarged, and the median tooth rows are composed of

bicuspid teeth only a little coarser than their lateral conge-

ners. In the exceptional species, C. mola (see Balon &

Stewart, 1983; fig. 12) the bone is greatly hypertrophied and

massive, with all but a few of its laterally situated teeth

enlarged and molariform or submolariform.

Anal fin spots usually small and fairly numerous (7-15) but in

one species, C. brevis, there are only 3 or 4 large spots.

Erde dew Maducottatrerenicoceeterceecs swcducoes seitecs Chetia

_ ACKNOWLEDGEMENTS. To Paul Skelton go my sincere thanks for the

_ many discussions we have had on the subject of serranochromine

_ cichlids, and for providing me with excellently documented material

jand field data. Once again it is a pleasure to thank Mike Bruton,

| Director of the J.L.B. Smith Institute for his hospitality and the fine

working facilities provided by the Institute, Huibre Tomlinson for her

| tenacity and good humour when producing the typescript, and Elaine

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South African Museum 46(5): 53-56.

—— 1964. A revision of the genus Serranochromis Regan (Pisces, Cichlidae).

Musee Royal de l'Afrique Centrale, Tervuren; Annales, serie in —8°, Sciences

Zoologique no. 125: 1-S8.

— 1983. Tilapiine fishes of the genera Sarotherodon, Oreochromis and

Danakilia viii + 583p. British Museum (Natural History), London.

Wickler, W. 1962. Ei-Attrappen und Maulbriiten bei Afrikanischen Cichliden.

Zeitschrift fiir Tierpsychologie 19(2): 129-164.

P.H. GREENWOOD

—— 1963. Zur Klassifikation de Cichlidae, am Beispiel der Gattungen

Tropheus, Petrochromis, Haplochromis und Hemihaplochromis n.gen.

(Pisces, Perciformes). Senckenbergiana Biologica 44(2): 83-96.

Winemuller, K.O. & Kelso-Winemiller, L.C. 1991. Serranochromis altus, a new

species of piscivorous cichlid (Teleostei:Perciformes) from the upper Zam-

bezi river. Copeia 3: 675-686.

Bull. nat. Hist. Mus. (Zool.) 59(1): 45-81 Issued 24 June 1993

A revision of Danielssenia Boeck and Psammis

Sars with the establishment of two new genera

Archisenia and Bathypsammis (Harpacticoida:

Paranannopidae)

RONY HUYS*

Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD

J. MICHAEL GEE

Plymouth Marine Laboratory, Prospect Place, The Hoe, Plymouth PLI 3DH

CONTENTS

LPOG NGL ect ocar op siciae Poon ont BEC OC CCCP CSTE IGO 2 00SEC TACGTt CROMER nce aici. SAA nea en art or ene anne ir at See nner ae 46

NEAL eTEAISHALC MICOS) a.com -etasee ese cenme ane tac taccctce cess xox ote teenm daaad ons ntaclte Dbestes cee ae abeutad dulssseSb uted. dod. dette 46

SU RECTATATICS HMA, SPER NCEE Eee tattoo tat ve ect res naea Me Nes «nn nn QUOP ECR MRMEEI RII Gueg. 12> mS Wlaliass EER oe letaes od cath woes 47

HALLE Ara ANNO PIGACTEOL MOG tater abatdys te gC e se « sc ncn ablch arab aati wanes: dh ane din debh exe sdcancasstckhe doads. 4. satteebh 47

MGeNTS: ALG /PISCHIIG PE UE NOM ws Itt. EMEI One u non acted Uat c's nt naaPenmetnUeh oMlange -h Sk ad dasa Nes «dciehindd ache ve -adeceucaveeee Pe 47

PAT CHISENIG: SIDEICH: (SAIS M1S9S) COM. NOVA Nees ss ... «deat tes pee asa <eetdo snes, cbuued shah beeketelee »tiedsee vee. dents -cuest 47

ROTIAEIG SRcReee 28. SRR ne ree math cda Adee te an Ieee teeth ene vans atett ates ee qeeey ee Tekaoans dasat sah sabia dvalucaacesSsadasns tess oaates 51

(Oy RO MMORVINY: J. .c55 . Grutaads. . SR ONEET, WUE , «cae setae oemv egy Shs Cauts sdtyreetes ty ees oneebtane «sso vach aueslidszeavagye 51

(iE ALILAPOMLOLDINES Mchomnere. tere h. hes Le EdRG TS «a0 ve cstler shone dnactsicdduaaermteadre® ot dninde uv doslecinde date ctilnceoe ave tcee 60

(ED) LEME T SO RURAL) coc cocctinn ous arden saree cnn cn can conan de tneh anda pins Giga un Same eed cde cena ee chests suade op emaeee 60

MVE TRIS LO AITICLSSCNEEE EXO ORS LUA Catccicoatrorccrougise genera siceic else ace one eT Ota aa nisleix Minis basaroscaiaAie 8 Gainiaicnis ath go;e.n enn cinte'o.oldieninacnsaidiss 62

(i) Danielssentajusijormis. (Brady,..1880) nec Sars (1910) « v.nc tins .dicescesctmnoundscccsdessnesassace convacunes 62

(Gu) BR AIIE ISS ITIL TODSIA SATS lO AUD a. den Res » tos « dcleabics Padaidocn cevuinss edeioidvomascbateside Goenteshirsoscegeenkis date 62

ELD LD AIILE LS SEPIEANSEITIEIES’ @ MISE KOS LO cect .ciicuespunteneot dees dechaens.s cangeecba yoaldses davis vee. ddckeddamucsseee 64

(iv) Darnrelssenrannvoted BOCK VS72 veces cies en's «ggutee satrampauteenny daswtcnaeed s+ divcks auneoes ruses tesauccessesses 64

QUJiee eC CRA PHIOSIG Metron. tentaeh fan se nce en cin vacvarns cacricartranecas dar saccqcagasc¥enpsaecuacecade snags oMeatinas 67

GETSTEAIILY ISAIITUST COLA MOVE scodrcteusor cori teeta as sacle neceve dente te neonea toes -Scsied sadnenescavenetioe ciecusagess suaetees 67

Bally PSAMIMIS'LON HUNCH (BOUIN, 1968) COMM DOV. .catuserues dances casts: aecserczecsccnescvets dec cadudccsctercsecseees 69

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(CaS BsanintisvDOreaisik lie M1959 ere. AO... SR a a. Sa IS 72

CMON saris: Riven SrsienOVynl 946 woe O ES «.« «aterm deere rete ed hanes Teese ches abote te cet slikatok baad aeoumeaee 72

O03) ARP SGIITISIIOTIGIDESHECCKETHIO TALOIT IRS. Bw cn << AINE eorex dete aaah aces tee sO ELA foc oooh eanalt 72

(LV) ae LE SOPPIYTLS LOPIGISCLOSCL SALSA SLO) oceania cata cotteepielasicicatsiqepetausa’ sess samnedae.adesinanaisatpmaeseepam opines «sin 72.

CY) ieee Amended dia ei SIS srreseeeseecaee rae eat aca sats dana sans sania tala sae sinnas cn eaniiasdaineedasienmanisinaiat snaiabinsle calsiineiin 75

MOSTSCUISSTONN, spehet as foactrcss sie vasa cioeirats sebteeaate sian aoa tne aice sole tcie cist sah pains vd wpisUPalds se nasiaiga' ciduenescsesinasumedsaleauniaonsudanessaamd 78

Maeva OrOenelal OlibardnannOplades eevcteccn tusk casvose se anas tosh evan aasGek asics no's stelepe Gay ceeculss Onis a)coiianaseeasae deacon seers 79

EXGRMOWICULEMICHIS MEIN. vonjecia scence cueatesae Gdn sincajeaeaviaveaeacndsderanccasnnqusagaettecetes etn tetey Cenc aitaatiense sees cina teeters 80

RO LENE COS iis toate sce cnren act aca cine ae etic auras snaewedeie tinea cas cu case sin maisciacacisine on caisltveiseaceinassitess cop siies nemeciiawan teers somes 80

Synopsis. Archisenia gen. nov. is proposed to accommodate the sibirica-group of the genus Danielssenia Boeck,

1872. Re-examination of Alaskan material of D. stefanssoni Willey, 1920 has shown the latter species to be a junior

synonym of D. sibirica Sars, 1898, the type and only species of the new genus.

Danielssenia robusta Sars, 1921 and Fladenia intermedia (Wells, 1965) are synonymous and consequently F.

robusta comb. nov. becomes the type species of the genus Fladenia Gee & Huys, 1990. Danielssenia similis

Chislenko, 1971 is regarded as species inquirenda and the genus Danielssenia is redefined from the type species D.

typica Boeck, 1872, and two other species (D. quadriseta Gee, 1988 and D. reducta Gee, 1988).

The status of D. fusiformis (Brady, 1880) nec Sars (1910) is reconsidered and as a result the genus Sentirenia Huys

& Gee, 1992 is relegated to a junior synonym of Jonesiella Brady, 1880 which is reinstated to accommodate J.

fusiformis Brady, 1880 and J. eastwardae (Coull, 1971) comb. nov.

Psammis borealis Klie, 1939 is removed from the genus Psammis Sars, 1910 but retained in the Paranannopidae as

species incertae sedis. P. longifurca Bodin, 1968 is transferred from Psammis to Bathypsammis gen. nov. The genus

Psammis is redefined on the basis of the type species P. longisetosa Sars, 1910, and P. longipes Becker, 1974.

* Visiting Research Fellow of the Institute of Zoology, University of Gent, B-9000 Gent, Belgium.

R. HUYS AND J.M. GEE

A detailed redescription of A. sibirica and new illustrations of D. typica, P. longisetosa, P. longipes and B.

longifurca are provided.

Intersexuality in copepods and the possible phylogenetic relationships of Danielssenia, Psammis, Fladenia,

Archisenia gen. nov. and Bathypsammis gen. nov. are briefly discussed.

A key to the genera of the Paranannopidae is presented.

INTRODUCTION

Throughout its taxonomic history up to the late 1980s, the

genus Danielssenia Boeck, 1872 has served as a repository to

accommodate different kinds of ‘tachidiid’ harpacticoid cope-

pods, in so far that the distinction between this genus and

Psammis Sars, 1910 almost became no longer tenable (Wells,

1965, 1967). Gee (1988a) pointed out that differences in

mandibular gnathobase structure, possibly reflecting different

diets, could indicate that both genera are trophically isolated,

but admitted that perhaps more solid morphological evidence

is necessary to maintain generic distinction.

The criteria applied by most workers to allocate newly

discovered species to Danielssenia generally had no phyloge-

netic significance as they were mainly based on plesiomorphic

character states (i.e. P1 not modified) which are diagnostic of

a wider group of families. Virtually no effort has been made

to correctly assess the sexual dimorphism on the swimming

legs and very little information on detailed mouthpart struc-

ture has been documented. Both categories of characters

have nevertheless proved to hold a high phylogenetic infor-

mation content that can be used to determine relationships

within the Danielssenia-Psammis core group of genera (Gee

& Huys, 1990, 1991; Huys & Gee, 1992, in press).

The impact of Lang’s (1944, 1948) classification of the

Tachidiidae also caused people to lose sight of the relation-

ships of this core group with taxa beyond the family bound-

aries. The fact that his artificial subdivision into three

subfamilies constrained the development of alternative phy-

logenetic scenarios for a long time is illustrated by the

ongoing discovery and description of numerous new species

of Paranannopus Lang, 1936 (placed in the Cletodidae and

subsequently in the Paranannopidae) and Danielssenia

(placed in the Thompsonulinae, Tachidiidae) in the post-

Langian era without any recognition of the close relationship

between these two taxa. Huys & Gee (1990) inevitably had to

break down the concept of the Thompsonulinae before they

could re-allocate the ’danielsseniid genera’ to the Paranan-

nopidae. This group of genera essentially represents the

continental shelf lineage of the family with a few species that

secondarily explored deeper habitats (e.g. Leptotachidia iber-

ica Becker, 1974). Its affinity to the predominantly deepwater

group, containing Paranannopus and Cylindronannopus

Coull, 1973, has recently been supported by the redescription

of Fladenia Gee & Huys, 1990, a possible ’missing link’

between both lineages (Gee & Huys, 1990).

This paper is the final contribution to a revision of the

genus Danielssenia, including the allocation of the sibirica-

group to a new genus Archisenia, thus reducing the number

of species previously referred to the genus from 14 to four

(Table 1). It also presents a revision of the other major genus

Psammis, resulting in the proposal of a new genus Bathyp-

sammis. With the revision of these taxa the establishment of

novel genera draws to a close and, accordingly, a key to

genera of the Paranannopidae is presented.

MATERIALS AND METHODS

Before dissection, the habitus was drawn and body length

measurements were made from whole specimens temporarily

mounted in lactophenol. Specimens were then dissected in

lactic acid, the parts mounted in lactophenol and the prepara-

tions sealed with glyceel® (BDH Chemicals Ltd, Poole,

England). All drawings of the specimens were prepared using

a camera lucida on a Leitz Dialux 20 or Leitz Diaplan

Table 1 Re-allocation of species previously referred to Danielssenia Boeck, 1872.

Species previously referred Current status

to Danielssenia

typica Boeck, 1872

fusiformis sensu (Sars, 1910)

quadriseta Gee, 1988

reducta Gee, 1988

similis Chislenko, 1978

sibirica Sars, 1898

stefanssoni Willey, 1920

fusiformis Brady, 1880

perezi Monard, 1935

paraperezi Soyer, 1970

eastwardae Coull, 1971

robusta Sars, 1921

intermedia Wells, 1965

spinipes Wells, 1967

minuta Coull, 1969

Danielssenia typica Boeck, 1872

Danielssenia quadriseta Gee, 1988

Danielssenia reducta Gee, 1988

Danielssenia similis Chislenko, 1978 [sp. inq.]

Archisenia sibirica (Sars, 1898) comb. nov.

Jonesiella fusiformis Brady, 1880

Jonesiella eastwardae (Coull, 1971) comb. nov.

Fladenia robusta (Sars, 1921) comb. nov.

Afrosenia spinipes (Wells, 1967)

Sentiropsis minuta (Coull, 1969)

Reference

Gee (1988)

Gee (1988), present account

Gee (1988)

present account

Huys & Gee (1992), present account

present account

Gee & Huys (1988), present account

Huys & Gee (in press)

REVISION OF DANIELSSENIA AND PSAMMIS

differential interference contrast microscope. The terminol-

ogy for body and appendage morphology is according to

Huys and Boxshall (1991). Abbreviations used in the text and

figures are P1—P6 for thoracopods 1-6; exp(enp)-1 (-2,-3) to

denote the proximal (middle, distal) segment of a ramus.

Body length was measured from the base of the rostrum to

the posterior margin of the anal somite.

SYSTEMATICS

Family Paranannopidae Por, 1984

Genus Archisenia gen. nov.

SYNONYMY. Danielssenia Boeck, 1872 (part).

DIAGNOSIS. Paranannopidae. Body large, slightly fusiform

and dorso-ventrally flattened. Rostrum not hyaline, with 2

pairs of small sensillae. Somatic hyaline frills minutely den-

tate. Female genital double-somite with lateral and ventral

sub-cuticular ridge marking original segmentation; genital

field with minute copulatory pore and sinusoidal copulatory

duct leading to transverse seminal receptacle partly located

anterior to genital slit; P6 with 1 outer plumose seta and 2

minute spiniform elements. Pseudoperculum hyaline with

dentate margin. Caudal rami slightly divergent and slightly

longer than broad. Female antennule 6-segmented; aes-

thetasc on segment 4; distal 2 segments with heavily pectinate

spines. Antennary exopod 3-segmented with armature for-

mula [2-1-3]. Mandibular coxa elongate, with blunt teeth on

gnathobase; basis with 4 setae; endopod 1-segmented; exo-

pod 2-segmented. Maxilliped subchelate with 1 large and 1

small seta on syncoxa; basis with naked seta on palmar

margin, endopodal claw with 2 accessory setae. Pl exopod

3-segmented, exp-3 with distal outer spine longer than middle

outer spine; endopod longer than exopod, 2-segmented,

enp-2 4.5 times longer than broad, inner seta implanted

medially. P2—P4 intercoxal sclerites with spinules or setules

on distal margin; rami 3-segmented; exp-1 with inner seta;

female P2-P3 enp-2 with small apophysis at outer distal

corner. Armature formula of P1—P4 as follows:

Exopod Endopod

al 0.1.023 1.121

P2 121.223 fo 221

P3 e323 EIS Al

P4 Ha 328) 1.1.221

Female fifth pair of legs not fused medially; exopod and

baseoendopod separate, each with 5 setae, inner seta on

exopod well separated from remaining 4 setae.

Male with sexual dimorphism on antennule, P1, P2 endo-

pod, P3 endopod, P5, P6 and in genital segmentation.

Antennule 9-segmented, subchirocer; segment 6 very swol-

len, with aesthetasc. P1 inner basal spine less strongly devel-

oped, segments of rami more slender and spinule rows on

outer margin of endopod much smaller. P2 enp-1 larger, with

inner seta transformed into a non-articulating process; enp-2

without inner seta, outer distal corner attenuated into a long

apophysis reaching far beyond the distal border of enp-3;

nn —e—EEeEeEeEeEeEeEeEeEeEeEeEeEeEeEeEeEeEeEee—eEeeEeEeEeEeeew

enp-3 with distal outer spine partially fused to segment, much

shorter and stronger than in female, with spinules reduced to

coarse teeth, other setae reduced in size. P3 enp-2 with inner

distal corner slightly attenuated, outer distal corner attenu-

ated into a hook-shaped apophysis. Fifth pair of legs fused

medially; baseoendopod and exopod separate with 2 and 5

setae, respectively. P6 symmetrical, fused to somite, with 3

setae each.

TYPE SPECIES. As a result of the arguments and analysis put

forward below we regard D. stefanssoni Willey, 1920 as a

junior synonym of D. sibirica Sars, 1898 and therefore A.

sibirica (Sars, 1898) comb. nov. is designated as the type

species.

OTHER SPECIES. None.

ETYMOLOGY. The generic name is derived from the Greek

prefix archi, meaning first in time and alludes to the primitive

position in the family. Gender: feminine.

Archisenia sibirica (Sars, 1898) comb. nov.

SYNONYMY. Danielssenia sibirica Sars, 1898; Danielssenia

stefanssoni Willey, 1920.

MATERIAL EXAMINED.

— National Museum of Natural History (Smithsonian Institu-

tion), Washington, D.C.: 8 99 and 1 CO from Point Barrow,

Nuwuk Lake, Alaska, U.S.A.; collected by R. Lewis et al.,

August 1 1960, bottom sample A974; identified as D. stefans-

soni by M.S. Wilson; 1 ? dissected on 13 slides, 1 0" dissected

on 7 slides, others preserved in alcohol: reg. no. USNM

204769.

— Naturhistoriska Riksmuseet, Stockholm: 1 @ and 1 oC’

from East Greenland, Barclay Bay; collected by Jespersen,

July 14 1932; identified as D. stefanssoni by K. Lang;

preserved in alcohol; reg. no. Cop. 31.

DESCRIPTION OF FEMALE. Body slightly dorso-ventrally flat-

tened (as for male, Fig. 9B); length 0.97-1.242 mm (x =

1.15 mm; n = 7); urosome clearly demarcated from prosome.

Cephalothorax rounded anteriorly, widest near posterior

margin. Rostrum (Fig. 2A) not hyaline; tapering anteriorly;

with 2 pairs of sensillae. Free prosomites each with a dorsal

row of spinules and some sensillae near posterior margin;

hyaline frill of prosomites minutely dentate. All urosomites

(Fig. 1A-B) with lateral row of spinules; first to third

urosomites with dorsal row of spinules, 2 rows dorsally on

genital somite; ventral spinule row on posterior border of

genital double-somite and succeeding urosomites, slightly

anterior to lateral rows. Genital double-somite (Fig. 1A—B)

with lateral and ventral subcuticular ridge. Genital field (Fig.

1C—D) with minute copulatory pore posterior to genital slit;

copulatory duct sinusoidal (Fig. 1D) leading to single, trans-

versely elongate seminal receptacle located at level of genital

slit; vestigial P6 with 1 plumose seta and 2 spinules (vestigial

setae?); paired, blind ending, cuticular invaginations poste-

rior to genital field (Fig. 1C). Hyaline frill of urosomites

minutely dentate, that of penultimate somite extended to

form pseudoperculum (Figs. 1B, 8C). Anal somite deeply

incised. Caudal rami (Figs. 1E, 5F, 8C) tapering posteriorly,

slightly longer than broad, with short spinule row medially on

lateral margin and a latero-ventral spinule row on distal

margin which also has a large pore near ventral outer corner

(Fig. 1E); seta I minute (Fig. SF); setae IV & V well

48 R. HUYS AND J.M. GEE

= \y eae marie] won

w LA

—

—~= —

Ga peMeeiay [ i" jen

ra Ba

VLE ae LD ty Se

SLL IE tay EE

cil LAM NG 9 i

Doc yg

| 6

( anim f

ne wey

—

eee

Vy iy Le a

LIE

LLLLL.

———

ypu

Fig. 1 Archisenia sibirica comb. nov. A, Female urosome (excluding P5-bearing somite), ventral view; B, same, dorsal view; C, female

genital field, ventral view; D, same lateral view; E, caudal ramus, ventral view.

REVISION OF DANIELSSENIA AND PSAMMIS

developed, spinulose in distal portion (Fig. 1E); seta VII

tri-articulate (Fig. 8C).

Antennule (Fig. 2A—B) 6-segmented; segment 1 with 2

spinule rows on outer margin and a plumose seta at outer

distal corner. Segment 2 with 5 pinnate and 1 naked setae on

outer margin and 2 pinnate and 1 naked setae posteriorly

directed on dorsal margin. Segment 3 with 2 pinnate and 6

naked setae at outer distal corner. Segment 4 with 6 naked

setae and an aesthetasc. Segment 5 with 3 pectinate spines, 3

naked and 2 pinnate setae. Segment 6 with 1 pectinate spine

and 7 naked setae.

Antenna (Fig. 2C-D). Coxa with a row of spinules proxi-

mally. Allobasis with long spinules at base of abexopodal,

pinnate seta. Exopod 3-segmented with armature formula

[2-1-3]; distal segment elongate with subterminal row of

spinules. Endopod with 2 spinule rows on outer margin; 2

spines, a geniculate seta and a naked seta subdistally (Fig.

2C); distal margin with a pectinate spine, 4 geniculate setae, a

small plumose seta (Fig. 2C) and a very small naked seta (Fig.

2D).

Mandible (Fig. 3A—B). Coxa (Fig. 3B) elongate, slender,

with 2 median rows of spinules; gnathobase with bidentate

and unidentate teeth and a pinnate seta at inner distal corner.

Palp biramous. Basis (Fig. 3A) with patch of spinules medi-

ally and 4 pinnate setae on distal margin. Exopod

2-segmented; proximal segment with 2 pinnate setae laterally

and a row of large spinules distally; distal segment with 3

apical setae. Endopod 1-segmented with 3 lateral and 6 distal

setae.

Labrum (Fig. 3C) with numerous spinule rows near median

distal margin of posterior face.

Maxillule (Fig. 3D). Praecoxal arthrite with 2 juxtaposed

setae medially on anterior surface and 9 bidentate or pinnate

spines and 1 naked seta on distal margin. Coxal endite with 5

armature elements on distal margin. Basal endite with 2

subdistal setae and 4 setae on distal margin. Rami

1-segmented and each with 3 setae.

Maxilla (Fig. 4B). Syncoxa with spinule row at outer

proximal corner and with 3 endites each with 1 fused and 2

articulating pinnate spines. Allobasal endite with a fused

pinnate claw, a pinnate spine and 2 setae. Endopod

1-segmented with a pinnate spine and 3 setae.

Paragnaths (Fig. 4A) well developed; with several rows of

fine spinules laterally and medially; anterior face with coarse

teeth.

Maxilliped (Fig. 4C). Syncoxa with numerous spinule rows,

_ 1 large subterminal and 1 smaller terminal pinnate seta. Basis

| with row of spinules and a naked seta on palmar margin.

Endopodal claw as long as basis, spinulose distally and with 2

accessory setae proximally.

P1 (Fig. 5A). Intercoxal sclerite rectangular with 2 groups

of setules on each side. Coxa with rows of spinules on

anterior face and outer margin. Basis with row of spinules on

inner and distal margin and around base of inner pectinate

spine (Fig. 1D) and outer pinnate seta. Exopod 3-segmented,

each with row of spinules on outer margin, outer spines

pectinate, distal outer spine on exp-3 longer than middle

Outer spine. Endopod longer than exopod, 2-segmented;

proximal segment slightly longer than broad, distal segment

about 4.5 times longer than broad, inner seta implanted

medially.

P2-P4 (Figs. 6A, 7A, 8A). Intercoxal sclerite with row of

spinules or setules on each side. Both rami 3-segmented,

equal in length in P2 but with endopod shorter than exopod in

P3 and P4; all segments with rows of spinules on outer

margin; P2 and P3 with a large spinule at base of each inner

seta on enp-2 and -3. Exp-1 with inner seta; enp-2 with outer

distal margin somewhat attenuated. Armature formula of

swimming legs as in generic diagnosis.

Fifth pair of legs (Fig. 11D) not fused medially; exopod and

baseoendopod separate. Baseoendopod with short row of

spinules at base of exopod and setophore of outer seta;

endopodal lobe well developed, tapering distally, with 5

pinnate setae, second outer seta longest. Exopod wider than

long, boundary with baseoendopod straight, not reaching to

distal margin of endopodal lobe; with 5 pinnate setae, 4

grouped together on distal outer margin and 1 well separated

near inner distal corner.

DESCRIPTION OF MALE. As in female except for following

characters.

Body (Fig. 9). Length 1.008 mm (n = 1); second and third

urosomites not fused and ornamental spinules on urosome

somewhat more robust (Fig. 11A).

Antennule (Fig. 10) 9-segmented, subchirocer with 6th

segment very swollen, geniculation between 6th and 7th

segments. Segmental fusion pattern: I, II, II-VI, [X—xII,

XIII, XIV-XX, XXI-XXIIT, XXIV-XXV, XXVI-XXVIII.

Armature formula: [1, 1, 11, 8, 1, 14+ae, 4, 3, 8]. Segment 6

very swollen with a complicated pattern of ridges and teeth

on anterior surface (Fig. 1OC-D). Segment 7 with 4 setae, 3

of which sagittiform, on anterior surface (Fig. 10E).

P1. Coxa with fewer spinule rows on anterior surface.

Inner spine on basis without spinule row at base; inner spine

less well developed and with finer spinules (Fig, SE) than in

female (Fig. 5D). Segments of both rami (Fig. 5B) more

elongate than in female. Spinules on outer and distal margin

of endopod segments much finer than in female, particularly

on distal margin of enp-1 (Fig. 5C).

P2 (Fig. 6B-C). Basal pedestal and articulating surface of

endopod enlarged. Enp-1 much larger than in female and

inner seta transformed into a non-articulating process with a

flagellate tip; outer spinules small. Enp-2 without inner seta

or spinule row on outer margin; outer distal corner attenu-

ated into an apophysis reaching well beyound the distal

margin of enp-3. Enp-3 (Fig. 6C) reduced in size with no

outer spinule row; outer distal spine shorter but stouter than

in female with spinules reduced to coarse blunt teeth; termi-

nal and inner setae also reduced in size compared to female.

P3 endopod (Fig. 7B—C). Enp-2 without outer spinule row;

outer and inner distal corners much more attenuated than in

female, apophysis at outer corner with hooked tip (Fig. 7C);

inner seta much smaller than in female.

PS (Fig. 11B). Baseoendopods of each leg fused medially;

not fused to exopod. Endopodal lobe reduced with 2 pinnate

setae of very unequal length. Exopod with 5 pinnate setae,

inner seta small, middle seta longest.

P6 a single plate fused to somite proximally (Fig. 11A),

with 3 pinnate setae on each side (Fig. 11C).

REMARKS

(i) Synonymy

The Alaskan material on which the above redescription is

based, was first described in detail in an excellent paper by

Fig. 2 Archisenia sibirica comb. nov. A, Rostrum and female antennule (armature omitted); B, female antennule (disarticulated); C,

antenna, anterior view; D, antennary endopod, posterior view of distal margin.

a SS eA cima anise ae

REVISION OF DANIELSSENIA AND PSAMMIS

SSS

os t

TAIHON

“AAI\( My Y i

So Be,

)

‘o)

ig. 3 Archisenia sibirica comb. nov. A, Mandibular palp; B, mandibular gnathobase; C, labrum; D, maxillule.

52 R. HUYS AND J.M. GEE

Fig. 4 Archisenia sibirica comb. nov. A, Right paragnath, posterior view; B, maxilla with disarticulated endopod; C, maxilliped.

REVISION OF DANIELSSENIA AND PSAMMIS

ig. rchisenia sibirica comb. nov. A, Female P1, anterior view; B, male P1, protopod and endopod, anterior view; C, male P1, distal

| margin of enp-1 of other side; D, female P1 inner basal spine; E, male P1 inner basal spine; F, caudal ramus, lateral view.

Y i I uw v/a

o. }

= Y,

view; C, male P3, detail of outer apophysis

view; B, male P3 endopod, anterior

. A, Female P3, anterior

omb. nov

sibirica c

ig. 7 Archisenia

of enp-2.

R. HUYS AND J.M. GEE

Gif

——

~ AF y

Lbs 5

Se ;

i <a

ys SF TT

WAX

ISN

\ Was Ay KS

REVISION OF DANIELSSENIA AND PSAMMIS

oe teprrnag! ME ae Wig

wo"

WALES

ESN

, “f

’ | ret a en,

vp fi " : ~ ‘m

ZZ ere — Nein

us .

yy y

per ee NTS SAS sy

IED

ly Nw?

Me cert SA

AES

Fig. 9 Archisenia sibirica comb. nov. A, Male habitus, dorsal; B, same, lateral. [Sensillae on cephalothorax omitted. ]

58 R. HUYS AND J.M. GEE

Fig. 10 Archisenia sibirica comb. nov. A, Male antennule (armature omitted); B, male antennule, disarticulated (armature of segment 6

omitted); C, male antennule segment 6, anterior view; D, same, ventral view; E, male antennule segment 7, anterior view.

REVISION OF DANIELSSENIA AND PSAMMIS

se at

mip} Mt —

Wilson (1966). There are slight discrepancies between these 2

descriptions which should be noted because of their possible

phylogenetic significance. The rostrum does articulate with

the cephalothorax and there are 2 accessory setae at the base

of the maxillipedal claw. In the female there is no aesthetasc

on the terminal segment of the antennule and the outer distal

corner of enp-2 in P2 and P3 is significantly attenuated but

that of enp-1 is normal (especially compared to the condition

in Psammis). In the male the antennule is distinctly

9-segmented with segment 6 being swollen and bearing an

aesthetasc; the distal outer element of P2 enp-3 is not

completely fused to the segment but articulates along the

anterior surface and partially articulates on the posterior

surface; and there is no sexual dimorphism in P4 enp-2 (Fig.

8B).

We have been unable to locate the type, or any other,

material of D. sibirica and therefore, like Wilson (1966), have

had to rely on Sars’ (1898) original description and figures. A

study of these has led us to conclude that there are no real

differences between D. stefanssoni and D. sibirica. We agree

with Wilson’s interpretation of the swimming leg setation.

The original copy of Sars’ paper in our possession also shows

faint lines where the seta should be on P2—P4 exp-1 and in no

other species in any paranannopid genus is this seta absent in

the female when it is present in the male. Although Sars

(1898) states in his text that the female antennule is

5-segmented, close scrutiny of his drawing (Plate X, Fig. 4)

reveals that the terminal segment is at least partially divided

into two segments and that the arrangement of the pectinate

spines (3 on segment 5 and 1 at the apex of segment 6) is

exactly the same as in D. stefanssoni. Further Sars has clearly

misinterpreted the segmentation of the male antennule in

that he has combined segments 1 and 2 as shown by the fact

that his segment 1 bears 2 setae, a condition found in no other

Paranannopidae. Wilson (1966) concluded that the only real

difference between D. stefanssoni and D. sibirica was the

absence in the latter of the distinctive outer distal spine on P2

enp-3 in the male. However, Sars’ drawing of this limb (Plate

X, Fig. 16) shows only 4 elements on enp-3 instead of 5, as in

the female, and it is possible that the crucial one (the outer

spine) is concealed behind the large apophysis on enp-2. This

interpretation is supported by the fact that, irrespective of the

degree of modification in males, the number of elements on

P2 enp-3 is always the same in both sexes, except in the

genera bearing claviform aesthetascs on the mouthparts (Gee

& Huys, 1991) where the distal outer spine is further modi-

fied into a non-articulating apical apophysis.

These observations, coupled with such factors as similarity

of size (they are the largest known members of the Paranan-

nopidae except for Psammis borealis Klie, 1939 whose taxo-

nomic position is unclear) and the peculiar identical

distribution of setae on the distal margin of the female P35,

lead us to the conclusion that the two species are synonymous

and have a circum-polar distribution. Fig. 13 shows that the

most easterly record of D. sibirica at Wrangell Island (Yash-

nov, 1935) is very close to the most westerly record of D.

stefanssoni on the Chukchi Sea coast of Alaska (Wilson,

1966). The only record of the species outside the Arctic Circle

is that of Wells (1965) from Loch Nevis on the west coast of

Scotland and this must be regarded as doubtful (original

specimens no longer available for re-examination). Further,

both D. stefanssoni and D. sibirica have been recorded from

estuaries and in brackish water, a most unusual habitat for

members of this family. All other species are found only in

R. HUYS AND J.M. GEE

sublittoral marine habitats although Danielssenia typica

Boeck, 1872, the other species with a known global distribu-

tion, has been recorded from the Baltic Sea (Veldre &

Maemets, 1956; Arlt, 1983), a region of lowered salinity.

(ii) Autapomorphies of Archisenia

Turning now to consider the taxonomic status of D. sibirica,

its distinction from other members of the genus Danielssenia

was suggested by Lang (1944) who divided Danielssenia into 2

groups, the typica group and the sibirica group. The latter he

characterized by: (i) antenna exp-1 with 2 setae; (ii) P4 enp-3

with 2 inner setae; (iii) the male P2 enp-1 with the inner seta

transformed into a non-articulating process; (iv) the male P3

enp-2 with an outer hooked apophysis. This last character is

now known to occur in all species of Paranannopidae and

might even be a diagnostic character for a wider group of

families. The first two characters, although diagnostic of D.

sibirica in combination, are the plesiomorphic condition in

the family and are found in a number of other genera.

Paranannopus, Psammis, Micropsammis Mielke, and

Bathypsammis gen. nov. also retain 2 setae on exp-1 of the

antenna (all other genera bear only 1 seta on this segment)

and Sentirenia Huys & Gee (= Jonesiella Brady, see below)

and the male of Fladenia retain 2 inner setae on P4 enp-3.

However, the transformation of the inner seta into a non-

articulating spine on P2 enp-1 in the male is unique to this

genus, as are the following autapomorphies: (i) the outer

extension on P3 enp-2 in both sexes; (ii) the sigmoid, heavily

sclerotized female copulatory duct; (iii) the sexual dimor-

phism of the inner basal spine of the male Pl. Another

character of phylogenetic significance is the loss of the inner

seta on the male P2 enp-2. This character is also found in

Afrosenia spinipes (Wells, 1967) and is regarded here as a

product of convergence. A further character which is unique

but difficult to quantify is the arrangement of the setation on

the exopod of PS, with the inner seta well separated from the

remaining setae. It is on the basis of all these characters that

we have removed D. sibirica to a new genus leaving the typica

group as the only species group in the genus Danielssenia.

(iii) Intersexuality

The male from East Greenland (Fig. 12), collected by Jes-

persen, proved upon examination to be an aberrant intersex-

ual specimen. It has the male body facies, including a

6-segmented urosome (Fig. 12A), a well developed testis and

vas deferens (however, a spermatophore has not been

observed), and the male form of the P5 and P6. The

endopods of P2 and P3 are also modified but differ from the

typical male condition by the retention of certain female

features.

The antennules resemble the female condition in all

aspects: they are 6-segmented, lack any trace of a genicula-

tion mechanism and possess the female armature pattern.

The P1 (Fig. 12B) basis and endopod show the same spinule

arrangement as in the female but size and shape of certain

spinule rows approach the male condition. The P2 endopods

are not identical on both sides (Figs. 12C-D) and show a

combination of male and female characteristics. The proximal

segment and its basal pedestal are moderately enlarged but

the spinule at the base resembles the female condition and

the inner seta is — though being shorter than in the female —

not transformed into a spinous process. The outer apophysis

Tig.

REVISION OF DANIELSSENIA AND PSAMMIS

Jkt See,

awit

—_—~

P er Soc AAG ge g

a = BELA

RSC SS

ee ee jit EE

eG ERA PRREN

EE SN <.

specimen from Greenland. A, Habitus, dorsal view; B, P1, endopodal segments; C, P2,

omb. nov. Intersex

sibirica c

endopod of right side; D, P2, middle endopod segment of left side; E, P3 endopod; F, PS.

g.12 Archisenia

of the middle segment is distinctly shorter than in the typical

male and its outer margin might bear spinules as in the

female; the inner seta — completely missing in the male — is

represented by a vestigial spine which is either entirely (Fig.

12C) or partly (Fig. 12D) invaginated. The distal segment is

almost identical to the female condition. The P3 endopod is

modified as in the male except that the inner seta of the

middle segment is distinctly longer than in the typical male

(but shorter than in the female). The P4 endopod grossly

resembles the male condition. The P5 also has the basic male

outline but the endopodal lobe is slightly more pronounced

and the inner exopodal seta is distinctly longer.

Intersexuality within the Harpacticoida appears to be very

rare. Klie (1944) describes a female specimen of Amphias-

coides debilis (Giesbrecht, 1881) from Helgoland which dis-

played the male condition for the antennules (i.e. haplocer)

and the first thoracopods (i.e. modified basis) and the female

condition for the genital somite and the remaining append-

ages except for the P2 endopod which combined both male

and female features. Recently, Moore & Stevenson (1991)

found that 90% of a population of Paramphiascella hyper-

borea (T. Scott, 1903) in the vicinity of a sewage outfall in the

Firth of Forth, Scotland, were intersex specimens. In the

majority of these the prosome (including the antennules and

swimming legs) exhibited the female condition whilst the

urosome had the male condition of 6 distinct somites and a

plate-like P6, although the PS was more similar to that of the

female. At the same site, a small number of intersex speci-

mens of Stenhelia gibba Boeck, 1864 and Halectinosoma

similidistinctum Lang, 1965 were also found. Intersexuality is

more common in other orders of copepods, particularly the

calanoids Eudiaptomus vulgaris (Schmeil, 1898), Arctodiap-

tomus (Rhabdodiaptomus) alpinus (Imhof, 1885), Eudiapto-

mus gracilis (Sars, 1863), Pseudocalanus elongatus (Boeck,

1864), Calanus hyperboreus Kroyer, 1838, Paracalanus par-

vus (Claus, 1863) (Bremer, 1914; Pirocchi, 1940; Cattley,

1949; Francois, 1949; Conover, 1965; Ianora et al., 1987) and

cyclopoids Megacyclops gigas (Claus, 1857) and Megacyclops

viridis (Jurine, 1820) (Mrazek, 1913; Coker, 1938).

In natural populations the frequency of occurrence of

intersexuality appears to be very low and may be a result of

infrequent chromosomal aberrations during embryonic devel-

opment. In cases of higher incidence, various causes of

intersexuality have been postulated. Coker (1938) attributed

it to low temperature during naupliar development; Cattley

(1949) to parasitism of the developmental stages by the

marine ectoparasitic dinoflagellate Blastodinium contortum

hyalinum Chatton; and Moore & Stevenson (1991) argued

that the very high incidence of intersexuality in the vicinity of

a sewage outfall strongly implicated some form of chemical

pollution as the causative factor.

Genus Danielssenia Boeck, 1872

Since the publication of Lang’s (1948) monograph a number

of new species have been assigned to the genus Danielssenia

but recent analyses have shown this to be a heterogeneous

assemblage. In previous papers (see also Table 1) we have

removed D. intermedia Wells, 1965 to the genus Fladenia; D.

perezi Monard, 1935 (syn. D. paraperezi Soyer, 1970) and D.

eastwardae Coull, 1971 to the genus Sentirenia and propose to

remove D. spinipes Wells, 1967 and D. minuta Coull, 1969 to

two other new genera (Gee & Huys, 1990; Huys & Gee,

1992, in press). This has restricted the genus Danielssenia to

R. HUYS AND J.M. GEE

the following species: D. typica; D. quadriseta Gee, 1988; D.

reducta Gee, 1988; D. robusta Sars, 1921 and D. similis

Chislenko, 1971. The status of D. fusiformis (Brady, 1880),

previously been synonymized with D. typica (cfr. Shen & Bai,

1956; Gee, 1988b) is reconsidered here.

(i) Danielssenia fusiformis (Brady, 1880) nec Sars (1910)

Brady (1880) created the genus Jonesiella to accommodate

two new species, J. fusiformis (Brady & Robertson) and J.

spinulosa (Brady & Robertson), and provided illustrated

descriptions for these species. Brady remarked that both

species had already been listed in an earlier report (Brady &

Robertson, 1876) as Zosime (?) fusiformis and Z. spinulosa,

respectively, and therefore unjustly concluded that both

authors had to be credited with the authorship. This state of

affairs has perpetuated in the nomenclature, even to the

present (e.g. Gee, 1988b), though it is clear that Brady &

Robertson’s species names are mere nomina nuda and only

Brady (1880) should be cited as the author of both Jonesiella

species. Norman & Scott (1906) were the first to list J.

spinulosa Brady, 1880 as a junior synonym of Danielssenia

typica Boeck, 1872 and also changed J. fusiformis Brady,

1880 into D. fusiformis. Both species were redescribed and

illustrated by Sars (1910) who admitted that they were very

similar. It were also Sars’ descriptions that led Shen & Bai

(1956) to conclude that both species were identical, and after

careful examination of Sars’ material Gee (1988b) formally

relegated D. fusiformis sensu Sars (1910) to a junior synonym

of D. typica. There is, however, considerable evidence that

what Sars (1910) considered to be D. fusiformis in Norway is

clearly different from Brady’s (1880) original material from

the Scilly Islands. Brady’s type material does no longer exist,

but his illustrations (Plate XLVIII, Figs 1-13) of the female

antennule, mandible, maxilliped, P1, the fifth legs in both

sexes and the male endopod P2 leave no doubt that his

species is identical with D. perezi Monard, 1935, originally

described from Roscoff and later also recorded from the

Scilly Islands (Wells, 1968) — the type locality of J. fusi-

formis. Huys & Gee (1992) recently synonymized D. para-

perezi Soyer, 1970 with D. perezi and established a new genus

Sentirenia to include the latter species and D. eastwardae

Coull, 1971. Sentirenia Huys & Gee, 1992, therefore, has to

be relegated to a junior synonym of Jonesiella, thus encom-

passing the type species J. fusiformis Brady, 1880 nec Sars

(1910) (syn. nov.: Danielssenia perezi Monard, 1935; D.

paraperezi Soyer, 1970) and J. eastwardae Coull, 1971 comb.

nov.

Thompson’s (1893) illustration of the female antennule

suggests that his record of J. fusiformis from Liverpool Bay is

correct. Re-examination of specimens (7 ?@ labelled D.

fusiformis; Norman collection, reg. no. 1911.11.8.43561-565,

gift from T. Scott; October 1899) collected in the Firth of

Clyde indicates that the species might be distributed along the

entire west coast of Britain. Lang’s (1936a,b) specimens from

the Oresund and Spitzbergen clearly belong to D. typica. All

other records of D. fusiformis have to await confirmation (see

list in Lang, 1948).

(ii) Danielssenia robusta Sars, 1921

Lang (1948) was of the opinion that D. robusta (and D.

perezi) probably would require the definition of additional

species groups inside the genus but as the males were still

REVISION OF DANIELSSENIA AND PSAMMIS

ig. 13 Distribution map of Danielssenia sibirica (circles) and D. stefanssoni (stars). Records of 1. Sars (1898); 2. Yashnov (1935); 3. Wells

(1965); 4. Willey (1920); 5. Jespersen (1939); 6. Wilson (1966). Arctic Circle shown by dashed line.

unknown at that time he regarded such an allocation as being

premature. We have re-examined Sars’ material of this

species from Risgr, Norway (13 2 9 and 1 copepodid V stage;

Zoologisk Museum, Oslo, Reg. No. F20257) and found the

following significant discrepancies from the original descrip-

tion of Sars (1921): (i) there is an inner seta on exp-1 of

P2-P4; (ii) the inner element on enp-1 of P2—P4 is a pinnate

spine; (iii) the PS baseoendopod has 4 setae, the second inner

one being much smaller than the others; (iv) the PS exopod is

fused to the baseoendopod on the posterior surface. Further,

we have made a detailed comparison of these females with

the recently discovered female of D. intermedia (which was

assigned to the genus Fladenia by Gee & Huys (1990)) and

have found them to be identical. Therefore D. robusta must

be referred to the genus Fladenia whose type species now

becomes F. robusta (Sars, 1921) comb. nov. as this has

priority over F. intermedia (Wells, i965). F. robusta has also

been recorded from the Mediterranean by Por (1964), who

found one female in 470 m off the coast of Israel, and Soyer

(1970) who found 18 adult females at depths ranging from 50

to 420 m in the vicinity of Banyuls-sur-mer. Both authors

state in their text that these specimens agree exactly with the

original description. However, Por’s (1964) Fig. 73 does show

an inner seta on exp-1 of what is probably P4 (this limb is

labelled P1 by Por but cannot possibly be so as the endopod is

3-segmented). This, taken in conjuction with his figure of the

P5 (1964, Fig. 74) leaves little doubt that the Mediterranean

material can be assigned to F. robusta, thus giving this species

a Boreo-Mediterranean distribution similar to that of Jone-

siella fusiformis (see Huys & Gee, 1992).

(iii) Danielssenia similis Chislenko, 1971

Chislenko (1971) distinguished D. similis from D. typica on

the basis of the following characters: (i) Size, the specimen

drawn in his Fig. 1 is approximately 0.9 mm long; (ii) a

maxilliped with only 1 seta on the syncoxa and a somewhat

longer seta on the basis; (iii) the sexual dimorphism on P2

endopod, with the loss of the inner seta on enp-1 and of | seta

on enp-3. The character of size is of no particular significance

as it is within (but near the upper limit of) the size range of D.

typica given by Gee (1988b). Similarly, the absence of a large

seta on the basis of the maxilliped is of doubtful significance

as this seta can be easily dislodged during dissection, as was

the case in Sars’ (1910) description of D. typica (see Gee,

1988b). The differences in sexual dimorphism of the male P2

endopod are more difficult to assess from drawings alone.

However, it is highly improbable that the inner seta on P2

enp-1 is missing in the male when it is present in the female as

this condition is found in no other member of this genus or

indeed of the family as a whole. The same goes for a

reduction in the number of setae on enp-3. In Danielssenia,

the 2 terminal setae on this segment are very reduced and

implanted close together and it is conceivable that Chislenko

(1971) has combined these 2 fine setae and drawn them as

one broad one. We believe that D. similis is referable to D.

typica which has been shown to be the most variable species

in the genus (Gee, 1988b) but without being able to examine

topotype material we must regard it as a species inquirenda.

(iv) Danielssenia typica Boeck, 1872

The following material of the Norman collection (The Natu-

ral History Museum) has been examined (species name given

R. HUYS AND J.M. GEE

on the original museum label presented in parentheses):

1911.11.8.43451-470: vial containing > 400 specimens,

mostly 99, a gift from T. Scott; collected near Duke Buoy,

Plymouth, 01 August 1889;

1911.11.8.43471-490: vial containing 23 99 and 10’, a gift

from T. Scott; collected from Varanger Fjord, East Finmark,

Norway, 1890;

1911.11.8.43491—-510: vial containing 31 99 and3 OC ,a

gift from T. Scott; collected from Vads6, East Finmark,

Norway, 03 July 1890;

1911.11.8.43511-530: vial containing 39 specimens (D.

typica), a gift from T. Scott; collected in Trondhjem Fjord,

Norway, 1893; 32 2@ belong to D. typica, the other 7 29

belong to two different species of Halectinosoma;

1911.11.8.43531-540: vial containing 16 specimens (D.

typica), a gift from T. Scott; collected from Inchkeith in Firth

of Forth, October 1895. None of these specimens belongs to

D. typica, instead the vial contained Bradya sp. (2 29, 8

copepodids), 2 O&'C’ Robertsonia tenuis (Brady & Robert-

son), 1 Q Idomene coronata (T. Scott) and 3 QQ of a

Fladenia-like paranannopid;

1911.11.8.43541—-560: vial containing > 1000 specimens,

mostly 2Q, a gift from T. Scott; collected from Kames Bay,

Isle of Cumbrae, 1888; a second lot of about 200 specimens

from the same locality is registrated under no. 1900.3.29.274;

1911.11.8.M.2299: 1 Q dissected on slide (Jonesiella spinu-

losa), dried out; collected in Trondhjem Fjord, Norway,

1893;

1911.11.8.M.2301: 43 specimens mounted in toto on slide

(Jonesiella spinulosa), dried out; collected near Duke Buoy,

Plymouth, 02 August 1889;

1911.11.8.M.2300: 8 specimens mounted in toto on slide

(Jonesiella spinulosa), dried out; collected from Vads6, East

Finmark, Norway, 1890;

1900.3.6.644: 5 QQ mounted in toto and 3 QQ (one

belonging to Halectinosoma sp.) dissected on slide (Jonesiella

spinulosa); collected in Trondhjem Fjord, Norway, 1893.

Gee’s (1988b) redescription of D. typica is updated here by

the following observations and illustrations (Figs 14-16)

based on specimens from Duke Buoy (closest to type local-

ity):

Somatic hyaline frills of pedigerous and abdominal somites

minutely dentate (Fig. 14A) except for the dorsal frill of

P5-bearing somite which is deeply incised, forming rectangu-

lar lappets (Fig. 14A, B). Frill of cephalothorax smooth.

Dorsal transverse spinule rows are found only on thoracic

somites bearing P3—PS5, the genital double-somite and second

abdominal somite. Genital double-somite with continuous

transverse chitinous rim dorsally, laterally and ventrally,

marking original segmentation (Figs. 14A, D; 16D-E).

Pseudoperculum (Figs. 14E-F) formed by deeply incised

posterior extension of penultimate somite. Pattern of caudal

rami setae as in Figs. 14E-F.

Rostrum (Fig. 161) large, hyaline, with 2 pairs of minute

sensillae; typically deflected (Figs. 14A—C).

Male antennule (Fig. 15G) 8-segmented or indistinctly

9-segmented; distal 2 segments very small and largely fused.

Mandible with blunt teeth and a single pinnate seta on

gnathobase (Fig. 15A). Palp with short, equally long,

l-segmented rami (Fig. 15B); basis with row of very long

setules proximally, inner margin with 1 short and 2 long

setae; endopod with 2 lateral and 6 apical setae; exopod with

00 oo,

pale

pp ptm 0 pene

geen PR Se

gests

oo,

‘ig. 14 Danielssenia typica. Female: A, habitus, dorsal; B, rostrum and anterior part of cephalothorax, ventral; C, same, lateral;

| _D, pleurotergite of P4-bearing somite, P5-bearing somite with fifth thoracopod and genital double-somite, lateral; E, pseudoperculum,

anal somite and left caudal ramus, lateral; F, same, dorsal. [Incised hyaline frill of P5-bearing somite arrowed in A and D.]

labled

VERN

YER

< <S

Ww Ves

wy a

Fig. 15 Danielssenia typica. A, Mandible, gnathobase; B, mandible, palp; C, maxillula, posterior, showing disarticulated praecoxa, coxa and

palp; D, maxilla, showing disarticulated syncoxa, basis and endopod; E, maxilliped, anterior; F, maxilliped, posterior; G, male antennule

(armature ommited). [Tubular setae arrowed in C-D.]

- REVISION OF DANIELSSENIA AND PSAMMIS

row of long setules at 1/3 distance from the proximal margin,

and 1 lateral, 1 subapical and 2 apical setae.

Maxillule (Fig. 15C). Praecoxal arthrite with 9 spines and 1

tubular seta around the distal margin, and 2 geniculate

tubular setae on the anterior surface; coxal endite with 1

pinnate spine, 1 setulose claw, 1 smooth setae and 3 tubular

setae; basal endites closely set, proximal with 2 setae, distal

with 2 setae and 2 spines; rami with 3 setae each.

Maxilla (Fig. 15D). Praecoxal endite with 2 unilaterally

pinnate spines and 1 basally fused spine bearing tubular

extension. Coxal endites with 1 spine and 2 tubular setae

each. Allobasis with 2 articulating claws, 1 pinnate spine and

1 tubular seta. Endopod with 1 tubular seta, 1 spine and 2

pinnate setae.

Maxilliped (Figs 1SE-F). Syncoxa with 1 large setulose

spine posteriorly and 1 smaller pinnate seta anteriorly. Basis

with anterior row of long spinules and small, pinnate seta on

palmar margin. Endopodal claw with 2 accessory setae.

Intercoxal sclerites of P2—P4 (Fig. 16H) not U-shaped and

provided with large spinules on anterior surface (as in Archis-

enia).

P2 endopod of male (Figs 16A—B). Inner setae of proximal

and middle segments reduced compared to the female. Outer

apophysis of middle segment very large, reaching far beyond

the distal segment. Inner setae of distal segment spiniform

and stouter than in the female; inner terminal seta reduced,

with spatulate tip; outer terminal seta represented by small

setule; outer spine curved at tip and standing on cylindrical

| process.

P3 endopod of male (Fig. 16C) with acutely recurved

| process anteriorly at outer distal corner of middle segment.

Genital field with minute copulatory pore (Fig. 16D)

_ leading via short sclerotized duct to multi-chambered seminal

receptacle (Figs 16F—G). Copulatory duct entering unpaired

| ventral chamber leading dorsally to paired reservoirs both

| anteriorly and posteriorly. Anterior reservoirs largest and

_ extending to posterior part of P5-bearing somite (Fig. 16D).

| P6 in female represented by 1 plumose seta and 2 minute

| spiniform elements (Fig. 16E). P6 of male with 1 plumose and

| 1 pinnate seta (Fig. 16J).

REMARK. Shen & Bai (1956) pointed out that either 1 or 2

setae can be found on the middle endopod segment of P2,

however, their figured specimen with 2 setae on this segment

| (Plate XI, Fig. 86) is almost certainly an aberrant case. The

| same applies to the armature of the baseoendopod of the

| male specimens reported on by Gee (1988b) where a ‘vari-

' able’ number of setae can be found; all setation patterns

| diverging from the typical bisetose condition are aberrations

caused by abnormal copepodid development.

(v) Amended diagnosis

As a result of the arguments put forward above, the genus

Danielssenia now contains only 3 well defined species and we

have re-diagnosed the genus accordingly:

Paranannopidae. Body variable in size, slightly fusiform

and dorso-ventrally flattened. Rostrum hyaline, large, typi-

cally deflexed, with 2 pairs of small sensillae. Somatic hyaline

frills minutely dentate except for deeply incised frill on dorsal

margin of PS-bearing somite. Original segmentation of

female genital-double somite marked by complete sub-

cuticular ridge; genital field with small copulatory pore; short

copulatory duct leading to seminal receptacle with paired,

anteriorly directed chambers extending to anterior margin of

genital double-somite; P6 with 1 outer plumose seta and 2

minute spiniform elements. Pseudoperculum hyaline with

deeply incised margin. Caudal rami parallel, broader than

long, seta I minute. Female antennule 4-segmented; aes-

thetasc on segment 3; terminal segment with strong pinnate

spines. Antennary exopod 3-segmented with armature for-

mula [1-1-3]. Mandibular coxa with blunt teeth and 1 seta on

gnathobase, basis broad with 3 setae on distal margin;

endopod 1-segmented; exopod 1-segmented, with 1 lateral

and 3 distal setae. Maxillule with 3 tubular setae, 1 pinnate

seta and 1 spine on coxal endite; basal endite with 4 setae and

1 spine. Maxilla with tubular setae on coxal endites, allobasis

and endopod; praecoxal endite with 3 pinnate spines. Maxil-

liped subchelate with 1 large and 1 small seta on syncoxa;

basis with small pinnate seta on palmar margin, endopodal

claw with 2 accessory setae. Pl exopod 3-segmented, exp-3

with distal outer spine longer than middle outer spine;

endopod 2-segmented, enp-2 4 times longer than broad, inner

seta implanted medially. P2—P4 intercoxal sclerites with

spinules on distal margin; rami 3-segmented; exp-1 without

inner seta. Armature formula of P1—P4 as follows:

Exopod Endopod

PA 0.1.023 1.121

P2 0.1.(1-2)23 220

P3 0.1.(1-2)23 1.1.(1-2)21

P4 0.1.(2-3)23 1.1.(0-1)21

Female fifth pair of legs not fused medially; exopod and

baseoendopod separate, each with 4 or 5 setae.

Male with sexual dimorphism on antennule, P2 endopod,

P3 endopod, P5, P6, and in genital segmentation. Antennule

8- to 9-segmented, subchirocer; segment 6 very swollen, with

aesthetasc. P2 enp-2 with inner seta, outer distal corner

attenuated into a long apophysis reaching far beyond the

distal border of enp-3; enp-3 with distal outer spine and 2

terminal setae very reduced, inner setae spiniform and larger

than in female. P3 enp-2 with outer distal corner attenuated

into a recurved apophysis. PS of each side fused medially;

baseoendopod and exopod separate with 2 and 4 or 5 setae,

respectively. P6 symmetrical, fused to somite, with 2 setae

each.

TYPE SPECIES. D. typica Boeck, 1872 (by monotypy). [syn.:

D. fusiformis (Brady, 1880) sensu Sars (1910)].

OTHER SPECIES. D. quadriseta Gee, 1988; D. reducta Gee,

1988.

SPECIES INQUIRENDA. D. similis Chislenko, 1971.

Key to species

1. P2—P3 exp-3 with 2 inner setae, P4 enp-3 with 1 inner seta ... 2.

P2-P3 exp-3 with 1 inner seta, P4 enp-3 without inner seta

baeT ABBR sanseee She ooe BOE Oot On SERCO TEETER EIT D. reducta Gee, 1988.

2. P3enp-3 with 2 inner setae, PS exopod with 5 setae in both sexes

Bat ect nie sae etse a eee elteiacmoecec eens creer D. typica Boeck, 1872.

P3 enp-3 with 1 inner seta, PS exopod with 4 setae in both

SEXES Hee ee ees haces dere rae decctuet teeaene D. quadriseta Gee, 1988.

R. HUYS AND J.M. GEE

sterior; C, P3 endopod, middle segment,

tal slit and copulatory p F, seminal

opod, po

1; E, genital slit and copulato ore; F, semina

al sclerite P3; J, sixth leg. Female: D, genital double-somite, ventra

receptacle, lateral; G, same, ve

2 end

ia typica. Male: A, P2 endopod, anterior; B, distal segment of P

1; H, intercox

Fig. 16 Danielssen

anterio

ntral; I, rostrum.

REVISION OF DANIELSSENIA AND PSAMMIS

Genus Bathypsammis gen. nov.

SYNONYMY. Psammis Sars, 1910 (part.).

DIAGNOSIS. Paranannopidae. Body large, more or less cylin-

drical. Rostrum not hyaline, with 2 pairs of sensillae, anterior

pair large. Somatic hyaline frills minutely dentate. Female

genital double-somite with lateral and ventral sub-cuticular

ridge, marking original segmentation; copulatory pore

minute; copulatory duct and seminal receptacle unconfirmed;

P6 with 2 setae and 1 setule in between. Pseudoperculum

hyaline, vestigial. Caudal rami divergent, elongate (length

about 5 times proximal width); with tuft of long setules near

inner distal corner; dorsal surface with chitinized rim in

anterior half. Female antennule 4-segmented; aesthetasc on

segment 3; distal 2 segments with heavily pectinate spines.

Antennary exopod 3-segmented with armature formula

[2-1-3]. Mandibular coxa elongate, gnathobase with blunt

teeth and spinule row; basis broad with 4 setae on distal

margin; endopod 1-segmented, slightly longer than exopod;

exopod 1-segmented with 1 lateral and 2 apical setae. Maxil-

lule without modified spines on coxal endite; basal endite

with 5 setae. Maxilla without tubular setae; praecoxal endite

with 3 pinnate spines (1 fused to endite). Maxilliped subche-

late; armature of syncoxa unconfirmed; basis with naked seta

on palmar margin, endopodal claw with 2 accessory setae. P1

with very long outer basal seta reaching to middle of exp-3;

exopod 3-segmented, exp-3 with distal outer spine longer

than middle outer spine; endopod shorter than exopod;

2-segmented, enp-2 as long as enp-1, inner seta implanted at

1/3 distance from proximal margin. P2—P4 intercoxal sclerites

with few long setules; rami 3-segmented; exp-1 with inner

seta; female P2—P3 enp-2 without apophysis at outer distal

corner. Armature formula of P1—P4 as follows:

Exopod Endopod

Pl 0.1.023 Lest

P2 1223 1.2.121

P3 1.1.323 10.120

P4 1.1.323 Plea 7

Female fifth pair of legs not fused medially; exopod and

baseoendopod fused to form a bilobate plate; exopodal lobe

with 2 spines and 2 setae; endopodal lobe with 2 setae and 3

spines, the outer 2 of which are stubby.

Male unknown.

TYPE SPECIES. Bathypsammis longifurca (Bodin, 1968)

comb. nov.

OTHER SPECIES. None.

ETYMOLOGY. The generic name is derived from the Greek

bathys, meaning deep, and Psammis, probably the most

closely related genus known in the Paranannopidae. Gender:

feminine.

| Bathypsammis longifurca (Bodin, 1968) comb. nov.

SYNONYMY. Psammis longifurca Bodin, 1968.

MATERIAL EXAMINED. From Dr Ph. Bodin: holotype dis-

sected on 3 slides and now deposited in the collections of The

\Natural History Museum under reg. no. 1992.1091; Bay of

Biscay, Stn 308 (46°07' N; 05°00’ W), depth 3950 m; coll.

August 13 1963, R/V Job-ha-Zelian.

Bodin’s (1968) excellent original description is supple-

mented here by the following observations and Figures

17-18.

Antennule 4-segmented, third segment homologous to

segments 3-4 in Archisenia. Distal segment with large, swol-

len seta anteriorly near proximal corner; as pointed out by

Bodin this segment is seemingly subdivided by the raised

insertion site of one of the large pectinate spines (Fig. 18A).

Armature formula: [1, 8, 14+ae, 16].

Mandibular gnathobase (Fig. 17B—C) with 4 long teeth,

one trifid, slender element and 1 pinnate seta; a comb of

spinules is present at the base of the smaller teeth. The

endopod has 1 outer, 1 subapical and 6 apical setae (2 of

which are fused basally).

Maxillule (Fig. 17D). Praecoxal arthrite with 9 spines and 1

tubular seta around the inner margin, and 2 geniculate

tubular setae on the anterior surface; coxal endite with 4

setae, pinnate spine and straight spine with defined flexure

zone and small pore near the apex; basal endite with 5 setae.

Maxilla (Fig. 18B—C) with praecoxal endite drawn out into

heavily pectinate spine and bearing 2 articulating elements;

coxal endites with 1 serrate spine and 2 setae each. Allobasis

with 1 short and 2 long setae; endopod with 4 setae.

Maxilliped (Fig. 18D). Syncoxa missing in preparation.

Basis with naked seta on inner margin, 2 spinular rows on

anterior surface and another one on posterior surface;

endopodal claw with 2 accessory setae.

Intercoxal sclerites of P2—P4 U-shaped (as in Fladenia) and

provided with few long setules near lateral margins.

Fifth leg (Fig. 17E) with 2 large tube pores on anterior

surface.

Genital field (Fig. 18E) with minute copulatory pore. The

internal structures of the genital double-somite were

destroyed during the dissection, so no observations of the

copulatory duct and the seminal receptacles could be made.

Pseudoperculum very weakly developed. Distribution of

caudal rami setae as in Fig. 18F—G; seta III dislodged in both

rami, insertion site indicated by small socle (Fig. 18G).

P. longifurca does have certain features in common with P.

longisetosa and P. longipes, namely: anterior pair of rostral

sensillae enlarged (Fig. 17A); only 1 lateral seta on both rami

of the mandible (Fig. 17B); 2 setae on exp-1 of the antenna; 2

setae on enp-2 of the P2; and fused rami in the female P5.

However, P. longifurca lacks certain important features

shared by the other two species, namely: no large strongly

pinnate seta on the basis of the maxilliped, the seta on this

segment being small and naked (Fig. 18D); the endopod of

P2 is not distinctly longer than the exopod; the proximal inner

seta of P2 enp-2 is not displaced to the posterior surface; the

inner distal seta of P3—P4 enp-3 is not reduced; and, there is

no attenuation of the outer distal corner of P2 enp-1. Finally,

P. longifurca has a number of characters which are not shared

by the other members of this genus such as: (i) a plume of

long fine setules at the inner distal corner of the caudal

ramus; (ii) an outer basal seta on P1 which is nearly as long as

the exopod; (iii) a Pl endopod which is shorter than the

exopod and in which both segments are equal in length; (iv) a

P5 with peculiar spines on the endopodal lobe and a minute

outer basal seta; (v) a primitive setal formula for the exopods

of the swimming legs which is shared only by Archisenia and

Jonesiella. On the basis of these characters we assign P.

R. HUYS AND J.M. GEE

SS Wwe

ts...

=SS SF

a See

DS SSS ESOS

Le eg

v. Female: A, rostrum; B, mandible; C, mandible, gnathobase; D, maxillule, praecoxa

mb. no

rticulated; E, PS, anterior.

Fig. 17 Bathypsammis longifurca co

disa

REVISION OF DANIELSSENIA AND PSAMMIS

\ vil

g- 18 Bathypsammis longifurca comb. nov. Female: A, antennule, distal segment; B, maxilla, allobasis and endopod; C, maxilla, syncoxal

endites; D, maxilliped, anterior (syncoxa missing); E, genital apertures and copulatory pore (arrowed); F, anal somite and left caudal

ramus, dorsal; G, caudal ramus, detail of posterior margin.

longifurca to a new genus Bathypsammis which is closely

related to Psammis.

Genus Psammis Sars, 1910

With the removal of P. longifurca to Bathypsammis gen.

nov., the number of species currently allocated to the genus

Psammis is reduced to four: P. longisetosa Sars, 1910; P.

borealis Klie, 1939; P. kliei Smirnov, 1946; and, P. longipes

Becker, 1974.

(i) Psammis borealis Klie, 1939

This species was first briefly diagnosed in 1939 from material

collected in deep water near Iceland. A more extensive

description, accompanied by illustrations, was published in

1941. Any justification for placing this species in Psammis is

missing from Klie’s (1939, 1941) papers, providing instead a

large number of fundamental differences with the type spe-

cies P. longisetosa. We have re-examined Klie’s type material

of P. borealis (Cop. 211-215; 4 99, 1 GC, all dissected on

slides; Zoologisches Museum der Universitat Kiel). The slide

of the male is somewhat confusing in that there seems to be 3

mounted antennules which do not show male characteristics

and only part of one which does have the features of a male.

Further, the limbs on this slide show no sexual dimorphism

on either P2 or P3. The genital somite is also missing and the

only appendage that differs from the slides of the females is

the PS. The fifth legs of both sexes are exactly as drawn in

Figs. 4 & 6 in Klie (1941). However, based on the mouthparts

and the setation of the female thoracopods, and pending

more information on swimming leg sexual dimorphism, we

propose to retain this species within the Paranannopidae as

species incertae sedis. It should be noted here that the

specimens labelled P. borealis and deposited in the Smithso-

nian Institution (reg. no. 00231018) by Prof. Dr B.C. Coull

are not the same genus as that of Klie (1939). This material (2

2) collected from the North Carolina continental shelf [this

record is not listed in Coull (1971)] closely resembles

Pseudotachidius similis T. Scott, 1902 and P. minutus It6,

1983.

(ii) Psammis kliet Smirnov, 1946

We have been unable to discover the type material of P. kliei

described by Smirnov (1946) from Henrietta Island (New

Siberian Islands, East Siberian Sea). However, the recent

recovery of a specimen from Spitsbergen which we believe is

referable to this species, indicates that it should be placed in

another genus close to Psammis and Danielssenia. This will

be discussed further in a future paper on the Paranannopidae

of Spitsbergen (Gee & Huys, in prep.).

(iii) Psammis longipes Becker, 1974

MATERIAL EXAMINED. Holotype @ dissected on 2 slides

(Becker collection; Zoologisches Museum der Universitat

Kiel, reg. no. 1009-1010); Peru Trough, R/V Anton Bruun

Sta. 179, 12°03’S 78°45'W, depth 5000 m, leg. W. Noodt.

This species is known from the type locality only. The

following redescription (Figs. 19-20) is confined to structures

that were misinterpreted or not well illustrated in Becker’s

(1974) original description:

Mandible (Figs. 19A—B). Gnathobase with multicuspidate,

elongate teeth descreasing in size dorsally, and with 2 pinnate

R. HUYS AND J.M. GEE

setae near the distal dorsal corner; coxa with large spinules

around the base of the palp. Basis with 3 setae, middle one

with shorter spinules. Endopod only slightly longer than

exopod, with 1 lateral and 3 apical setae; exopod with 1

lateral and 2 apical setae.

Maxillule (Fig. 19C—D). Praecoxal arthrite with 9 pinnate

spines and 1 tubular seta around the distal margin and with 2

geniculate tubular setae on the anterior surface. Coxal endite

specialized; armature consisting of 3 tubular setae and 3

spines; largest (= anterior) spine with broad base, a comb of

flat spinules along the inner margin and ending in a tubular

extension; middle spine also swollen at base and with fan of

non-articulating flat spinules arranged around the apex; pos-

terior spine with large spinule. Basal endite with 3 plumose

setae and 1 short spine with tubular extension. Endopod and

exopod with 3 setae each.

Maxilla (Fig. 19E). Praecoxal endite with 2 pinnate spines,

distal one with tubular extension. Coxal endites with 2 spines

and 1 seta each, distal spine and posterior seta with tubular

extension. Allobasis with 2 articulating claws and a tubular

seta on either anterior and posterior surface. Endopod with 1

simple and 3 tubular setae.

Maxilliped (Fig. 19F) as described by Becker (1974) except

that the endopodal claw bears an accessory seta.

The armature formula given by Becker for the swimming

legs is erroneous on two points: P3 enp-2 has only 1 inner

seta, the proximal one shown in his figure being an enlarged

spinule; P3—P4 exp-3 have and extra element distally, repre-

senting the reduced inner terminal seta (Fig. 20A—B).

Fifth leg (Fig. 20C). An incomplete furrow on the posterior

surface marks the original proximal margin of the endopodal

lobe. The 3 distal setae of this lobe are multipinnate.

Genital field (Fig. 20D) with small copulatory pore leading

via linear duct to bilobate seminal receptacle largely located

anterior to genital slit. P6 armature represented by pinnate

seta and 2 minute spinules (vestigial setae?).

Hyaline frill of all body somites finely dentate; pseudoper-

culum well developed (Fig. 20E). Pattern of caudal ramus

setae as in Fig. 20E.

(iv) Psammis longisetosa Sars, 1910

MATERIAL EXAMINED.

— Zoologisk Museum, Oslo: (a) G.O. Sars collection:

F20223: 1 Q (in alcohol) and 1 © (dissected); collected

from Farsund (type locality), Norway;

F20224: vial containing 19 9? and 6 O'C;; collected from

Risgr, Norway;

(b) F20929: 4 99 (2 on slides, 2 in alcohol), 3 OC (1 on

slide, 2 in alcohol); collected by J.A. Berg, deposited by J.M.

Gee, from Bjgrnehodebukta (59°42.8’N, 10°32.2’E),

Oslofjord, 35 m depth, June 1984;

— The Natural History Museum: 1992.1096: 1 OC (in

alcohol), 1 2 (on 6 slides), 1 © prosome (on 7 slides);

collected by R. Huys, from Frierfjord-Langesundfjord, 55 m

depth, spring 1985.

The original descriptions of P. longisetosa given by Sars

(1910, 1921) have been supplemented since by a complete

redescription by Gee (1988a). The single female collected

from Raunefjorden and figured by Por (1965) in all probabil-

ity does not belong to P. longisetosa. In addition to the

differences in the shape and armature of the P5 mentioned by

Por, substantial discrepancies appear from his illustrations of

the P1 (relative proportions of endopodal segments), last

- 19 Psammis longipes. Female: A, mandible, posterior; B, mandible, gnathoba erior; C, maxillule, posterior;

i or. [Tubular setae arro in

of coxal endite; E, maxilla, posterior (showing syncoxal spines enlarged); F

se, ant ‘

, maxilliped, posteri

D, maxillule, detail

wed in C-E.

| | R. HUYS AND J.M. GEE

~~ >>

fag oo

3 SS 77

Fig. 20 Psammis longipes. Female: A, P3 endopod, distal segment; B, P4 endopod, distal segment; C, PS, anterior; D, genital apertures and

copulatory pore (arrowed); E, posterior abdominal somites and left caudal ramus, dorsal. [Vestigial seta arrowed in A-B].

REVISION OF DANIELSSENIA AND PSAMMIS

abdominal somites (ornamentation) and caudal rami (shape).

Re-examination of P. /ongisetosa has revealed a number of

features that were overlooked or misinterpreted in earlier

descriptions. In many cases these observations have shown an

astonishing similarity in the detailed structure of the cephalic

appendages between P. longipes and the type species.

The rostrum is not hyaline (Fig. 22A); the anterior pair of

sensillae is enlarged. In the male the antennule is

9-segmented (Fig. 22A) and the segmental pattern is homolo-

gous to that of Archisenia.

Mandible (Fig. 21A—B). The gnathobase has similar multi-

cuspidate teeth and 2 pinnate setae. The basis has 4 setae; the

ornamentation of these setae shows that it is either the

proximalmost or following seta that is missing in P. longipes.

Both species have the same armature on the rami.

Labrum (Fig. 22B) with 1 large, median and a pair of

smaller secretory pores on the anterior surface, and long

spinules around the distal margin.

The detailed structure of the maxillule and maxilla is

exactly the same as in P. /ongipes, including the presence and

position of tubular setae and the modifications of the maxillu-

lary coxal endite.

The maxillipedal syncoxa has been invariably described as

possessing a single, very large, spinulose seta, corresponding

to the posterior seta in P. longipes; the smaller, setulose,

anterior seta in this species is further reduced to a minute,

pinnate spine in P. longisetosa (arrowed in Fig. 21C) and

approaches the length of the largest ornamental spinules, the

reason why it had been overlooked in previous descriptions.

The sexual dimorphism on the P2 endopod includes modi-

fications of the middle and distal segments (Figs 21D-—-E). The

anterior, spinous apophysis on the outer margin of the

proximal segment is not a sexually dimorphic feature since it

is also found in female specimens. The middle segment is

drawn out into a large apophysis not reaching to the end of

the distal segment and provided with an anterior secretory

pore near the apex; the inner margin has 2 distally serrate

setae, the proximal one being slightly displaced to the poste-

rior surface; these setae are distinctly longer in the female.

The distal segment possesses 4 articulating armature elements

corresponding to the 2 inner and 2 terminal setae in the

female; the outer spine in the female is modified in the male

and replaced by a short, spinous process distally.

As in P. longipes, the reduced inner terminal seta of P3—P4

enp-3, represented by a setule, has been overlooked thus far

(arrowed in Figs 21F-G). In the male the outer distal corner

of the P3 middle segment is transformed into an acutely

recurved process (Fig. 21F); the inner seta on this segment is

distinctly longer in the female.

The fifth legs of both sexes are as in Figs 22E and F,

respectively.

The original segmentation of the female genital double-

somite is marked by a transverse chitinous rib dorsally and

ventrally (Fig. 22D). The seminal receptacle is relatively

small (Fig. 22C); the P6 is represented by 1 plumose seta and

1 small spinule in the female; in the male the sixth legs are

fused and symmetrical, and bear 2 naked setae on either side

(Fig. 22G).

(v) Amended diagnosis

Only P. longisetosa and P. longipes are retained in the genus

Psammis, which is here redefined.

WS)

DIAGNOSIS. Paranannopidae. Body large, slightly fusiform

and dorso-ventrally flattened. Rostrum not hyaline, with 2

pairs of sensillae, anterior one large. Somatic hyaline frills

minutely dentate. Female genital double-somite with lateral

and ventral sub-cuticular ridge marking original segmenta-

tion; genital field with minute copulatory pore and linear duct

leading to transverse seminal receptacle located anterior to

genital slit; P6 with 1 plumose seta and 1-2 minute spinulose

elements. Pseudoperculum hyaline with dentate margin. Cau-

dal rami divergent and longer than broad, tapering slightly.

Female antennule 4-segmented; aesthetasc on segment 3; all

segments with pinnate setae and spines. Antennary exopod

3-segmented with armature formula [2-1-3]. Mandibular

coxa elongate, with finely pointed teeth and 2 setae on

gnathobase; basis broad with 3-4 setae on distal margin;

endopod l-segmented, equal in length to exopod, with

strongly reduced armature; exopod 1-segmented, with 1

lateral and 2 distal setae. Maxillule with 2 large comb-like

spines and 3 tubular setae on coxal endite; basal endite with 3

plumose setae, 1 spine and 1 tubular seta. Maxilla with

tubular setae on coxal endites, allobasis and endopod; prae-

coxal endite with 2 pinnate spines. Maxilliped subchelate with

1 large and 1 small seta (both pinnate) on syncoxa; basis with

long plumose seta on palmar margin, endopodal claw with 1

accessory seta. Pl exopod 3-segmented, exp-3 with distal

outer spine longer than middle outer spine; endopod at least

as long as exopod, 2-segmented, enp-2 longer than enp-1,

inner seta implanted medially. P2—P4 intercoxal sclerites

without ornamentation; rami 3-segmented; exp-1 with an

inner seta. P2 endopod distinctly longer than exopod; enp-1

with outer distal margin attenuated in both sexes; enp-2 with

1 inner margin seta and 1 seta implanted on posterior surface.

Inner distal seta enp-3 P3—P4 extremely reduced and repre-

sented by setule. Armature formula of P1—P4 as follows:

Exopod Endopod

Pl 0.1.023 L2H

P2 Laat 23 Meeal

P3 1-223 sleale 37211

P4 17223 1.1.221

Female fifth pair of legs not fused medially; exopod and

baseoendopod fused to form a bilobate plate; exopodal lobe

with 4-5 setae, endopodal lobe with S setae.

Male with sexual dimorphism in antennule, P2 endopod,

P3 endopod, P5, P6 and in genital segmentation. Antennule

9-segmented, subchirocer; segment 6 swollen, with aes-

thetasc. P2 enp-2 with long outer apophysis not reaching to

distal margin of enp-2; enp-3 with outer spine transformed

into non-articulating process, distal setae reduced and inner

setae enlarged compared to the female. P3 enp-2 with outer

distal corner attenuated into a recurved apophysis. Fifth pair

of legs not fused medially; endopodal lobe with 2 spines,

exopod with 4 setae/spines. Sixth legs symmetrical, fused to

somite, with 2 setae each.

TYPE SPECIES. P. longisetosa Sars, 1910 (by monotypy).

OTHER SPECIES. P. longipes Becker, 1974.

Gee (1988a) concurred with Wells’ (1967) opinion that a

generic distinction between Danielssenia and Psammis on the

base of P5 segmentation alone can hardly be justified.

R. HUYS AND J.M. GEE

| EST \ M@ \\ VXs

Nf a . S

rior (small seta on syncoxa arrowed)

d); G, P4 endopod

ndopod (small seta arrowe

bey one

al segments, posterio

, middle and dist:

. Female: A, mandible, gnathobase; B, mandible, palp; C, maxilliped, ante

‘oie pba : :

Fig. 21 Psammis longisetosa

D, P2 endopod, anterior; E, P2 endo

Male:

. |

owed).

dopo r

d).

REVISION OF DANIELSSENIA AND PSAMMIS

> S—S

“5

ee eee Ee =

ig. 22 Psammis longisetosa. Male: A, antennule and rostrum (armature omitted); F, P5, anterior; G, P6. Female: B, labrum, anterior

C, genital apertures and copulatory pore; D, genital double-somite, ventral; E, PS, anterior.

However, Gee also pointed out that the mandibular gna-

thobase in all Psammis species bears long, relatively fine,

sharply pointed and widely separated teeth compared to the

species of Danielssenia where these teeth are short, stout,

blunt and closely set. On the base of this difference he

suggested that both genera probably utilize different food

items and to a certain extent are trophically isolated. In

combination with the fused rami in the female PS, this

evidence was considered as sufficient to maintain Psammis’

separate generic status. Close examination of the mouthparts

in P. longisetosa and P. longipes and comparison with D.

typica has now revealed several other characters that can be

used to distinguish both genera. Unique features for Psammis

are the specialized comb-like spines on the coxal endite of the

maxillule, the presence of only two spines on the praecoxal

endite of the maxilla, and the extremely enlarged, spinulose

seta on the maxillipedal basis. The presence of tubular setae

and modified spines with tubular extensions on the maxillule

and maxilla is a character that is shared by both genera

though the precise number is not identical. It is conceivable

that these specialized structures might perform a sensory role

(as chemo- or probably mechanoreceptors) in remote food

detection and/or manipulation. Both genera are predomi-

nantly found in the upper flocculent layer of muddy sub-

strates where selection of food-particles probably requires a

different mechanism. This could be particularly true for

deepwater bottoms (fjords, abyss) where either turbidity is

high or the proportion of suspended food-particles might fall

below a subsistence level. The unique specialization of the

mandibles, maxillules and maxillae might be viewed collec-

tively as the result of a different dietary discrimination

mechanism based on successful remote selection of food

particles and thus avoiding the unnecessary high energy costs

of rejecting unsuitable items upon initial capture. It is noted

here that the claviform aesthetascs found on the mouthparts

of certain other Paranannopidae (Gee & Huys, 1991) are not

homologous to the tubular setae or modified spines bearing

tubular extensions.

Another unique apomorphy of Psammis is illustrated by

the setation pattern on the endopods of P3 and P4 (Fig. 23).

The ancestral condition of P3 enp-3 is shown by e.g. Archise-

nia and consists of 1 outer spine (a), 2 distal spines (b—c) and

3 inner setae (d-f). This full complement of armature ele-

ments is also found in Psammis but is obscured by modifica-

tions in the distal part of the segment. The extreme reduction

of the inner terminal spine (c) and the distad displacement of

the distal inner seta (d) are the main reasons why the setal

formula was erroneously cited as 221 (or 121 in P4) in

previous descriptions. The distal elements expressed in this

formula are b and d, rather than b and c. The spiniform and

pinnate nature of seta d in Psammis did certainly contribute

to this misunderstanding. The reduced condition in Bathy-

psammis (Fig. 23) has not evolved from the Psammis pattern

but resulted through the loss of 2 inner setae. It is impossible

to determine which seta (d, e or f) has been retained in B.

longifurca.

Both species of Psammis can be differentiated by the

number of setae on the mandibular basis (3 in longipes, 4 in

longisetosa), the length of the anterior seta on the syncoxa

which is distinctly longer in P. longipes, the ratio of endopod

length to exopod length in P1 to P3 being much higher in P.

longipes, the number of setae on the 9 P5 exopod (4 in

R. HUYS AND J.M. GEE

longisetosa, 5 in longipes), and the gross difference in body

size (+ 550 um in longisetosa, + 900 um in longipes).

DISCUSSION

Within the Paranannopidae, aesthetascs on the mouthparts

are a powerful synapomorphy for separating a number of

genera which have recently been created or redefined, viz.

Jonesiella (cf. Huys & Gee, 1992), Paradanielssenia, Microp-

sammis, Telopsammis and Leptotachidia (cf. Gee & Huys,

1991), Sentiropsis and Peltisenia (Huys & Gee, in press). The

absence of such sensory appendages in Archisenia excludes it

from this lineage and allies it with the more primitive

danielsseniid genera, namely Fladenia, Danielssenia, Psam-

mis and Bathypsammis. However, the phylogenetic relation-

ships amongst these more primitive danielsseniid genera are

somewhat unclear at the moment particularly with respect to

the position occupied by Archisenia. The problem is that this

genus shows a mosaic of primitive plesiomorphic characters

(6-segmented female antennule; setal formula of legs P2—P4

with 7.8.8 setae/spines on exp-3 and 5.6.5 setae on enp-3; PS

with 5 setae on baseoendopod and exopod), but at the same

time a number of unique autapomorphies in the sexual

dimorphism on P1 basis, P2 enp-1 and P3 enp-2.

Within this group of genera it is clear that Fladenia is the

most primitive genus because it retains both vestiges of sexual

dimorphism involving a difference in the number of elements

(in this case setae) on the endopod of P3 and P4 (Gee &

Huys, 1990) and a primitive setal formula particularly in the

exopods of P3 and P4. It is also clear that Danielssenia,

Psammis and Bathypsammis are linked by a 4-segmented

female antennule, a reduced number of setae on P4 enp-3 and

probably by having only 2 setae on the P6 in the male (though

the latter character cannot be scored for Bathypsammis since

the male is unknown). Since it has no vestige of P3 and P4

setal sexual dimorphism and does not show the apomorphies

of the Danielssenia lineage, it is likely that Archisenia

diverged from the main evolutionary line after Fladenia and

probably before the Danielssenia-grouping.

Within the Danielssenia-Psammis-Bathypsammis lineage,

Danielssenia is considered the most advanced genus on

account of the loss of a seta on exp-1 of the antenna, the basis

of the mandible, exp-1 of P2-P4 and enp-2 of P2. Unique

apomorphies for this genus are the typically ventrally

deflected rostrum, the blunt teeth on the mandibular gna-

thobase, and the dorsal, incised, hyaline frill on the

P5-bearing somite. Another diagnostic character for Daniels-

senia is illustrated by the shape of the seminal receptacle.

Multi-chambered receptacles have been described for a num-

ber of Paranannopidae such as Leptotachidia, Telopsammis,

Psammis and Paranannopus (Gee & Huys, 1990, 1990) and

might well be the ancestral state in this family. However, in

none of these genera the paired anterior chambers are ©

elongate, cylindrical reservoirs extending into the posterior

part of the P5-bearing somite.

Analysis of the precise relationships within the Danielsse-

nia grouping is hampered by the absence of male Bathypsam-

mis. The specialized tubular structures on the endites of the

maxillule and maxilla provides a robust synapomorphy to link |

Danielssenia and Psammis. A close relationship is also indi-

cated by the armature of the female sixth legs bearing one

REVISION OF DANIELSSENIA AND PSAMMIS

(

LNeone:

a a

Archisenia Psammis Bathypsammis

Fig. 23. Comparison of armature on distal endopod segment of P3 in Archisenia, Psammis and Bathypsammis.

plumose seta and 2 inner, minute spiniform elements (com-

pared to 2 setae and 1 setule in between in Bathypsammis),

and by a detailed comparison of the distal transformations in

the male P2 endopod. Potential synapomorphies grouping

Psammis and Bathypsammis are: (i) rostrum with enlarged

anterior sensillae; (ii) the mandibular exopod with only 1

lateral and 2 apical setae; (iii) the fusion of the exopod and

baseoendopod in the female P5. Some species of Danielsse-

nia, however, also show a reduction in the setation of the

mandibular exopod (e.g. D. typica), and the fused PS in

Bathypsammis might have been evolved convergently, since,

in other respects, it is very different from the condition in

Psammis. The rostral character might also be a product of

convergence since the enlargement of the anterior pair of

sensillae has evolved independently in a number of other

deepwater genera such as Paranannopus and Cylindronanno-

pus.

Unique apomorphies for Psammis are: (i) reduction of the

mandibular endopod (1 lateral, 3 apical setae); (ii) the

specialized comb-like spines on the maxillulary coxal endite;

(iii) praecoxal endite of maxilla with only 2 spines; (iv)

extreme development of the posterior seta on the maxillipe-

dal basis; (v) elongation of P2 endopod, being longer than the

exopod; (vi) the apophysis on P2 enp-1 in both sexes; (vii)

reduction of the inner terminal seta on P3—P4 enp-3. In

Bathypsammis the unique apomorphies are confined to the

female as the male is unknown: (i) a very long outer basal

seta on the basis of P1; (ii) a very long caudal ramus with a

plume of setules on the inner distal corner; (iii) the form of

the setae on the endopodal lobe of the female P5.

KEY TO GENERA OF PARANANNOPIDAE

REMARK. This key also includes Psammis kliei Smirnov,

1946, which will be placed in a genus by itself in a forthcom-

ing paper (Gee & Huys, in prep.), and the genus Carolinicola

Huys & Thistle, provisionally assigned to the Paranannop-

idae by Huys & Thistle (1989).

1. P4 endopod 3-segmented

P4 endopod 2-segmented, 1-segmented or absent

2. Antennary exopod 1-segmented

Carolinicola Huys & Thistle, 1989.

Antennary exopod 3-segmented

3. Body short, robust; caudal rami setae [TV and V long and

spinulose; PS well developed, covering entire width of thoracic

somite Paranannopus Lang, 1936.

Body slender, cylindrical to vermiform; caudal rami setae IV

and V short and plumose; P5 a minute plate, located midven-

BLAU Y cee hivisoweceneenecnccteceencss Cylindronannopus Coull, 1973.

4. P2-P4 exp-1 without inner seta ..............ceeeeeeceeeee een eeees a

P2—PAlexp-liwithanner seta 95.. seacctacst 3. he, -zarsteepellc «ques the

5. Antennules without plumose or _ pinnate spines/

SCLAG) entries nr avoscnuvspennpienanesas Sentiropsis Huys & Gee, 1993.

Antennules with plumose and/or pinnate spines/setae ....... 6.

6. Caudal ramus with distinct cluster of long setules at the inner

distal corner; P2 enp-2 with large apophysis in 2 (and presum-

ably in & also) Psammis kliei Smirnov, 1946.

Caudal ramus without such cluster; P2 enp-2 with large apophy-

sis in O’ only Danielssenia Boeck, 1872.

ee Aexp-3) With OISCtae/ SPINES! cese.s- caress ee-csecceneeesereaceeaeene 8.

P4 exp-3 with at most 7 setae/spines .................ceeeeeeee ees 13:

Someb2.enp-2: witht INnemSetacy ce -ssss.cceseracs cs uecaveeueoseaesceete oO:

P2ienp-2iwithylamner|setal -fse28, eee sae ee eeenea eerie ee 10.

9. Caudal rami 5 times as long as maximum width; P1 endopod

shorter than exopod; P5 9 with fused exopod and baseoen-

COPOd are sac ee erie scue ee wee eReaes Bathypsammis gen. nov.

Caudal rami broader than long; P1 endopod longer than exo-

pod; P5 9 with separated exopod and baseoendopod ............

eR, S8) 8 REE. Rae ese teee Jonesiella Brady, 1880.

10. Body dorsoventrally flattened; caudal rami setae IV and V

stubby and spiniform; Pl enp-1 1.5 times as long asenp-

DP SEES DO ee SE FOR 3 Peltisenia Huys & Gee, 1993.

Body not dorsoventrally flattened; caudal rami setae [TV and V

long and setiform; P1 enp-1 at most as long as enp-2 ....... sil

11. Antennule 9 4-segmented; club-shaped aesthetascs present on

mandible (endopod), maxillule (basis) and maxilla (endopod);

P2 enp-2 CO without distinct outer apophysis

smacgran etepmaead serps = beasts ct tee Paradanielssenia Soyer, 1970.

Antennule @ 6-segmented; no club-shaped aesthetascs on

mouthparths; P2 enp-2 C& with long outer apophysis ....... 12?

12. Antennary exopod with 1 seta on proximal segment; P3 exp-3

with 7 setae/spines; P2 enp-3 with inner distal seta transformed

into large pinnate spine reaching beyond apophysis of enp-

Di ES. RISO A oie USI eames net ane Afrosenia Huys & Gee, 1993.

Antennary exopod with 2 setae on proximal segment; P3 exp-3

with 8 setae/spines; P2 enp-3 OC’ with inner distal seta not

transformed and shorter than apophysis of enp-2

gap leche Guise Bilstagsjculsoe aelsactue sh Soh ewsislte agaticamatas Archisenia gen. nov.

13. P2 enp-2 with 2 inner setae

LA ayoall yd (S11 RENEE coococcansanr acoonpocedoqaccacoacdect 14.

14. Club-shaped aesthetascs present on mandible (endopod), max-

illule (basis) and maxilla (endopod); P2 exp-3 with at most 6

SCtAC/SPIMES oja.eccrenctagaais 9 Passe ninasiasie-(asieelalacaee desacbe yaseoeossercme 15.

No club-shaped aesthetascs present on these appendages; P2

exp-3 with 7 setae/spines .......... Fladenia Gee & Huys, 1990.

15. P1 enp-2 with 2 terminal setae geniculate; PS 2 baseoendopod

and exopod indistinguishable, with 5 setae; P2 enp-2 CO’ without

apophiysiswPOIG@), withi2 Setaew..-.c-ce-e-ce-eeeee-eeeeeseeece ease 16.

P1 enp-2 with 1 terminal seta geniculate; PS 2 baseoendopodal

and exopodal lobes indistinguishable, with 3 and 4 setae,

respectively; P2 enp-2 O' with small apophysis; P6 O’ with 3

SELAC east Waesceteenane eS eS eanetos Micropsammis Mielke, 1975.

16. Antennule in both sexes with densely opaque, bulbous append-

age on distal segment P2—P4 exp-2 without inner seta ............

reise ssi Cisse is ceisiie = eagle ecldseee Stee ates Leptotachidia Becker, 1974.

Antennule in both sexes without densely opaque, bulbous

appendage on distal segment P2—P4 exp-2 with inner seta

Pash A sumed «scear ens demeceueen cess Telopsammis Gee & Huys, 1991.

ACKNOWLEDGEMENTS. The authors wish to thank Dr. Philippe Bodin

and the Curators of Crustacea at the Zoologisches Museum der

Universitat Kiel, the Naturhistoriska Riksmuseet Stockholm, the

Zoologisk Museum Oslo and the National Museum of Natural

History (Smithsonian Institution) for the loan of material. Dr Geof-

frey A. Boxshall is acknowledged for his comments on an earlier

draft of the ms. For the senior author this is communication No. 551

of the Centre for Estuarine and Coastal Ecology, Yerseke and for the

junior author the work forms part of the Community Ecology

Programme of the Plymouth Marine Laboratory.

R. HUYS AND J.M. GEE

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Bull. nat. Hist. Mus. (Zool.) 59(1): 83-94

Issued 24 June 1993

A new species of Syrticola Willems & Claeys,

1982 (Copepoda: Harpacticoida) from Japan

with notes on the type species

RONY HUYS

Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD

SUSUMU OHTSUKA

Fisheries Laboratory, Hiroshima University, 1294 Takehara-cho, Takehara, Hiroshima 725, Japan

CONTENTS

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Synopsis. A new species of Syrticola Willems & Claeys, 1982 (Harpacticoida: Cylindropsyllidae) is described from

Okinawa, Japan. Morphological notes on the type species S. flandricus Willems & Claeys, 1982 and a key to the

species are given. The inadequately described S. trispinosus A. Scott, 1896 is ranked as species inquirenda. The

diagnosis of the genus is amended and its position in the Cylindropsyllidae re-assessed. Both sexes of S. intermedius

sp. nov. were found to be infested by early parthenogenetic female stages of an as yet undescribed genus of

Tantulocarida.

INTRODUCTION

The interstitial harpacticoid fauna of Japan is very poorly

known, as is that of most east Asian countries. The paucity of

data on marine interstitial species stands in marked contrast

with the number of studies on subterranean copepods pro-

duced by workers like Miura and Takashi Ito. In fact, with

the possible exception of Microsetella norvegica (Boeck,

| 1864) only 11 genuinely interstitial harpacticoids have been

recorded from marine and brackish water habitats in Japan

(Table 1) and the majority of these was described by the

latter author’s namesake, the late Tatsunori It6, whose

activities were mainly focussed on the fauna from Hokkaido

in the north and the Bonin Islands in the southeast. The only

other information on mesopsammic harpacticoids is con-

tained in the papers of Kikuchi (1970, 1972) and Kikuchi &

Yokota (1984), reporting on species from Lake Hinuma, a

brackish lagoon near the central east coast of the Japanese

mainland.

In the course of a survey of the sandy bottom copepods off

Nagannu Island, Okinawa (Ryukyu Archipelago) by one of

us (S.O.), several interstitial harpacticoids were found to be

infested with tantulocaridans (Huys et al., in preparation).

This paper describes a new species of Syrticola Willems &

Claeys (Cylindropsyllidae) based on two specimens that were

parasitized by parthenogenetic females of an as yet unde-

scribed tantulocarid.

MATERIALS AND METHODS

Specimens of Syrticola intermedius sp. nov. were collected by

dredging of a sandy bottom off Nagannu Island, Okinawa,

South Japan (26° 14’ N, 127° 32’ E, depth 46.49 m; leg. S.

Ohtsuka) on 9 April 1992. The dredge (mouth area: 50 cm

wide x 15 cm high; mesh size 5 mm) was towed along the

bottom at a speed of 2 knots by the T/V Toyoshio-maru of the

Hiroshima University for about 5 minutes. Copepods were

R. HUYS AND S. OHTSUKA

Table 1 Interstitial harpacticoid copepods reported from marine localities in Japan.

Species

ECTINOSOMATIDAE

Microsetella norvegica (Boeck, 1864)

Arenosetella bidenta Ito, 1972

Noodtiella sp.

DARCYTHOMPSONIIDAE

Leptocaris brevicornis (van Douwe, 1904)

PARAMESOCHRIDAE

Paramesochra sp.

LEPTASTACIDAE

Cerconeotes japonicus (It6, 1968)

Paraleptastacus unisetosus Ito, 1972

CYLINDROPSYLLIDAE

Arenopontia ishikariana It6, 1968

Arenopontia sakagamii It6, 1978

Stenocaris intermedia Ito, 1972

Psammopsyllus imamurai Kikuchi, 1972

PARASTENOCARIDIDAE

Parastenocaris hinumaensis Kikuchi, 1970

' Brackish lagoon.

fixed and preserved in 10% neutralized formalin/sea-water.

Females of S. flandricus Willems & Claeys, 1982 were

collected by the senior author in different localities along the

coast of The Netherlands in the course of the biological

monitoring programme BIOMON. All specimens have been

deposited in the collections of The Natural History Museum,

London.

Specimens were dissected in lactic acid and the dissected

parts were placed in lactophenol mounting medium. Prepara-

tions were sealed with glyceel (Gurr®, BDH Chemicals Ltd,

Poole, England). All drawings have been prepared using a

camera lucida on a Leitz Diaplan differential interference

contrast microscope. The descriptive terminology is adopted

from Huys & Boxshall (1991). Abbreviations used in the text

are: P1—P6, first to sixth thoracopod.

DESCRIPTIONS

Family Cylindropsyllidae

Subfamily Leptopontiinae Lang, 1948

The genus Syrticola was established by Willems & Claeys

(1982) to accommodate the type species S. flandricus Willems

& Claeys, 1982 and Tetragoniceps trispinosus A. Scott, 1896.

Previously, the latter species had been considered ‘species

incerta in the genus Evansula T. Scott and thus placed in the

subfamily Cylindropsyllinae (Lang, 1948). The close relation-

ship between Syrticola and Notopontia Bodiou noted by

Willems & Claeys (1982) was already hinted at by Bodiou

(1977) who recognized a certain resemblance between T.

trispinosus and N. stephaniae Bodiou, 1977, and indirectly

also by Mielke (1982) who described (?) N. galapagoensis, a

Locality Reference

Hokkaido Itd (1968)

Hokkaido Itd (1972, 1984)

Hokkaido Itd (1984)

Lake Hinuma! Kikuchi & Yokota

(1984)

Hokkaido Itd (1984)

Hokkaido Itd (1968, 1984)

Hokkaido Itd (1972, 1984)

Hokkaido Itd (1968, 1984)

Bonin Islands Itd (1978)

Hokkaido Itd (1972)

Lake Hinuma! Kikuchi (1972)

Lake Hinuma! Kikuchi (1970)

species provisionally placed in Notopontia but subsequently

allocated to Syrticola (Bodiou & Colomines, 1986; Willems et

al., 1987). However, none of these authors has formally

assigned either of these genera to any of the subfamilies of

the Cylindropsyllidae recognized at that time. The only

attempt was that by Bodiou (1977) who suggested that

Notopontia is closest to Evansula (Cylindropsyllinae) but to a

certain extent is also related to Arenopontia Kunz and

Leptopontia T. Scott (Leptopontiinae).

Lang (1948) subdivided the family into the Cylindropsylli-

nae, Leptastacinae and Leptopontiinae and a fourth subfam-

ily, the Psammopsyllinae, was added by Krishnaswamy —

(1956). Recently, the Leptastacinae has been upgraded to full

family status (Huys, 1993). The diagnostic sexual dimorphism |

displayed on thoracopods 2 and 3 by all genera of the |

Cylindropsyllinae excludes Notopontia and Syrticola from

this subfamily since their swimming leg sexual dimorphism is

only slightly developed (and therefore might well have been

overlooked in Notopontia for which it has been recorded as

being completely absent). A detailed comparison with the

Leptopontiinae, currently encompassing Arenopontia,

Pararenopontia Bodiou & Colomines and Leptopontia, |

reveals a suite of apomorphic characters supporting a sister- |

group relationship between Leptopontia and the Notopontia-

Syrticola lineage. These characters include: (i) anal

operculum drawn out into spinous process(es); (11) outer

distal corner of caudal ramus produced into backwardly

directed spinous process; (ili) first antennulary segment

extremely elongated, much longer than second; (iv) mandib-

ular gnathobase stylet-like with teeth along one side; (v)

distal exopod segment P1 with 3 armature elements (proximal

outer spine lost); (vi) middle exopod segment P1 without

outer spine (in Syrticola and Notopontia the middle and distal

segment are fused or have failed to separate); (vii) apical

spines of distal exopod segments P3-P4 setiform; (viii) sexual :

NEW SPECIES OF SYRTICOLA FROM JAPAN

dimorphism endopod P3 involving fusion of distal spine to

segment; (ix) PS exopod with 3 elements in both sexes. There

is little evidence that Arenopontia and Pararenopontia share a

close relationship with this core group, however pending a

revision of these genera it is preferable to retain them in the

Leptopontiinae.

Syrticola Willems & Claeys, 1982

DIAGNOSIS (AMENDED). Leptopontiinae. Body cylindrical,

but not particularly vermiform. Hyaline frill of all body

somites incised. Antennule 6- or 7-segmented in 2. Maxilla

with one syncoxal endite. Midventral spinous process ante-

rior to intercoxal sclerite of Pl. Pl exopod 2-segmented.

Distal segment P1 endopod with 1 geniculate seta and 1 claw.

Distal segment P3—P4 exopods with 1 outer spine. P2—P4

endopods 1-segmented in 9, P3 endopod 1- or 2-segmented

and sexually dimorphic in O’. P5 with fused baseoendopod

and exopod in both sexes; endopodal lobe drawn out into

triangular process with 0-1 seta, exopodal lobe a tubercle with

3 elements. Genital apertures not fused in 2. Anal opercu-

lum with a series of small spinous processes or one large

median spike. Caudal ramus seta III inserted proximal to seta

Vv.

TYPE SPECIES. Syrticola flandricus Willems & Claeys, 1982

OTHER SPECIES. S. trispinosus (A. Scott, 1896), S. galapa-

goensis (Mielke, 1982), S. mediterraneus Willems et al. , 1987,

S. intermedius sp. nov.

Syrticola intermedius sp. nov. (Figs. 1-4, 5A-C, 6)

MATERIAL EXAMINED. Holotype Q dissected on 8 slides,

deposited under reg. no. 1992.1075. Paratype C dissected on

6 slides, deposited under reg. no. 1992.1076. Drawings based

on the paratype are Figs. 2E-F, 4D-F, 5A-C, 6A-G; all

others were drawn from the holotype 9°.

FEMALE. Body length measured from tip of rostrum to

posterior margin of caudal rami 485 um (Figs. 1A—B). Maxi-

mum width 75 wm measured at rear margin of cephalothorax.

Integument pitted. Pleural areas of cephalothorax not well

developed so that appendages are clearly exposed in lateral

aspect (Fig. 1B). Posterior margin of body somites (except

cephalothorax and anal somite) fringed dorsally and laterally

with finely incised hyaline frill; this frill also present ventrally

on genital double-somite and abdominal somites (Figs. 1B,

4A-B). Abdominal somites also with transverse spinular row

in anterior half which is usually concealed beneath the

hyaline frill of the preceding somite as shown in Fig. 5A.

Rostrum triangular, with 2 delicate sensillae (as in male,

Fig. 6B).

Genital double-somite (Fig. 4B) about as long as wide;

original segmentation not marked by any external or internal

cuticular structure; anterior margin with 2 transverse spinular

rows. Genital apertures located in anterior quarter of genital

double-somite, closely set together but separate and each

closed off by small operculum derived from sixth leg; no

armature observed but posterior margin of operculum with

minute spinous processes and a circular scar at the outer

distal corner (probably indicating insertion site of long seta as

in S. flandricus, cfr. Fig. 5G). Copulatory pore located far

anteriorly between genital apertures (arrowed in Fig. 4B).

Seminal receptacles not confirmed. Paired widely separated

secretory pores at about 2/5 distance from anterior margin.

Anal somite (Figs. 1A—B; 4A; 5A-B) with dorsal opercu-

lum drawn out into median, posteriorly directed, spike;

process about as long as anal somite proper; ventral posterior

margin spinulose medially.

Caudal rami (Figs. 4A; SA-B) divergent; outer distal

corner drawn out into backwardly directed, acutely recurved,

spinous process; with 7 setae; seta I minute, setae II and III

located anterior to seta I, seta IV tiny and located between

spinous process and large seta V, seta VII long and tn-

articulate at base, seta VI minute.

Antennule (Fig. 2A) 7-segmented, articulating on a small

pedestal as in the male (Fig. 6B); slender, anteriorly directed

(Fig. 1A); first segment extremely elongate, about 4 times as

long as maximum width, with 1 short seta distally; aesthetasc

on fourth segment fused basally to long seta; distal 2 setae of

last segment fused basally. All setae bare; setal formula: [1,

Ssperae, 1, 2,9].

Antenna (Figs. 2B—D). Coxa small, not ornamented. Basis

and proximal endopod segment fused to form allobasis,

original segmentation marked by internal chitinous rib anteri-

orly and incomplete suture line posteriorly near exopod; basis

with serrate seta located on inner lateral surface; exopod

small, 1-segmented, with 1 small, apical seta; free endopod

articulating with allobasis at right angle (Fig. 1B), lateral

margin with 2 spines, distal margin with 1 pinnate spine and 4

geniculate setae, the largest of which is fused basally with

vestigial seta and bearing coarse spinule at about midway.

Labrum (Fig. 5C) a ventrally projected, elongate, membra-

nous outgrowth, distinctly tapering distally. Paragnaths small

membranous lobes.

Mandible (Figs. 2E-F) with conspicuous coxa, drawn out

to form a slender, stylet-like gnathobase bearing small teeth

and a long serrate seta near the apex. Palp elongate,

2-segmented; proximal segment representing basis, slightly

sigmoid, swollen in distal half, with 1 seta and spinular row;

distal segment representing endopod, with 2 lateral and 3

apical setae.

Maxillule (Fig. 2G). Praecoxa with large, cylindrical arth-

rite bearing 2 anterior surface setae and 6 setae along the

distal margin; coxal endite with 2 setae; palp representing

fused basis and rami; exopod, endopod, proximal and distal

basal endites represented by 1, 2, 2 and 3 setae, respectively.

Maxilla (Fig. 2H) reduced, 2-segmented. Syncoxa with

single endite bearing unipinnate seta and conspicuous

aesthetasc-like structure representing modified seta with chi-

tinized dorsal margin and tubular membranous part ventrally;

exit of maxillary gland discernible in proximal half. Allobasis

drawn out into pinnate claw bearing serrate seta at its base.

No trace of endopod.

Maxilliped (Fig. 4F) subchelate. Syncoxa and basis without

armature but with 3 spinular rows each. Endopod repre-

sented by strong claw bearing tiny spinules along distal half of

inner margin; an accessory setule is located at the base of the

claw.

P1 (Fig. 3A). Praecoxa a small sclerite located around the

outer lateral margin of the limb base. Intercoxal sclerite a

minute rounded plate. Coxa with spinular row. Basis with

inner and outer basal seta and with spinules at middle distal

margin. Exopod 2-segmented, proximal segment with blunt

spine bearing long setules, distal segment with 2 geniculate

setae and 1 unipinnate spine. Endopod 2-segmented, elon-

gate, prehensile; proximal segment about twice as long as

exopod, with serrate inner seta near proximal margin; distal

segment with 1 geniculate seta, 1 short claw and a patch of

NEW SPECIES OF SYRTICOLA FROM JAPAN

latera

ry endopod, outer

inner lateral view; C, distal end of antenna

lateral view; G, maxillule; H, maxilla. Male. E, mandible; F, Mandible, gnathobase.

. female. A, Antennule; B, antenna,

ola intermedius sp. nov

ntennary exopod, outer

g.2 Syrtic

view; D, a

88 R. HUYS AND S. OHTSUKA

Fig. 3 Syrticola intermedius sp. nov. female. A, P1, anterior; B, P2, anterior; C, P3, anterior; D, P4, anterior.

NEW SPECIES OF SYRTICOLA FROM JAPAN

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g.4 Syrticola intermedius sp. nov. female. A, Urosome, ventral; B, genital double-somite, ventral; C, P5, anterior. Male. D, P5, anterior;

E, sixth pair of legs; F, maxilliped. [Arrows in C-E indicating vestigial seta; copulatory pore arrowed in B.]

90 R. HUYS AND S. OHTSUKA

Fig. 5 Syrticola intermedius sp. nov. male. A, Anal somite and left caudal ramus, dorsal; B, anal somite and right caudal ramus, ventral; C,

labrum, anterior. Female of undescribed tantulocaridan. D, Cephalic shield, dorsal; E, same, lateral. Syrticola flandricus female. F, P5,

anterior; G, genital double-somite, ventral; H, anal operculum and left caudal ramus, dorsal.

ig.6 Syrticola intermedius sp. nov. male. A, Habitus, dorsal; B, rostrum and antennule, dorsal; C, antennule, segments 3 and 4, anterior

[armature of these segments omitted; segment 4 stippled]; D, antennulary segment 3, anterior; E, antennulary segment 4, anterior; F,

proximal part of antennulary segment 5, anterior; G, P3 endopod, anterior.

fine spinules. A distinct, ventrally directed, spinous process is

located at the ventral midline between the maxillipedal

syncoxae and the coxae of the first leg (Fig. 3A).

P2-P4 (Figs. 3B-D) with 3-segmented exopods and

l-segmented endopods. Intercoxal sclerites small, rectangu-

lar, bare (Fig. 3C). Spines of distal exopodal segment elon-

gate and slender in P3 and P4. Inner seta of P2 endopod

serrate and typically recurved (Fig. 3B). Inner margin of

endopod P3 with serrate seta and vestigial seta represented

by setule (see inset Fig. 3C). Distal spines of endopod pinnate

in P2—P3, bare in P4. Armature formula as follows:

coxa basis exopodsegment endopod segment

1 1 Z

Pil 0-0 1-1 1-0; 1,2,0 0-15 OLE

P2 0-0 1-0 1-0; 1-0; 1, 11,0 0,11,1

P3 0-0 1-0 1-0; 1-0; Hele 0,1,2

P4 0-0 1-0 1-0; 1-0; [1,1 0,11

Fifth legs (Figs. 4A, C) closely set together, no intercoxal

sclerite. Baseoendopod and exopod fused to form a single

plate with 2 secretory pores and 4 armature elements in total;

endopodal lobe represented by long, triangular, spinous

process without setae but with tiny spinules along proximal

outer margin and on posterior surface; exopod presumably

represented by weakly developed process bearing outer pin-

nate spine, inner slender seta and a vestigial seta in between.

Outer basal seta elongate and bare.

MALE. Body length measured from tip of rostrum to poste-

rior margin of caudal rami (Fig. 6A) 460 um. Ornamentation

of body somites generally as in female; genital and first

abdominal somites separate, with spinulose hyaline frill each.

Sexual dimorphism in antennule, P3 endopod, PS, P6 and in

genital segmentation. Spermatophore not observed.

Antennule (Figs. 6B—F) indistinctly 8-segmented, articulat-

ing on a small pedestal. Relative lengths of first two segments

as in female. Third and fourth segment (= ancestral segment

XIII) interdigitating as shown in Fig. 6C. Major geniculation

between segments 6 and 7. Segmental fusion pattern: I,

II-VI, (X-XII, XII, XIV-XVIII, XIX-XX, XXI-XXII,

XXITI-XVII. Segment 6 with 1 modified flat spine and 1

setule, segment 7 with similar spine and 1 stubby pinnate

element. Armature formula: [1, 9, 5, 2, 4+ae, 2, 2, 9].

P3 endopod (Fig. 6G) 2-segmented. Proximal segment

unarmed. Distal segment drawn out into pinnate process

(derived from distal spine in 2) with spatulate tip bearing 2

rows of denticles; inner margin with short pinnate seta and

minute setule.

P5 (Fig. 4D). Relative position, shape and armature largely

similar to female except for the inner exopodal and outer

basal seta being distinctly shorter. Ornamentation of endopo-

dal lobe also slightly different with fewer spinules along the

proximal outer margin and tiny spinules along the inner

margin.

Sixth pair of legs (Fig. 4E) positioned midventrally, sym-

metrical; inner distal corner with numerous minute spinules

and produced into a small process; armature consisting of

inner strong spine, outer slender seta and a vestigial setule in

between.

VARIABILITY. An aberrant left P3 was noticed in the holo-

type @ (Fig. 3C).

R. HUYS AND S. OHTSUKA

ETYMOLOGY. The species name is derived from the Latin

inter, meaning between, and medius, meaning middle, and

refers to the intermediate position between S. galapagoensis

and the European species of the genus.

Syrticola flandricus Willems & Claeys, 1982 (Figs.

5F-H)

MATERIAL EXAMINED. 3 99 from off Walcheren, The Neth-

erlands, southern North Sea, 51° 57'25” N, 02° 40'45” E,

depth 44.5 m, coarse sandy sediment, 08 May 1991, coll. R.

Huys. One Q in alcohol deposited under reg. no. 1992.1077.

The description given by Willems & Claeys (1982) is

detailed and therefore only a few corrections to the original

figures are noted here.

Antenna. The exopod possesses only one seta as in S.

intermedius and S. galapagoensis. The oblique suture line has

probably been mistaken for the lateral seta (compare Fig. 2D

with Fig. 2B in Willems & Claeys (1982)), and it is conceiv- |

able that the same misinterpretation applies for S. mediterra- _

neus (cf. Willems et al., 1987: Fig. 3A).

Mandible. The basis bears only one seta; the supernumer- _

ary proximal ‘setae’ figured by Willems & Claeys are part ofa

transverse row of long spinules running around the lateral

margin of the basis.

Maxillule. The arthrite of the praecoxa has 6 marginal and

2 surface setae, the coxal endite 2 setae and the distribution

pattern of the palp setae is identical to S. intermedius (Fig.

2G).

Maxilliped. The endopodal claw bears an accessory setule

at its base.

P1. A seta is located at the inner distal corner of the basis.

P5. The armature of the exopodal lobe consists of an outer

spine, an inner seta and a setule in between (Fig. 5F).

The genital field is basically the same as in S. intermedius

(Fig. 5G).

REMARKS

A single probably parthenogenetic female of a tantulocaridan

was found attached to the pleurotergite of the P3-bearing

somite of the holotype 2 of S. intermedius (Fig. 1A). The

specimen is about 160 um long and is at an early stage of

development. The larval postcephalic trunk had been

sloughed already but no differentiating tissue could be

observed inside the sac. The male paratype was also infested

by a parthenogenetic female (Fig. 6A) which was larger

(235 um) and attached to the pleurotergite of the genital

somite. Inside the sac a large number of small eggs of about

20-25 um in diameter is contained. Both tantulocaridan

stages most likely belong to an as yet undescribed species;

which was found to infest harpacticoids belonging to at least

two other families (Huys et al., in preparation). Since only)

the head shield (Figs. 5D-E) is left for comparison this)

identification has to be considered provisional.

NEW SPECIES OF SYRTICOLA FROM JAPAN

DISCUSSION

Syrticola intermedius is the second species to be reported

from the Indo-Pacific, the other species (under the name (?)

_Notopontia galapagoensis) being originally described from a

sandy beach in the Galapagos (Mielke, 1982). Both species

resemble each other morphologically. A comparison of the

major diagnostic characters (Table 2) reveals two species

groups in the genus Syrticola. The European group includes

S. trispinosus, S. flandricus and S. mediterraneus and is

characterized by a 6-segmented antennule (segments 6 and 7

fused) and the anal operculum possessing small spinous

processes (Fig. 5H). The number of these projections ranges

from (rarely) 0 to 5, though specimens with a single small

process have not been recorded yet (Willems et al., 1987).

The second species group encompasses the two Indo-Pacific

species which share a 7-segmented antennule and an opercu-

lum drawn out into a single median strong spike. Both species

also share the plesiomorphic 2-segmented condition of the

male P3 endopod, but the significance of this character is

limited since not all the males are known in the European

species group. The zoogeographical and morphological sepa-

ration does not warrant the upgrading of these groupings to

generic rank, however, since S. intermedius exhibits certain

characters found in the European species. Outgroup compar-

ison with Notopontia and Leptopontia suggests that the

spiniform nature of the outer exopodal spine and the loss of

the inner baseoendopodal seta are apomorphic character

states for the fifth legs, linking the Japanese species with its

European congeners. The outline of the anal operculum links

S. intermedius to (?) N. galapagoensis, justifying the latter’s

re-allocation to Syrticola by Willems et al. (1987).

The possession of an aesthetasc-like structure on the syn-

coxa of the maxilla in S. intermedius is unusual.

Re-examination of S. flandricus showed an unmodified seta

in this position, in addition to the pinnate one also present in

S. intermedius. Two setae are also reported on the syncoxal

endite of S. mediterraneus and in the outgroup taxa Notopon-

tia and Leptopontia. The report of 3 setae on this endite in S.

alapagoensis (Mielke, 1982) therefore probably results from

N misinterpretation of an aesthetasc-like structure. Without

ifferential interference contrast microscopy the flaccid distal

art is easily overlooked, thereby accentuating the lateral

hitinized margins as setoid structures. The relative lengths of

he enditic ‘setae’ in Mielke’s (1982: Abb. 18E) illustration

re suggestive of this interpretation.

The status of S. trispinosus remains enigmatic as ever. A.

able 2 Comparison of Syrticola species.

trispinosus

ntennule ? 6-segmented

flandricus

6-segmented

Scott’s species is clearly closely related to S. flandricus.

Willems & Claeys (1982) list a number of differences but

except for the structure of the fifth leg, all of these can be

attributed to deficiencies in the original decription. This,

however, does not rule out S. trispinosus as a distinct species

since the discovery of an as yet undescribed species of

Syrticola in the North Sea has proven species discrimination

in this genus to be rather unreliable. Pending re-examination

of topotypes from the Isle of Man, S. trispinosus is relegated

to species inquirenda.

KEY TO SPECIES

1. Antennule 2 7-segmented, anal operculum with one large,

IC CUI AIS IR CM re acii's 9: Jee vptlaawintls «meine samacisents= 15. see nage Ps

Antennule 2 6-segmented, anal operculum with several small,

SILOS PLOCESSES Was cee citaowtis s seieadetdw arate coamindelss sficlala cheeses 3).

2. P3 endopod without inner seta; outer exopodal element PS

NEMMOUMI ES cee we eects deecee et eee ces galapagoensis (Mielke. 1982).

P3 endopod with inner seta; outer exopodal element PS spini-

LOIRE Ree eee een oot cise cnwn vane ct sacce react intermedius sp. nov.

3. PS5 exopodal lobe with 2 spiniform elements .....................04+

Pee eee ee ce daca Chics vosercewse trispinosus (A. Scott, 1896).

Only one element of PS exopodal lobe spiniform ............. 4.

4. ESIEDGOPOd WIthOULINNEL SETA .5...< cdevec cate westecencencadeene desea

Me eWeomessan cide sisinsieescieaisanogma ss mediterraneus Willems et al., 1987.

ACKNOWLEDGEMENTS. Drs Geoffrey Boxshall (The Natural History

Museum) and Philippe Bodin (Université de Bretagne Occidentale)

are gratefully acknowledged for commenting on earlier drafts of the

manuscript. We also would like to acknowledge the captain and the

crew of the TR/V Toyoshio-maru of the Hiroshima University and

Mr. M. Okada and Mr. Y. Endo for assistance at sea. Part of this

study was supported by research grants of the Research Institute of

Marine Invertebrates (1989) and the Narishige Zoological Science

Award (1992) awarded to one of us (S.O.). R.H. is a visiting research

fellow of the Institute of Zoology, University of Gent, Belgium. This

is communication no. 623 of the Centre for Estuarine and Coastal

Ecology, Yerseke, The Netherlands.

mediterraneus galapagoensis intermedius

6-segmented 7-segmented 7-segmented

_ {nner seta P3 endopod 9 ? present absent absent present

ndopod P3 i @ l-segmented 2-segmented 2-segmented

uter element exopodal lobe PS9C’ spiniform spiniform spiniform setiform spiniform

nner seta endopodal lobe PS 2 absent absent absent present absent

nal operculum processes several, several, several, one, one,

small small small large large

ody length ? (um) 500 460-530 540-660 280-340 485

ody length O’ (um) ? 2 460-500 310-350 460

istribution Irish Sea North Sea Mediterranean Galapagos Japan

REFERENCES

Bodiou, J.-Y. 1977. Harpacticoides (Crustacés, Copépodes) des iles Kerguelen.

III. Description de deux formes nouvelles de la famille des Cylindropsyll-

idae. In: Le benthos du plateau continental des iles Kerguelen. CNFRA 42:

277-286.

—— & Colomines, J.-C. 1986. Harpacticoides (Crustacés, Copépodes) des iles

Crozet. I. Description d’une espéce nouvelle du genre Arenopontia Kunz.

Vie et Milieu 36: 55-64.

Huys, R. 1993. The amphiatlantic distribution of Leptastacus macronyx (T.

Scott, 1892) (Copepoda: Harpacticoida): a paradigm of taxonomic confu-

sion; and, a cladistic approach to the classification of the Leptastacidae Lang,

1948. Academiae Analecta, in press.

—— & Boxshall, G.A. 1991. Copepod Evolution. The Ray Society, London.

468 pp.

It6, T. 1968. Descriptions and records of marine harpacticoid copepods from

Hokkaido. I. Journal of the Faculty of Sciences, Hokkaido University,

Zoology 16: 369-381.

— 1972. Descriptions and records of marine harpacticoid copepods from

Hokkaido. IV. Journal of the Faculty of Sciences, Hokkaido University,

Zoology 18: 305-336.

— 1978. A new species of marine interstitial harpacticoid copepod of the

genus Arenopontia from the Bonin Islands, southern Japan. Annotationes

Zoologicae Japonenses. 51: 47-SS.

— 1984. Studies on the interstitial animals in the Ishikari beach, Hokkaido,

northern Japan — a preliminary report. Benthos Research, Bulletin of the

R. HUYS AND S. OHTSUKA

Japanese Association of Benthology 26: 1-14. {In Japanese with English

summary.]

Kikuchi, Y. 1970. A new species of Parastenocaris (Har-pacticoida) from a

sandy beach of the Lake Hinuma. Annotationes Zoologicae Japonenses 43:

170-173.

— 1972. Psammobiontic harpacticoid copepods of Lake Hinuma, II. Anno-

tationes Zoologicae Japonenses 45: 170-177.

— & Yokota, K. 1984. New records of two freshwater harpacticoid cope-

pods, Nannopus palustris Brady and Leptocaris brevicornis (van Douwe), in

Lake Hinuma. Publications of the Itako hydrobiological Station 1: 1-9.

Krishnaswamy, S. 1956. Sewellina reductus gen. et sp. nov., a new sanddwell-

ing copepod from Madras. Zoologischer Anzeiger 157: 248-250.

Lang, K. 1948. Monographie der Harpacticiden. Hakan Ohlsson, Lund. I:

1-896, figs. 1-361; II: 897-1682, figs. 362-607, maps 1-378.

Mielke, W. 1982. Interstitielle Fauna von Galapagos. XXIX. Darcythompsoni-

idae, Cylindropsyllidae (Harpacticoida). Mikrofauna Meeresbodens 87:

1-52.

Scott, A. 1896. Description of new and rare Copepoda. In: Report on the

Investigations carried on in 1895 in connection with the Lancashire Sea-

Fisheries Laboratory at University College, Liverpool. Proceedings and

Transactions of the Liverpool Biological Society 10: 134-158

Willems, K.A. & Claeys, D. 1982. Syrticola flandricus n. g., 0. sp., a

harpacticoid copepod from the Southern Bight of the North Sea. Crustaceana

43: 1-8.

—, — & Fiers, F. 1987. Syrticola mediterraneus n. sp., a harpacticoid

copepod from the Bay of Calvi, Corsica. Hydrobiologia 153: 71-78.

Bull. Nat. Hist. Mus. (Zool.) 59(1): 95 Issued 24 June 1993

Erratum

The following replaces Fig. 5, Bull. Nat. Hist. Mus. (Zool.) 58(1), p. 42.

7 *

CF.

Nes & i £ a rf ip br Pitan

} id

be." ¢ 2,7

Fig. 5. Pharyngochromis acuticeps (Steind.) Adult male; $.L. 98.0 mm. Okavango system (RUSI 34974). Photographed by Paul Skelton and

R. Stobbs.

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CONTENTS

1 Anew snake from St Lucia, West Indies

G. Underwood

11. Anatomy of the Melanonidae (Teleostei: Gadiformes), with comments on its phylogenetic

relationships

G.J, Howes

33 Areview of the serranochromine cichlid fish genera Pharyngochromis, Sargochromis, Serra-

nochromis and Chetia (Teleostei: Labroidei)

P.H. Greenwood

45 A revision of Danielssenia Boeck and Psammis Sars with the establishment of two new

genera Archisenia and Bathypsammis (Harpacticoida: Paranannopidae)

R. Huys and J.M. Gee

83 Anew species of Syrticola Willems & Claeys, 1982 (Copepoda: Harpacticoida) from Japan

with notes on the type species

R. Huys and S. Ohtsuka

95 Erratum

Bulletin of The Natural History Museum

ZOOLOGY SERIES

Vol. 59, No. 1, June 1993

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Zoology Series

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Bull. nat. Hist. Mus. Lond. (Zool.) 59(2): 97-105 Issued 25 November 1993

THE NATURAL

HISTORY MUSEUN

-6 DEC 1993

The status of the Persian Gulf sea snake [-

Hydrophis lapemoides (Gray, 1849)

(Serpentes, Hydrophiidae)

F hooEN | eD

alvavas BAF

aren el e

ARNE REDSTED RASMUSSEN

Zoological Museum, University of Copenhagen, Universitetsparken 15, DK-2100 Copenhagen @,

Denmark

CONTENTS

REE ECILU CELORN gece cee asn oe cee eC oee eNc EEE eoreoe av alesse ean eetna.s Helaeu ets saan Sasaan dues seb einanate san aes ameeteacion 97

NAME ITI ST ALICL EME LINO CLS Imereti nrs acres hececiecraat tea arc ioiaa,.seararOatetiorsiarteelee sapiaters aera cen ataraareie ers eran GA SelM oro cidteatielelctercaiace 97

RSV SEE TAAMNC HAC COIL mire ste dsrcmtaetr caine seats fee eprele= ce te dao saltete ears eeilelnge ee ee ieisites"9 slriein os siete twin lee MERE pete le dpe eons 98

PN CRMC MLE On ENILS Aetmemtetsiay icc ecaetanited acicnaes seit ee s'sece ct esa suisleh oocidhiaaieh sclaeaetumaniicasn dvessicanad ne seus attiaanieanes vsleieviere seine 103

ERC ST CLIC C QUEER a aN ten rae ee creat ine Ae laraiclnn'nnids nas Datei cclaadcattate oo evista talban/aicine nan taaceeswaatnntterttets sc sine ste aateveme 104

Synopsis. A redescription is given of the two syntypes of Hydrophis lapemoides together with a description of

specimens from the whole range of the species. Information on breeding and feeding biology and epizooic organisms

of H. lapemoides is provided. Geographical variation was found between the following three areas: Andaman Sea

and Malacca Strait, India and Sri Lanka, and Persian Gulf and Gulf of Oman. Finally the validity of H. lapemoides

is tested against its congeners.

INTRODUCTION

In 1849, Gray described Hydrophis lapemoides on the basis of

two specimens from Sri Lanka and Madras, respectively. In

addition to the syntypes Smith (1926) had eight specimens

available for characterizing the species. Smith recorded H.

lapemoides from the Persian Gulf, and coasts of India and

Ceylon, and considered it as a rare species. Volsge (1939)

collected eight specimens in the Persian Gulf and the Gulf of

Oman, and concluded that H. lapemoides is a fairly common

snake in these waters. Minton (1966) mentioned two speci-

mens from the coast of Pakistan, and in 1981 Toriba & Sawai

extended the known range of H. lapemoides from the East

coast of India to Penang Island, Malaysia. Tamiya et al.

(1983) identified some sea snakes from the Philippines as H.

lapemoides, the identification, however, was questioned by

Rasmussen (1987). Rasmussen recorded specimens from Sin-

gapore and Phuket Island, Thailand, thereby confirming the

presence of H. lapemoides in the Malacca Strait and Anda-

man Sea, respectively. Recently Gasperetti (1988) considered

H. lapemoides as the most frequent sea snake of the Persian

Gulf.

On the basis of my own collections in 1985, 1987 and 1989

from Phuket Harbour, and in 1990 from Bahrain, Persian

Gulf, I have a most welcome opportunity to describe H.

lapemoides from its whole range, and to test the validity of H.

lapemoides against its congeners. Some comments on the

biology of H. lapemoides are also given.

MATERIALS AND METHODS

Material examined

Hydrophis lapemoides BMNH: 1946.1.7.2 (syntype) (for-

merly III.3.3.a) Ceylon. 1946.1.6.91 (syntype) (formerly

I11.3.3.b) Madras. 1946.1.3.88 (type of H. stewartii) (for-

merly 83.7.30.10) Orissa, Poorie. 1946.1.9.25 (type of H.

holdsworthii) (formerly 72.1.26.41) Ceylon. 72.1.26.43 Cey-

lon. 80.11.10.199 Gwadur, Baluchistan. 1904.6.13.19 Mekran

coast, Charbar. 1969.2902 Persian Gulf. 1972.689 Dubai,

Trucial Oman. 1971.135-136 Bahrain. 1970.753 east coast of

Bahrain. 1971.1461 Bahrain harbour. 1973.410 Sharjah, Tru-

cial coast. 1983.1169 Najwa, Darninam reef. 1983.1163-1164

Dammam channel. 1983.1170 Ogair Bay. 1983.1172 Half

Moon Bay, Saudi Arabia. 1985.646 Azaiba, Batinah. FMNH:

28310-11 Bahrain. 64432 Tarut Bay, Ras Tonura. 73996-97 Al

Khobar, Arabia. 82577 Persian Gulf 26° 39'N, 50° 07’E.

121473 Ceylon. USNM: 127993 Ras Tanura, Saudi Arabia.

132402 Saudi Arabia. RMNH: 18026 Bahrain. ZMUC: R

66166-173 Persian Gulf (map in Volsge, 1939). 66101 Mal-

acca Str. (Singapore) 1° 35’N, 103° 01’E. R 66460, 66587-603,

66605-616, 66618-627, 66629-639 all collected from trawling

boats in Phuket port, Phuket Island, west coast of peninsular

Thailand. R 66927, 937, 939-941, 945, 950, 951, 954, 958, 961,

964, 965, 967, 968, 970, 971, 973-975, 980-1001, 1004-1006,

1009, 1011, 1013 Persian Gulf 100 km north-northeast of

Bahrain.

Fig. 1 Sulcate and asulcate side of everted hemipenis of H.

lapemoides (ZMUC R 66620) from Phuket Harbour. Drawing by

M. Andersen.

5mm

Fig. 2. Lateral aspect of anterior braincase of H. lapemoides in

which the sphenoid is included in the margin of the anterior

orifice of cavum epiptericum. Drawing by M. Andersen.

Methods

The measurements and counts follow Smith (1926) with some

alterations as described below. For lateral head scales, both

sides of the head have been examined and numbers are given

separately. Number of maxillary teeth is given for the right

side only.

Scale rows are counted directly around body (Thomas

1976). Minimum and maximum number of rows are given for

comparison with the counts of Smith (1926, p.XVI).

Position of the tip of the heart and the anterior tip of the

liver are determined in relation to the number of the adjacent

ventral scales (VS). Relative position of the tip of the heart

and the anterior part of the liver is expressed as the percent-

age of the total number of ventral scales (% VS). Vertebral

counts are obtained from soft radiographs. Three counts are

obtained from each snake: number of body vertebrae (VB-

body), number of tail vertebrae (VB-tail), and number of

vertebrae from the head to the tip of the heart (VB-heart).

Body and tail are separated by the presence of the first pair of

forked ribs in the cloacal region; this pair of ribs is included in

the number of tail vertebrae. Tip of the heart was indicated in

A. REDSTED RASMUSSEN

the x-ray radiographs by inserting a needle perpendicular to

the long axis of the body pointing at the vertebrae opposite

the tip of the heart. Relative position of the tip of the heart is

expressed as the percentage of the total number of body

vertebrae (% VB). Terms and description of the hemipenis

follow Dowling & Savage (1960). Hemipenis was described in

everted condition. All measurements are given to the nearest

centimeter. Meristic and mensural data given as x +SD.

In the following description of the two syntypes H.

lapemoides I11.3.3.a, a subadult female (syntype a) is given

first, and III.3.3.b, a juvenile male (syntype b) is given when

different from type a. The description of the examined

specimens is given in parentheses when different from the

types.

The following are abbreviations (Leviton et al., 1985) used

for the collections: BMNH: The Natural History Museum,

London; FMNH: Field Museum of Natural History, Chicago;

RMNH: Rijksmuseum van Natuurlijke Historie, Leiden;

USNM: National Museum of Natural History, Smithsonian,

Washington; ZMUC: Zoological Museum, University of

Copenhagen.

SYSTEMATIC ACCOUNT

Hydrophis lapemoides (Gray, 1849).

Aturia lapemoides Gray, 1849:46.

Hydrophis holdsworthii Gunther, 1872:33.

Hydrophis stewartii Anderson, 1872:399.

Distira lapemoides, Wall, 1909:227.

Hydrophis lapemoides, Smith, 1926:86, 1943:461. Volsge,

1939:19. Minton, 1966:146. McDowell, 1972:229. Voris,

1977:91. De Silva, 1980:399. Toriba & Sawai, 1981:134.

Rasmussen 1987:57, 1989:413, 1992:92. Gasperetti, 1988:312.

Bussarawit et al. 1989:222. McCarthy & Warrell, 1991:163.

Diagnosis

Eight to 13 maxillary teeth behind poison fang, 28-35 scale

rows on neck, 40-57 scale rows on body. Number of ventral

scales 288-395, tip of heart extending to ventral number

106-155. Number of body vertebrae 164-188, tip of heart

extending to vertebrae number 73-94. Head dark dorsally

with curved white mark, disappearing with age. Body with

black bands forming rhombic spots dorsally and disappearing

with age ventrally. Tail with black bands, disappearing with

age, posterior part normally black.

Description of the syntypes and the examined

specimens

EXTERNAL MORPHOLOGICAL CHARACTERS. Maxillary teeth

behind poison-fang 10. Dentary teeth, pterygoid teeth and

palatine teeth not counted on syntypes; for the examined

specimens see Table 1. One pre- and two postoculars on both

sides (one pre- and two or three postoculars). Three anterior

temporals on both sides (two or three). Eight supralabials on

both sides (7-10 in males, 7-10 in females). First and second

supralabials in contact with nasal, second and third in contact

with preocular, third and fourth in contact with eye, syntype

b; only third in contact with eye, fourth is divided horizon-

tally. Eight infralabials on both sides, first, second and third

on each side in contact with anterior pair of sublinguals,

THE STATUS OF HYDROPHIS LAPEMOIDES

JANETTE UETYTEE ATAU EATS ATE TATE ETAT TT A TTT

eer 2 3 6glCU ana |

2 5

8 Bo AA a = oe eee

Fig. 3 Habitus of the juvenile type specimen of H. lapemoides (BMNH 1946.1.6.91) from Madras, India. Photo by G. Brovad.

which are well developed and in contact with one another;

third and fourth infralabials touching posterior pair of sublin-

guals, which are well developed and separated from one

another posteriorly. A series of small cuneated scales at the

oral margin after the third infralabial, syntype b; second

infralabial. Scale rows on neck 29, syntype b; 32 (28-34 in

males, 28-35 in females), on body 45, syntype b; 51 (40-51 in

males, 41-57 in females). Ventrals 349, syntype b; 318 (288-

365 in males, 293-395 in females), distinct throughout, bicari-

nate, about twice as broad as adjacent scales anteriorly,

narrower posteriorly. Subcaudals 44, syntype b; 49 (37-56 in

males, 36-53 in females).

INTERNAL MORPHOLOGICAL CHARACTERS. Tip of heart

extending to ventral scale number 127, syntype b; 119 (106-

141 in males, 106-155 in females), %VS heart 36.38%,

syntype b; 37.42% (34.2-41.5 in males, 33.8-40.9 in females).

Anterior end of liver situated at ventral scale number 133,

syntype b; 120 (110-144 in males, 107-157 in females), %VS

liver 38.10%, syntype b; 37.74% (34.4-41.5% in males,

34.4-41.2%). In type a, a small interval separates the heart

and the liver. Number of body vertebrae 171, syntype b; 165

(164-188 in males, 171-186 in females). Number of tail

vertebrae 33, syntype b; 30 (31-40 in males, 28-38 in females).

Tip of heart extending to vertebrae number 81, syntype b; 82

(73-90 in males, 79-94 in females), % VB heart 47.36%, in

syntype b; 49.70% (43.5-51.1% in males, 45.1-51.7% in

females).

HEMIPENIS. Hemipenis feebly bilobed with a bifurcate sulcus

spermaticus (Fig. 1). Bifurcation near apical end of organ

(Fig. 1). Organ covered with spines gradually decreasing in

size and becoming more scattered at the distal end. A

finger-like fold at the proximal portion opposite the sulcus

spermaticus.

SKULL MORPHOLOGY (based on skulls from the Persian Gulf

and Andaman sea (Phuket)). Posterior half of parietal with a

distinct ridge being about 1/3 of the total length (midline).

Supratemporals (squamosals) reach parietal, and extend as

far posteriorly as posterior part of exoccipitals. Postorbital

bones barely touch frontals. Ventral extensions of frontals do

not overhang trabecular grooves. Sphenoid enters broadly

into margin of anterior orifice of cavum epiptericum (Fig. 2).

Sphenoid with low but distinct keel. Both anterior and

posterior Vidian foramen on the ventral side of sphenoid, and

100

Table 1 Geographic variation in number of the teeth on maxilla,

palatine, pterygoid and dentary bone.

n Maxillary n Palatine Pterygoid Dentary

teeth teeth teeth teeth

Andaman Sea

Malacca Str. 53 10-13 20 8-10 23-30 20-23

India

Sri Lanka 5 9-10 - - - =

Persian Gulf

Gulf of Oman 73 8-11 1 7-10 21-26 20-21

>= Se é

TTT LOT

A. REDSTED RASMUSSEN

going back, yellowish ventrally, some older specimens with-

out a curved mark. Body with black bands (29-52) forming

rhombic spots dorsally and disappearing with age ventrally

(Figs. 4 and 5). Tail with black bands (5-8), disappearing with

age, posterior part black (Fig. 4).

BREEDING BIOLOGY. Six of 17 females (collected in

September-November 1987) from Phuket were pregnant.

Three specimens contained 2 full-term embryos each, two

specimens contained 4 full-term embryos each, and one

specimen contained 1 full-term embryo. Pregnant females

were collected in the period 3rd October to 4th November.

None of the females collected in February-March 1989 were

pregnant. The smallest embryo measured 9 cm (3rd October)

Fig. 4 Habitus of a juvenile and an adult H. lapemoides (ZMUC R 66992, 66993) from the Persian Gulf. Photo by M. Andersen.

in respect of the length of the Vidian canals they are

symmetric. Palatine exceeding maxilla in forward extension,

and without a flange for the anterior medial process of

maxilla. Palatine-pterygoid articulation anterior to maxilla-

ectopterygoid articulation. Fangs separated from solid maxil-

lary teeth by a diastema. Maxillary bone slightly longer than

ectopterygoid. Solid maxillary teeth shorter than fangs. For

number of teeth on maxilla, palatine, pterygoid, and dentary

bones see Table 1.

COLourR. Juveniles: Head black with a yellowish curved

mark above, body yellowish or whitish, encircled by black

bands broadest dorsally (Figs. 3 and 4). Adults: Head dark

dorsally with curved white mark above, starting forehead

and the largest 26 cm (19th October). The female collected

4th November had embryos measuring 22 cm. Thus H.

lapemoides seems to be a k-strategist (Lemen & Voris, 1981)

producing small clutches of relatively large offspring.

None of the females examined from the Persian Gulf were

pregnant, however, Volsge (1939) mentioned three females

with eggs, and again the clutch size was very small (two

females with 2 eggs, one female with 3 eggs). Only two of the

three specimens have a collection date, and both were from

April (Volsge, 1939).

FEEDING BIOLOGY. Remains of the following four fish fami-

lies were identified in stomach contents from H. lapemoides

collected at Phuket harbour; Gobiidae, Labridae, Mullidae,

THE STATUS OF HYDROPHIS LAPEMOIDES

101

(HYNN)UTTTY HEEL) Avid ! (ili Wit

9 1 2 3 4 5 ' 8

{}{1| Wyk | itty

| , § 4 F r j 4

ee]

Fig. 5 Habitus of the subadult type specimen of H. lapemoides (BMNH 1946.1.7.2) from Sri Lanka. Photo by G. Brovad.

and Pseudochromidae. Pseudochromidae was found as prey

in a sea snake stomach for the first time, and Labridae and

Mullidae are new prey records for H. lapemoides (Voris &

Voris, 1983). The stomach contents from H. lapemoides

collected in Bahrain were too digested to be identified,

however, Volsge (1939) mentioned Gobiidae in stomachs of 5

specimens of H. /apemoides from the Persian Gulf. Further-

more Voris & Voris (1983) mentioned Anguilliformes and

Ophicthidae as stomach contents from H. lapemoides.

_ EPIZOOIC ORGANISMS. Five of the 51 specimens examined

from the Andaman Sea and Malacca Strait had between one

and five barnacles (Platylepas ophiophilus) on the skin. Two

of the seven specimens from India and Sri Lanka had three

and 20 P. ophiophilus on the skin, respectively. 25 of the 71

specimens from the Persian Gulf and the Gulf of Oman had

between one and 181 P. ophiophilus on the skin. Most of the

barnacles were on the posterior part of the body. P. ophio-

philus is found only on sea snakes (Zann et al., 1975), and has

_ been found on many species (Rasmussen, 1992; Zann, 1975).

DISTRIBUTION. H. lapemoides is found from the Persian Gulf

in west, to the Malacca Strait (Singapore) in east. Specimens

have been collected from the Persian Gulf, the Gulf of

Oman, the coast of Pakistan, India, and Sri Lanka, the west

coast of peninsular Thailand, Penang (Malaysia), and Sin-

gapore. (Ahmed, 1975; Bussarawit et al., 1989; Gasperetti,

1988; McCarthy & Warrell, 1991; Minton, 1966; Rasmussen,

1987; Smith, 1926, 1943; Toriba & Sawai, 1981; Vols¢e,

1939;).

RECENT COLLECTION DATA. H. lapemoides was collected in

different periods of 1985, 1987, and 1989 from fishing boats in

Phuket harbour, Phuket Island, on the west coast of peninsu-

lar Thailand. The most common sea snake brought to the

harbour by fishing boats was Lapemis hardwickii (over 80%

of all sea snakes caught by trawl) followed by H. ornatus, and

then H. lapemoides. According to the fishermen, the sea

snakes were caught mainly by sea-going trawler-boats, fishing

in waters more than 10 m deep. No further information was

available, as the fishermen were rather secretive about the

exact position of their fishing grounds. During collection in

the Persian Gulf (Bahrain) in February 1990, we went to an

area about 100 km north-northeast of Bahrain, on board a

trawling boat. On a 3 days trip we collected 7 specimens of H.

lapemoides, 2 specimens of Thalassophina viperina, and 1

specimen of H. ornatus. They were all caught by trawl in a

depth of 27-30 m, the bottom material was gravel. We also

collected sea snakes at Bahrain harbour from 6 trawling

boats, all working in the same area as mentioned above. Ina

period of 10 days (each boat was out 3 to 4 times in that

period), a total of 110 sea snakes was collected, and 96% of

the snakes were identified as H. lapemoides.

102

SPECIES ASSIGNMENT. The material examined is separated

into three geographical regions: Andaman Sea and Malacca

Strait, India and Sri Lanka, and Persian Gulf and Gulf of

Oman (Tables 2 and 3, Figs. 6 and 7). When comparing

specimens from the three areas mentioned above, geographi-

cal variation is found in general body form (specimens from

the Persian Gulf look more robust than specimens from

Andaman Sea and Malacca Strait), in scale rows on neck in

relation to scale rows on body (Fig. 6), in number of

vertebrae (Table 3), and in number of vertebrae in relation to

VB heart (Table 3, Fig. 7). However, it is difficult to decide

whether the variation indicates a cline or distinct geographic

forms, as material is still missing from Bangladesh and

Burma, and so are representative samples from Pakistan,

India, and Sri Lanka.

Both Boulenger (1896), Wall (1909) and Smith (1926)

referred the type specimen described by Anderson in 1872

under the name H. stewartii to H. lapemoides. Having

examined the specimen, I have serious doubt about its

assignment. 52 Scale rows on body in relation to 30 scale rows

on neck (Fig. 6), and 182 vertebrae in relation to 94 VB heart

(Fig. 7) indicate that the specimen belongs to a distinct taxon.

But as representative material is lacking from India and Sri

Lanka, I tentatively assign it to H.lapemoides. Further mate-

rial may show whether it is a valid taxon.

Dunson & Minton (1978) caught some sea snakes in the

Philippines, during the Visayan Sea Expedition of R/V Alpha

Helix, and identified them as H. ornatus. In 1983 Tamiya et

al. reclassified the specimens as H. lapemoides, and later

Rasmussen (1989) reexamined the specimens and identified

them as H. lamberti. Comparison of the above mentioned

specimens with H. lapemoides from Andaman Sea and Mal-

acca Strait, shows that they differ in following characters:

Scale rows on neck (H. lamberti, 37-45), VS heart tip (H.

lamberti, 87-109), and VS liver (H. lamberti, 86-108), VB

heart tip (H. lamberti, 65-71), and color pattern (Rasmussen,

1989). Comparing the skull, H. lamberti shows a more robust

parietal, with a longer ridge (from 1/2 to 2/3 of the total

length of parietal bone in midline), and with a less globular

form than H. lapemoides. A single specimen of H. lamberti

(FMNH 313058) was collected sympatrically with H.

lapemoides (ZMUC 66101) in the area of Singapore, and also

here the two species are distinct on the characters mentioned

above.

A. REDSTED RASMUSSEN

McCarthy & Warrell (1991) referred to a specimen

(BMNH 1987.172) from the Gulf of Siam (Samut Sakhon) as

‘H. sp near H. lapemoides’. 1 have examined this specimen

and agree that it is very similar to H. lapemoides, however, it

differs in number of scale rows around body in relation to

scale rows on neck (Fig. 6) and in number of vertebrae in

relation to VB heart (Fig. 7). Compared to H. lapemoides

from Malacca Strait and Andaman Sea it is very long (1.1 m)

and very robust in body and head form. In general shape it is

much closer to H. ornatus and H. lamberti, although the

characters differ here, too. Accordingly I think ‘H. sp. near

H. lapemoides’ should be separated from H. lapemoides, but

further studies are needed to find out whether the specimen

belongs to some of the more robust species in the Gulf of

Siam or is an unknown species.

Generic assignment

H. lapemoides has a combination of characters which places it

in the genus Hydrophis as defined by Smith (1926): maxillary

bone not extending forward beyond the palatine; poison-fang

followed, after a diastema, by from 1 to 18 teeth; palatine

straight; nostrils superior; nasal shields in contact with one

another; head shields large, regular; and ventrals small,

distinct throughout and normally entire.

McDOWELL’S SUBGENERIC ASSIGNMENT. In 1972 McDowell

recognized three subgenera in the genus Hydrophis, how-

ever, making a cladistic analysis (Rasmussen, in press) of the

subgenus Chitulia (formerly Aturia, see Williams & Wallach,

1989), the results indicated that the group was paraphyletic,

held together by plesiomorphic character states. Neverthe-

less, many of McDowell’s character states are most useful in

making a congeneric comparison.

Comparison with sympatric species

In the genus Hydrophis the following species are sympatric with

H. lapemoides: H. bituberculatus, H. brookii, H. caerulescens,

H. cantoris, H. cyanocinctus, H. fasciatus, H. gracilis, H.

inornatus, H. klossi, H. lamberti, H. mamillaris, H. melano-

soma, H. obscurus, H. ornatus, H. spiralis, H. stricticollis, and

H. torquatus. (Bussarawit et al., 1989; De Silva, 1980; Gasper-

etti, 1988; McCarthy & Warrell, 1991; Minton, 1966; Murthy,

Table 2. Geographic variation of external and internal characters in H.lapemoides.

Sex n Ventrals

Andaman M 28 288-347

Sea and x+SD 317413

Malacca F 23 299-378

Str. x+SD 341+20

India M 2 313-318

and Sri x+SD BISEESES

Lanka F 5 313-376

xtSD 347+24

Persian M 45 293-369

Gulf and x+SD 320+16

Gulf of F 25 300-395

Oman x+£SD 347423

VS-heart % VS-heart VS-liver % VS-liver

106-131 34.2-40.6 110-133 35.7-41.3

1186.7 37 SMe: 120+6.2 Stoned lS)

106-140 33.8-38.5 107-143 34.4-39.1

122==7 oi, Spy eel 124+7.7 36.3212

114-119 36.4-37.4 114-120 34.4-37.7

116435 36.9+0.7 117+4.2 37.1+0.9

117-145 35.1-39.0 117-146 35.1-39.3

127=WD B7-S2leS 129+13 37.8+1.8

111-141 35.0-41.5 113-144 36.3-41.5

I Pgjas 7/7) 38.5+1.4 124+7.1 38.8+1.4

114-155 34.6-40.9 114-157 34.6-41.2

129+10 37.5+1.6 130+10 37TEMG

VS-heart, VS-liver = position of tip of the heart and anterior tip of liver in relation to the number of the adjacent ventral scales, respectively. % VS-heart,

%\S-liver = relative position of tip of the heart and anterior tip of the liver in number of ventral scales, expressed as percentage of total number of ventral scales.

THE STATUS OF HYDROPHIS LAPEMOIDES

Table 3 Geographic variation of internal characters in H. lapemoides.

Sex n VB-body

Andaman M 28 164-174

Sea and XESD 170257

Malacca F 23 171-180

Str. x+SD 174+2.5

India M 2 165-174

and Sri x+SD 169+6.4

Lanka F 5 171-182

x+SD 176+4.9

Persian M 45 171-188

Gulf and +SD 17733

Gulf of F 26 172-186

Oman x+SD 181+3.6

103

VB-heart % VB-heart VB-tail

73-83 43.5—48.0 31-38

79+2.3 46.7+1.1 34+2.0

79-86 45.1-49.1 28-38

82+1.8 47.0+1.1 3142.5

79-82 45.4-49.7 37(n=1)

80+2.1 47.5+3.0

81-94 45.8-51.7 30-35

85==5e1 48.5+2.3 3341.9

79-90 45.6-51.1 33-40

8542.3 47.9+1.3 37217

81-90 46.6-49.2 30-36

86+2.4 48.0+0.8 Base ley

VB-body = number of body vertebrae. VB-heart = position of the tip of the heart in relation to the number of vertebrae. % VB-heart = relative position of tip of

the heart in number of vertebrae, expressed as percentage of total number of vertebrae. VB-tail = number of tail vertebrae.

1985; Rasmussen, 1987, 1989, 1992; Smith, 1926, 1930, 1943;

Taylor, 1965; Toriba & Sawai, 1981; Tweedie, 1983).

The sympatric species differ from H. lapemoides in the

following characters: H. cyanocinctus and H. spiralis have the

sphenoid nearly excluded from the ventral margin of the optic

fenestra (McDowell, 1972; Rasmussen, 1992: Fig. 5), a lesser

number (< 9) of maxillary teeth, (< 20) pterygoid teeth, and

(< 20) dentary teeth, and a different colour pattern (Bussa-

rawit et al., 1989; McDowell, 1972; Rasmussen, in press;

Smith, 1926). H. brookii, H. cantoris, H. fasciatus, H.

gracilis, H. klossi, H. melanosoma, and H. obscurus have a

triangular flange on the palatine (McDowell, 1972; Rasmus-

sen, 1992: Fig. 4), and a lesser number (< 8) of maxillary

teeth, (< 17) pterygoid teeth, and (< 16) dentary teeth

(McDowell, 1972). H. bituberculatus has a lesser number

(25-29) of scale rows around neck, a lesser number (247-290)

of ventrals, a lower position (90-105 VS) of heart tip, and a

different colour pattern (Rasmussen, 1992). H. caerulescens

has a higher number (14-18) of maxillary teeth (Smith, 1926),

a higher position (96-99 VB, based on 3 specimens from

Phuket harbour) of heart tip, and a different colour pattern

(Bussarawit et al., 1989; Smith, 1926; Tweddie, 1983). H.

inornatus (type specimen BMNH_ 1946,1.1.27 formerly

III.7.1.a.) has a lesser number (253) of ventrals, a lower

position (86 VS) of heart tip, a lower position (67 VB) of

heart tip, and a different colour pattern (Rasmussen, 1989).

H. lamberti is compared with H. lapemoides in the section

concerning species assignment. H. mamillaris has a smaller

head, a lesser number (25-29, 35-43) of scale rows on neck

and body, and a different colour pattern (Minton, 1966;

Smith, 1943). H. ornatus has a lesser number (224-294) of

ventrals, a lower position (72-104 VS) of heart tip, a lower

position (59-65 VB) of heart tip, and a different colour

pattern (Rasmussen, 1989). H. stricticollis has a smaller head,

a higher number (> 200 VB, Voris, 1975, and own observa-

tion) of vertebrae, and the hemipenis is bilobed half way

down. H. torquatus has a higher position (91-105 VB) of

heart tip, and a lesser number (7-8) of maxillary teeth (only in

Malacca strait) (own observation).

Comparison with allopatric species

In the genus Hydrophis the following species are allopatric

with H. lapemoides: H. belcheri, H. coggeri, H. czeblukovi,

H. elegans, H. geometricus, H. macdowelli, H. melanoceph-

alus, H. pacificus, H. parviceps, and H. vorisi. (Bussarawit et

al., 1989; Cogger, 1975; Kharin, 1983, 1984a, 1984b; McCar-

thy & Warrell, 1991; Smith, 1986; Smith, 1926, 1930, 1935).

The allopatric species differ from H. lapemoides in the

following characters: H. coggeri, H. czeblukovi, H. elegans,

H. melanocephalus, and H. pacificus have the sphenoid

nearly excluded from the ventral margin of the optic fenestra

(Kharin, 1984b; McDowell, 1972; Rasmussen, 1992: Fig. 5)

and a lesser number (< 9) of maxillary teeth, (< 20)

pterygoid teeth, and (< 20) dentary teeth (Kharin, 1984b;

McDowell, 1972). H. parviceps and H. vorisi have a triangu-

lar flange on the palatine (Kharin, 1984a; McDowell, 1972;

Rasmussen, 1992: Fig. 4), and a lesser number (< 8) of

maxillary teeth, (< 17) pterygoid teeth, and (< 17) dentary

teeth (Kharin, 1984a; McDowell, 1972; Smith, 1935). H.

belcheri has a lesser number (24-26, 32-36) of scale rows on

neck and body, a lesser number (14-17) of pterygoid teeth,

and no cuneate scales at infralabials (McCarthy & Warrell,

1991). H. geometricus has a high number (51-58, a small

overlap) of scale rows on body, and a different colour pattern

(Smith, 1986:152 Fig. 1). H. macdowelli has a lesser number

(< 8) of maxillary teeth, (< 16) pterygoid teeth, and (< 17)

dentary teeth, and a lesser number (256-266) of ventrals

(Kharin, 1983).

ACKNOWLEDGEMENTS. I thank the staff of The Phuket Marine Bio-

logical Center, Thailand, CODEC Project, Chittagong, Bangladesh,

Ministry of Commerce and Agriculture, Directorate of Fisheries,

Bahrain, S. Bagge, Cowi-Almoayed Gulf, Bahrain, J. Jensen,

Danida, and M. Andersen who helped me during the collection. The

Natural History Museum, London, Field Museum of Natural His-

tory, Chicago, Rijksmuseum van Natuurlijke Historie, Leiden,

National Museum of Natural History, Smithsonian, Washington for

loan of specimens. M. Andersen, A.B Helwigh, and especially Dr.

C. McCarthy (BMNH) and Dr. J. B. Rasmussen (ZMUC) for

valuable advice and constructive criticism of the manuscript. The

study was supported by Dansk Naturhistorisk Forening, The

Johannes Schmidts Grant, The Krista and Viggo Petersens Grant,

The Danish Research Academy, and The Danish National Research

Council, Grant no. 11-8209.

104

38 - + Andaman sea and Malacca Str.

(J) India and Sri Lanka

36 / ‘ Persian Gulf and Gulf of Oman

S 34 1 4 +

(= |

c =F + + 4

aoe OO mR ke L- t+ + &

= + K + +

oO |

(Ss) |

” 30 | O

26 1 +r r , 1 : + 1

38 40 42 44 46 48 50 52 54

Scale rows on body

+ Andaman sea and Malacca Str.

405

India and Sri Lanka

38 | < Persian Gulf and Gulf of Oman

H. sp. near H. lapemoides

36 5

3 > ea = ies ls li

5S 344 + se

4 KR ++

o 32) as +

3 | ae

” |

30 4 <8 oO (lType of H. stewarti

28 -

——————— oe

388 40 42 44 46 48 50 52 54 56 58 60

Scale rows on body

Fig. 6 Relation between number of scale rows on body and

number of scale rows on neck in males (top) and females

(bottom) of H. lapemoides, showing geographic variation.

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A. REDSTED RASMUSSEN

92 5

90 +

88 = ¥

86 - x

2 | x

tc 84,

= +

= 82 - Oo * =r

iS) | see +

S 80, tear ae

= | 2p SE aes lel

Cane 4 cee eee

+> coe

76, s + Andaman Sea and Malacca Str.

Ap és (India and Sri Lanka

72 Persian Gulf and Gulf of Oman

70 T T = T T T 1

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Number of vertebrae

96 - + Andaman Sea and Malacca Str.

| © India and Sri Lanka

94 - (_]*Type of H. stewartl

“Persian Gulf and Gulf of Oman

925 H. sp. near H, lapemoides

@ 90 - x

bs, |

gas

2 | x

So 86. O + rs

S +

= 84 + * + (+

82 a5 Ap Se jae

| Ox*+4+ 4+

80 - + i

78 -

7668 170 172 174 176 178 180 182 184 186 188

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Fig. 7 Relation between number of body vertebrae and position of

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Bull. nat. Hist. Mus. Lond. (Zool.) 59(2): 107-124

Issued 25 November 1993

Taxonomic revision of some Recent

agglutinated foraminifera from the Malay

Archipelago, in the Millett Collection, The

Natural History Museum, London

P. BRONNIMANN +

9G, Chemin de Bedex, 1226 Thénex/Geneva, Switzerland

J.E.WHITTAKER

Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD

Synopsis. Eleven species of Recent agglutinated foraminifera in the Millett Collection from the Malay Archipelago,

have been re-examined and revised systematically. They were originally described and illustrated in 1899 and 1900

with excellent lithographic drawings in the hand of Millett. With rare exceptions, the actual figured specimens,

though not marked as such, have been recognised in his Collection. The species are here redescribed, re-illustrated

by SEM photography and, where necessary, supplemented by new data, notably from similar environments in

Brazil. All, with the exception of Paratrochammina simplissima (Cushman & McCulloch) and possibly Trocham-

mina? milletti sp.nov., are brackish-water forms.

New taxa are Trunculocavus durrandi gen. et sp.nov. and Trochammina? milletti sp.nov. A lectotype is designated

for Acupeina triperforata (Millett), while Ammobaculites salsus var. distinctus Cushman & Br6énnimann is elevated

to specific rank and placed in Ammotium. All the species reviewed in this paper belong to the Suborder

Trochamminina.

INTRODUCTION

Durrand (1898) gives what locality information there is about

the Malay samples from which Fortescue William Millett

made his classic study of the foraminifera. Durrand had for

several years, out of his own interest, obtained small samples

from around the SW Pacific and had examined them for the

microscopical fauna and flora. In 1889 he had succeeded in

getting the Netherlands India Steam Navigation Company

(then controlled by the British India Steam Navigation Com-

pany) ... ‘to instruct the commanders of their fleet plying

about the islands of the Archipelago, to collect bottom from

each port of call’. The ‘cleaned material’ was picked over first

by Durrand and then the foraminifera were determined by

Millett and published (1898-1904) in 17 parts in the Journal

of the Royal Microscopical Society. In all, 468 species and

varieties were listed by Millett, 45 of them new. The descrip-

tions were accompanied by 19 plates of quite exceptional and

accurate drawings, from Millett’s own hand.

__ The samples came from anchor mud where the ships were

moored, more or less close inshore . . . ‘in about 12 or 14

fathoms’ (22-25 m). Unfortunately, a number of labels on

| the flasks of sediment became illegible through getting

soaked by leakage, so the locality information is somewhat

sketchy. The original samples each contained about 4 lbs

(1.8 kg) of solid matter.

The material came from two areas. Area 1 (‘from Celebes

in the north and west, to Java in the south and New Guinea,

# Deceased 6.1.1993.

Aru, and the Islands in the east, including such stations as

Banda, Amboina, Flores, Sumbawa and Timor’) contains

stations 1-16; area 2 (‘Singapore in the north, Banka in the

south, Sumatra in the west, and Borneo in the east’) contains

stations 17-31.

As part of a major revision of shallow-water agglutinating

foraminifera of the Indo-Pacific region (see also Br6nnimann

et al. 1992), eleven species belonging to the Trochamminina

are here redescribed, and illustrated by scanning electron

microscopy for the first time. The fauna has, for the most

part, strong affinities with the foraminifera of brackish,

mangrove sediments from other parts of the tropics, notably

Brazil. Comparison is therefore made with material described

by us (Bronnimann & Zaninetti, 1984a; b; Zaninetti et al.,

1977) from the mangroves of Guaratiba, Acupe and Baia de

Sepetiba, Brazil.

For a recent review of mangrove foraminifera in general

and their potential for palaeoenvironmental interpretation,

the reader is referred to an important paper by Culver (1990).

LOCALITY INFORMATION

Of relevance to the present revision are the following stations

from whence the specimens came; where the name of the

station is not mentioned, the label has become illegible. The

sample descriptions are in Durrand’s own words.

Area 1 Station 2 [no locality]. Plastic mud, brownish

tinted, rich in floatings.

108

Station 3 [no locality]. Brownish mud with lumps

of blueish clay throughout, residue about one

quarter-pound and floatings small.

Station 5 [no locality]. Blue ooze, residue and

floatings small.

Station 9 [no locality]. Results poor.

Station 11 [no locality].

Station 12 [no locality].

Station 14. Similar to Station 13 [Segaar, New

Guinea, coral sand and mud, residue about six

ounces, floatings rich].

Station 15 [no locality].

Area 2 Station 17. Muntok Banka, blue mud, residue

eight ounces, floatings rich.

Station 19 [no locality]. Earthy coloured, river-

looking mud, few foraminifera.

Station 21. Paney, northeast coast of Sumatra.

Station 27 [no locality].

Station 28 [no locality].

Durrand (1898:257) adds a postscript, stating that. . . ‘it is

important to bear in mind all this series was obtained from

shallow water close inshore . . .’. It is clear from the aggluti-

nating foraminifera revised here, that most of the localities

were in fact brackish, associated with mangroves.

SYSTEMATIC DESCRIPTIONS

Order FORAMINIFERIDA Eichwald, 1830

Suborder TROCHAMMININA Bronnimann & Whittaker,

1988

Apart from the hierarchy listed above, no further suprage-

neric taxa will be used. Until we can be certain that the

families and superfamilies of agglutinating foraminifera used

by Loeblich & Tappan (1987) represent homogeneous units

with respect to the wall structure, then it is better, for the

present, not to use them. Similarly, genera are used in

‘inverted commas’ when the wall structure of their type

species has not yet been examined. The eleven species

described here, at least, all have a Trochamminina-type wall,

defined by Bronnimann & Whittaker (1988) as . . . ‘consist-

ing of organic and agglutinated phases. Agglutinant bound by

organic cement and outer and inner organic sheets. Devoid of

perforations or alveolar pseudopores’.

The synonymies are not meant to be comprehensive, they

are selective, merely listing the original reference, junior

synonyms, changes of generic combination and important

citations from the study area.

P.BRONNIMANN AND J.E. WHITTAKER

Genus ACUPEINA Bronnimann & Zaninetti, 1984b

TYPE SPECIES. Haplophragmium salsum Cushman & Bronni-

mann, 1948a (= junior subjective synonym of Haplophrag-

mium agglutinans d’Orbigny vat. triperforata Millett, 1899).

Acupeina triperforata (Millett, 1899) Figs 1.2, 13-15

1899 Haplophragmium agglutinans d’Orbigny var. triper-

forata Millett: 358(pars); pl. 5, figs 2a,b (lectotype)

only; non figs 3a,b.

Haplophragmium salsum Cushman & Bronnimann:

16,17; pl. 3, figs 10-13.

1965 Lituola salsa (Cushman & Bronnimann); Bronni-

mann & Zaninetti: 608-615; figs 1-3.

Acupeina salsa (Cushman & Bronnimann); Brénni-

mann & Zaninetti: 219-222; figs Al-4, B1,2.

Acupeina triperforata (Millett); Bronnimann &

Zaninetti: 222 (addendum).

1988 Acupeina triperforata (Millett); Bronnimann & Whit-

taker: 112; pl. 4, figs 1-7.

1948a

1984b

REMARKS. Millett (1899, pl. 5, figs 2,3; here reproduced as

Figs 1.2, 3) illustrated four views of his new variety triperfo-

rata. Examination of the original material shows that two

different brackish species are involved: Acupeina triperforata

(Millett) and Arenoparrella mexicana (Kornfeld).

The individual drawn by Millett (1899, pl. 5, figs 2a,b; here

reproduced in Fig. 1.2a,b) in side and apertural views, has

been re-illustrated by SEM in Figs 13-15. The micrographs

show side and edge views of a test, initially streptospiral then

uniserial, and the detail of the multiple aperture which

consists of three closely spaced, virtually equidistant pores (of

around 25 um diameter) with upturned rims. Millett appar-

ently believed that the aperture of his new variety invariably

consisted of the three rounded pores, hence the name. The

individual in Figs 13-15 is undoubtedly Millett’s figured

specimen and is here formally designated lectotype.

The specimen drawn by Millett (1899, pl. 5, figs 3a,b;

reproduced here in Fig, 1.3a,b) in side and apertural views,

has been re-illustrated by SEM in Figs 9-12 not only to show

both sides of the test but the details of the composite

aperture. Its morphology is quite different from the lectotype

of H. agglutinans var. triperforata. It represents, in fact, a

typical specimen of Arenoparrella mexicana (Kornfeld,

1931)(see below). It is unfortunate that Loeblich & Tappan

(1987: 21, pl. 71, figs 3,4) illustrated this very specimen,

together with the lectotype, as A.triperforata. It is also worth

noting that Millett’s pl. 5, fig. 3b is the edge view of fig. 3a,

but as can be seen from our SEM illustration, rather mislead-

ing. It purports to show only three large pores with everted

borders. In reality, it has a single oblique-perpendicular slit

and 12 small peripheral pores of 5—6 um diameter, devoid of

rims. Closer examination of Millett’s apertural view (see Fig.

1.3b) may just show the termination of the slit (the specimen

is tilted forward), but the determination of the pores is still

seriously in error.

Fig. 1.1-1.10, 1.12 Reproduction of part of Plate 5 of Millett (1899). The original identifications were as follows: Fig. 1.1, Haplophragmium

agglutinans (d’Orbigny), X112; Fig. 1.2, 3, H. agglutinans var. triperforata var.nov., X112; Fig. 1.4-6, H. cassis (Parker), X112; Fig. 1.7,

A. cassis (Parker) or ?Reophax, X75; Fig. 1.8, H. compressum Goés, X75; Fig. 1.9, H. nanum Brady, 112; Fig. 1.10, H. anceps Brady,

X56; Fig. 1.12, Trochammina ochracea (Williamson), X75. Reproduced by permission of the Royal Microscopical Society.

Fig. 2.1

Reproduced by permission of the Royal Microscopical Society.

Reproduction of part of Plate 1 of Millett (1900). It was originally identified as Bigenerina digitata d’Orbigny var., X169.

TAXONOMIC REVISION OF FORAMINIFERA IN MILLETT COLLECTION 109

JOURN.R.MICR. SOC.1900. Pl.1,

JOURN.R.MICR. SOG. 1899. Pl. V.

110

Bronnimann & Zaninetti (1984b: 222, Addendum) have

shown that Haplophragmium agglutinans d’Orbigny var. trip-

erforata Millett (1899) is identical with H.salsum Cushman &

Bronnimann (1948a),which is the type species of Acupeina

Bronnimann & Zaninetti, 1984b.

LECTOTYPE. The individual illustrated by Millett (1899, pl. 5,

figs 2a,b; Figs 1.2, 13-15 herein) is designated lectotype of H.

agglutinans vat. triperforata, now Acupeina triperforata. It is

deposited in the collections of the BMNH, no.

1955¢ 151 1076:

DIMENSIONS (LECTOTYPE). Height of test — 380 wm; diam-

eter of coiled portion — 235 um; maximum diameters of

apertural pores — 25 um, with everted rims 4 um high.

ENVIRONMENT. This species. . . ‘is not uncommon at Station

9, and occurs also, but very sparingly, at Station 5’ of Area 1.

At Station 9, Millett (1899: 359) also reported Haplophrag-

mium cassis (Parker) (= Ammoastuta salsa Cushman &

Broénnimann and Ammotium spp.), all brackish water spe-

cies. Both Acuipeina triperforata and Arenoparella mexicana

are also exclusively brackish forms, occurring commonly in

tropical to subtropical mangrove swamp sediments.

Genus AMMOSASTUTA Loeblich & Tappan, 1984

TYPE SPECIES. Ammoastuta salsa Cushman & Bronnimann,

1948a.

Ammoastuta salsa Cushman & Bronnimann, 1948a

Figs 1.6, 35

1899 Haplophragmium cassis (Parker); Millett (pars): 359;

pl. 5, figs 6a,b only (non figs 4,5,7) (non Lituola

cassis Parker, 1870).

Ammoastuta salsa Cushman & Bronnimann: 17; pl.

3, figs 14-16.

1970 Ammoastuta salsa Cushman (sic); Hofker: 3.

1986 Ammoastuta salsa Cushman & Broénnimann; Bronni-

mann: 29-44; figs 1-7. (q.v. for synonymy).

1948a

REMARKS. Millett (1899: 359, pl. 5, figs 6a,b; here repro-

duced as Fig. 1.6a,b) figured side and edge views of a slightly

damaged, but clearly recognizable specimen of Ammoastuta

salsa under the name of Haplophragmium cassis (Parker). He

also illustrated two different species of Ammotium (pl. 5, figs

4a,b, 5a,b; Figs 1.4, 5) and, used for all these different

morphologies the same name, as he thought . . . ‘the Malay

specimens of this species [H. cassis] are very variable in form,

some of them being extremely compressed, and composed of

numerous chambers’.

The SEM photograph of the side view (Fig. 35), although

now slightly more damaged, is demonstrably of the same

specimen as in Millett’s drawing. The tight initial coil cannot

be seen, but on the other hand, the final two chambers of the

P.BRONNIMANN AND J.E. WHITTAKER

juvenile stage are clearly visible. The adult consists of at least

7 elongate uniserial chambers which make up the main

portion of the compressed test.

Bronnimann’s (1986) morphological revision of A. salsa

has shown that the test starts with a tight early spiral

consisting only of a proloculus and deuteroloculus. On the

basis of this arrangement, Ammosastuta is correctly placed in

the Lituolidae. Loeblich & Tappan (1987: 79) accepted this

interpretation, but stated that the second chamber is growing

in the . . . ‘opposite direction’ (without saying in respect to

what). This is simply not the case. The second chamber

develops from a porus in the side of the proloculus. It is just

the normal forward continuation, considering the flow of the

protoplasm, which produces the elongate deuteroloculus with

a porus at its apex. Hence the embryonic chambers form a

tight, reduced spiral (see Bronnimann, 1986: 32, fig. 3).

Ammoastuta salsa is occasionally placed in synonymy with

Ammobaculites (=Ammoastuta) ineptus Cushman & McCul-

loch, 1939. Cushman & Brénnimann (1948a) regarded the

two as distinct, as did Bronnimann (1986). An examination

by Bronnimann of the two paratypes of A. ineptus, deposited

in the collections of the U.S. National Museum of Natural

History, Washington, confirms this separation. Of the

paratypes, only one, registration no. 35826, is well preserved.

It is definitely an Ammoastuta, but differs from the com-

pressed A. salsa by having a strongly inflated test.

DIMENSIONS OF FIGURED SPECIMEN (BMNH no.

1955.11.1.1121). Maximum height (damaged) — 280 um.

ENVIRONMENT. Recorded from Station 9, Area 1. It occurs

together with Acupeina triperforata, Ammotium spp. and

Arenoparella mexicana, all typical brackish water species.

Genus AMMOBACULITES Cushman, 1910

TYPE SPECIES. Spirolina agglutinans d’Orbigny, 1846. Lecto-

type designated by Loeblich & Tappan (1964: C241, figs

251.6a,b).

REMARKS. The genus Ammobaculites Cushman (1910) con-

tains free agglutinated tests with a simple interior; the early

portion is planispiral, the later part uncoiled and rectilinear.

It is radially-symmetrical in transverse section. The single

aperture is terminal, areal and radially symmetrical. The wall

structure of the type species is unknown.

This definition is more restrictive than Loeblich & Tap-

pan’s (1987: 74) as it not only excludes streptospiral and

trochospiral initial coils, but also laterally compressed tests.

The transverse sections of the chambers of the uncoiled

portion of the test and the outlines of the terminal apertures

are radially symmetrical; these features are regarded as

important generic criteria.

The wall structure of Ammobaculites exiguus, the species in

the Millett Collection, is of the Trochamminina type. If A.

Figs 3-8 = Trunculocavus durrandi gen. et sp.nov. Figs 3,4, Detail of aperture (X900) and side view (X160), respectively. Holotype, BMNH

no. 1955.11.1.187; Fig. 5, Side view (X175). Paratype, BMNH no. 1911.11.1.189; Figs 6-8, Detail of initial coil (X540), aperture (730)

and side view (X160), respectively. Paratype, BMNH no. 1955.11.1.188.

Figs 9-12 Arenoparrella mexicana (Kornfeld). Detail of apertures (X480), side, edge and view of other side (X160), respectively. BMNH

no. 1955.11.1.1075.

Figs 13-15 Acupeina triperforata (Millett). Edge and side view (X160) and detail of aperture (X700), respectively. BMNH no.

NOTA. LeLO76:

All from the Millett Collection, Malay Archipelago.

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112

agglutinans, the type species, should have the same wall type,

then exiguus would be correctly placed in Ammobaculites. If

not, then it would have to be placed in a new genus. In view

of these uncertainties, Ammobaculites is placed in inverted

commas in our treatment of ‘A’. exiguus.

‘Ammobaculites’ exiguus Cushman & Bronnimann,

19485 Figs 1.1, 42-44

1885 Haplophragmium agglutinans (d’Orbigny); Balkwill

& Wright: 330; pl. 13, figs 18?, 19,20 (non d’Orbigny,

1846).

1899 Haplophragmium agglutinans (d’Orbigny); Millett:

357, pl. 5, figs 1a,b.

1938 Ammobaculites agglutinans (d’Orbigny); Barten-

stein: 391; fig. 14.

1948b Ammobaculites exiguus Cushman & Bronnimann:

38; pl. 7, figs 7,8.

1952b Ammobaculites cf. exiguus Cushman & Bronnimann;

Parker: 443;pl. 1, figs 16,17.

1952. Ammobaculites agglutinans (d’Orbigny); Rottgardt:

180; pl. 1, fig. 4. 1954 Ammobaculites exiguus Cush-

man & Brénnimann; Phleger: 633; pl. 1, fig. 5.

1956 Ammobaculites sp. B, Warren: 139; pl. 1, figs 22-24.

1957 Ammobaculites exiguus Cushman & Brénnimann;

Todd & Bronnimann: 23; pl. 2, fig. 7.

1962 Ammobaculites exiguus Cushman & Br6énnimann;

Benda & Puri: 335; pl. 1, fig. 15. 1973 Ammobacu-

lites balkwilli Haynes: 25-27; pl. 2, figs 2,3; pl. 29,

figs 5,6; text—fig. 4.1—S.

1978 Ammobaculites dilitatus (sic) Cushman & Bronni-

mann; Schafer & Cole: pl. 3, fig. 9 (non Cushman &

Bronnimann, 19485).

1980 Ammobaculites dilatatus Cushman & Bronnimann;

Scott & Medioli: 35; pl. 1, figs 9,10.

Ammobaculites exiguus Cushman & Bronnimann;

Haman: 72; pl. 5, figs 14.

1983. Ammobaculites diversus Cushman & Bronnimann;

Haman: 72; pl. 4, figs 14,15 (non Cushman & Bron-

nimann, 1948b).

1986 Ammobaculites exiguus Cushman & Brénnimann;

Bronnimann & Keij: pl. 3, fig. 7.

21983

REMARKS. Millett (1899: pl. 5, figs 1a,b; here reproduced as

Figs 1.1a,b) illustrated, without description, a typical speci-

men of exiguus under the name of Haplophragmium agglutin-

ans (d’Orbigny). The same specimen (BMNH_ no.

1955.11.1.1057) is re-illustrated by SEM in our Figs 43,44.

The oblique view (Fig. 43) shows the radially-symmetrical

areal and terminal aperture, which is larger than in Millett’s

drawing. It is not bordered by a rim as that shown by

Haman’s (1983, pl. 5, figs 1-4) ‘A. exiguus’, which may

represent a different species. Millett’s specimen has four

uniserial chambers which follow from a planispiral, tightly

enrolled early test. The agglutinant is coarse and the sutures

in the initial portion are not well defined; on the uniserial

portion, they are distinct, however, and run perpendicularly

to the elongate axis of the test.

Illustrated in Fig. 42 (BMNH no. 1911.11.1.1058) is a

smaller, albeit damaged specimen, which is more typical of

the size of the Malay material. Four radial sutures can be

recognized in the coiled portion and there are three chambers

in the uniserial part; the final chamber is crushed.

One of us (P.B.) has re-examined the holotype of A.

P.BRONNIMANN AND J.E. WHITTAKER

exiguus (registration no. 56761) in the U.S. National Museum

of Natural History. Its overall morphology corresponds well

with Millett’s illustrated specimen of H. agglutinans. How-

ever, in its uniserial portion there are five chambers and the

agglutinant is finer than in the Malay specimen. Nevertheless,

the two both have a circular transverse section and a large

radially-symmetrical, terminal aperture without a rim; the

intercameral sutures run perpendicular to the elongate axis of

the test, there being no suggestion of Ammotium-type

sutures. In addition to the holotype of exiguus, there are two

slides with paratypes: in slide no. 56762 there is a single

paratype; under no. 56763 there are, amongst typical speci-

mens, some very small individuals which differ from the type

by their thin, elongate tests. These latter have also been

encountered by us in the mangrove sediments of Acupe,

Brazil. They represent a new species of brackish ‘Ammobacu-

lites’ which will be published elsewhere. It should be noted

that ‘A’. exiguus and this new, minute species, are the only

true representatives of ‘Ammobaculites’ occurring in brackish

waters.

DIMENSIONS OF FIGURED SPECIMEN (BMNH ono.

1911.11.1.1057). Height of test — 385 um; diameter of initial

planispiral portion — 135 um; diameter of final chamber —

125 um; diameter of aperture — 50 um.

ENVIRONMENT. In the Millett Collection, specimens are

labelled ‘Haplophragmium agglutinans’ from _ stations

2,9,12,14,15,19,21 and 27; Millett notes (p. 358) that . . . ‘the

specimens are all minute, and although they occur at most of

the Stations, are not very numerous’. According to Parker et

al. (1953), ‘A’. exiguus is a species which lives in brackish as

well as in marine waters.

Genus AMMOTIUM Loeblich & Tappan, 1953

TYPE SPECIES. Lituola cassis Parker (in Dawscn), 1870.

REMARKS. Ammomarginulina Wiesner, 1931 (type species:

A. ensis Wiesner, 1931) is a deep-water genus, with a

morphology close to that of the supposedly exclusively

brackish-water genus, Ammotium. After having compared

the definitions of Ammomarginulina and of Ammotium in

Loeblich & Tappan (1987), the question arises as to whether

the two are really synonymous. The sutures of the former are,

however, less slanting that those of Ammotium, and the test

is strongly compressed. Of the shape of the aperture of

Ammomarginulina ensis nothing is known except for the fact

that it is rounded. Small morphological differences such as

these may not be considered sufficient to retain the two

genera. However, they seem to represent two disparate

homogeneous environmental groups which, should this be

sustained, must be separated taxonomically, even if the

morphological differences were even less pronounced (see

also Resig’s (1982: 977-978, pl. 1, figs 3-5,9) description of

Ammomarginulina hadalensis Resig from the Peru-Chile

Trench, depth 5846 m). Clearly, the wall structure of Ammo-

marginulina must also be investigated.

Figs 32-34, 54

1899 Haplophragmium cassis (Parker): 359 (pars) (non

Lituola cassis Parker, 1870).

1940 Ammobaculites morenoi Acosta: 272; pl. 49, figs 3,8

(holotype) only (non Fig. 1).

Ammotium morenoi (Acosta, 1940)

TAXONOMIC REVISION OF FORAMINIFERA IN MILLETT COLLECTION

1948a Ammobaculites salsus Cushman & Brénnimann: 16;

pl. 3, figs 7a,b,8,9 (holotype figs 7a,b).

1952b Ammoscalaria fluvialis Parker: 444; pl. 1, fig. 24

(holotype) only (non fig. 25).

1953. Ammobaculites salsus (et vars.) Parker et al.: 5; pl. 1,

figs 18-25 only (non fig. 17).

1954 Ammobaculites exilis Cushman & Br6nnimann;

Phleger: pl. 1, fig.6 (non Cushman & Bronnimann,

1948).

1954. Ammobaculites salsus Cushman & Br6énnimann;

Phleger: pl. 1, fig. 7 only (non fig. 8).

1954 Ammoscalaria fluvialis Parker; Phleger: pl. 1, fig. 11.

1957 Ammobaculites salsus Cushman & Bronnimann;

Todd & Bronnimann; 24, pl. 3, fig. 8

1958 Ammobaculites salsus Cushman & Bronnimann;

Arnal: 37; pl. 98, figs 4-7.

1968 Ammotium salsum (Cushman & Bronnimann);

iutze: 25; pl.1; figs 5,6.

1978 Ammotium salsum (Cushman & Bronnimann); Poag:

405; pl. 5, figs 1-39.

1980 Ammotium salsum (Cushman & Br6énnimann); Scott

& Medioli: pl. 1, figs 11-13.

1983. Ammotium morenoi (Acosta); Haman: 72; pl. 5, figs

6-9.

REMARKS. The specimen illustrated by us in Figs 32-34 was

not figured or described by Millett but comes from a slide in

the Millett Collection labelled Haplophragmium cassis Parker

(BMNH no. 1955.11.1.1118-1133) and was undoubtedly part

of his concept of that species. It is a typical representative of

Ammotium morenoi. The small test is axially compressed and

consists of a short, completely coiled planispiral initial stage,

followed by an uncoiled portion of about 5 low, elongate,

laterally compressed chambers which on the interior side

reach back toward the initial planispire. The single aperture is

a narrow elongate slit with rounded extremities, situated at

the apex of the final chamber, in a marginal or outer position.

Under the name of H. cassis, Millett (1899, pl. 5, figs 4a,b,

5a,b) did illustrate two specimens, which belong to different

species of Ammotium. The latter (reproduced here as Fig.

1.5a,b) is the upper part of an Ammotium pseudocassis

(Cushman & Bronnimann, 1948b)(see p. , below) but was not

found in the Millett Collection. The former (Fig. 1.4a,b) is a

complete specimen of A. directum (Cushman & Bronnimann,

19485) and is refigured in Fig. 31.

In addition to this specimen, we have also illustrated in Fig.

54, for the purpose of comparison, the lateral view of a

typical specimen of A. morenoi from the mangrove sediments

of Guaratiba, Brazil (see Zaninetti et al., 1977). It consists of

an initial, almost involute planispire, followed by two unise-

rial, laterally flattened, low and elongate chambers, which on

the inner side extend backwards toward the early spire.

In common with other brackish foraminifera, A. morenoi is

highly variable in its overall morphology, in particular in size

| and in outline of the test in lateral view. From small, almost

triangular forms, as represented by the holotype of morenoi

or the holotype of Ammoscalaria fluvialis Parker (1952b), we

find all possible transitions to the elongate slender specimens

of Ammobaculites salsus described by Cushman & Bronni-

mann (1948a) from Trinidad, or to the large and elongate

individuals recorded by Poag (1978) from Gulf Coast estuar-

ies. Brodniewicz (1965: 187-194, text-figs 21-25) has shown

that a similar form, identified by her as Ammotium cassis

(Parker), from the Baltic, is also characterized by a great

113

morphological variability. She tried to distinguish six differ-

ent morphological types on the basis of outline, chamber

form, and dimensions of the test and chambers. A study of

Brodniewicz’s paper, however, suggests to us that it is

virtually impossible to separate her different morphotypes.

DIMENSIONS OF FIGURED SPECIMENS (MALAY SPECIMEN,

BMNH no. 1991.11.1.1122). Height of test — 170 wm; width

(length) — 105 um; thickness — 35 um.

(BRAZILIAN SPECIMEN). Height of test — 370 um;

maximum width — 190 um; final chamber — 225 um high;

maximum diameter of oblong aperture — 50 um.

ENVIRONMENT. Found only in Station 9 (Area 1) in associa-

tion with Ammotium pseudocassis, A. directum, Acupeina

triperforata, Ammoastuta salsa and Arenoparella mexicana,

all typical brackish water species. A. morenoi is normally

abundant in tropical and subtropical mangrove sediments but

has also been recorded, albeit rarely, in brackish sediments of

temperate climes (Parker, 1952b; Lutze, 1968). We have

never encountered this species in the British Isles or in the

Mediterranean.

OBSERVATIONS ON CERTAIN SYNONYMS AND NON-SYNONYMS

(NEAR ISOMORPHS) OF AMMOTIUM MORENOI ACOSTA.

1. Ammobaculites salsus Cushman & Br6nnimann,

1948a and A. distinctus Cushman & Bronnimann,

1948b.

Haman (1983) was the first author to place Ammobaculites

(=Ammotium) salsus into synonymy with Acosta’s species.

In the introduction to his paper, Acosta (1940: 269) wrote

that the agglutinating species were rare in the shallow water

assemblages from the Gulf of Santa Maria, Camaguey Prov-

ince, Cuba, which were dominated by miliolids and nonion-

ids. The Gulf of Santa Maria is bordered by extensive

mangrove swamps. It is therefore assumed that the tests of

the brackish agglutinated species, such as A.morenoi, had

been transported by wave action into the marine environment

of the open Gulf and were not in situ at the locality where

Acosta collected them. Acosta (1940: 275) claimed to have

deposited the types of his species in the Cushman Collection,

which were later transferred from Sharon, Massachusetts to

the U.S. National Museum of Natural History, Washington,

D.C. A search by P.B. for the type specimen of A. morenoi

proved unsuccessful and it seems that Acosta never did

deposit his types in the Cushman Collection. Acosta’s draw-

ings (op.cit. pl. 49, figs 3,8, non fig. 1) leave no doubt,

however, that Ammotium morenoi and A. salsum, originally

described from Trinidad mangrove swamps, are one and the

same.

When comparing the two ‘species’, the apertural view of

the holotype of Ammotium morenoi (Acosta, 1940: pl. 49,

fig. 8) is of interest. It shows a slit-like opening at the apex of

the final chamber, in a marginal or outer position; the same

type of aperture occurs in A. salsum. In both holotypes the

peripheral outline of the initial planispire, as seen laterally, is

perfectly rounded and not angular as in Ammotium distinc-

tum (Cushman & Bronnimann (19485: 40, pl. 7, fig. 14),

which has also been described from the brackish mangrove

sediments of Trinidad. This latter form was originally intro-

duced as a variety of Ammobaculites salsus. As there are no

intermediates between distinctum and salsum, the former is

here elevated to specific rank. Authors, however, normally

114

make no distinction between the two (see Phleger, 1954: pl.

1, fig. 8).

We have illustrated in Fig. 55 a lateral view in oil of

Ammotium distinctum, from the mangrove sediments of

Acupe, Brazil. The angular outline of the early planispire is

clearly shown. The test begins with a relatively large prolocu-

lus of 65 um diameter, followed by a larger deuteroloculus of

75 um diameter. The total number of chambers in this

specimen is eight, including the embryonic chambers. The

height of the test is 220 um, width (length) 125 wm, and

length of aperture 45 um. Apart from the distinct angular

periphery, there are no other important differences between

Ammotium morenoi and A. distinctum.

2. Ammoscalaria fluvialis Parker, 1952b.

Parker (1952b: 444, pl. 1, fig. 24) first described this species

from the Housatonic River, Long Island Sound, depth 3 m.

From its association with other brackish species in her Facies

1, such as Trochammina inflata, Jadammina macrescens and

Miliammina fusca, it can be inferred that A. fluvialis is also a

brackish-water form. The morphology of the holotype is

virtually identical with the holotype of A. morenoi, and for

this reason we regard it as a junior synonym of the latter.

3. Lituola cassis Parker, in Dawson, 1870.

We have compared Ammotium morenoi with Lituola cassis

Parker, the type species of Ammotium Loeblich & Tappan

(1953). The lectotype of Ammotium cassis (BMNH no. ZF

4637), designated by Hodgkinson (1992), on our advice, is

re-illustrated in Figs 38-41. It is from Gaspé Bay, Gulf of St.

Lawrence, Canada, and came from the W.K. Parker Collec-

tion; it was collected in 16 fathoms (30 m), which suggests a

marine environment, but the specimens could have been

washed in from a brackish locality. Loeblich & Tappan (1987,

pl. 60, figs 1,2) illustrate a ‘Holocene’ specimen from off

Alaska in 223 m of water; should this specimen have been in

situ it would further undermine the supposedly exclusively

brackish nature of the genus, a factor that needs further

investigation.

The lectotype clearly shows the initial planispire, then the

uniserial inward slanting narrow and low chambers; the

oblong aperture is at the apex of the final chamber, in a

marginal or outer position (see also Goés, 1894, pl. 5, figs

152-157). The lectotype and paralectotypes are five times

larger and much more massive than A. morenoi, though the

two in several other respects are quite similar. It is our

opinion that A. cassis should only be used for large and

massive individuals, but at the same time we have our

reservations that ecological factors (?marine salinities) may

be responsible for the massive development of the cassis test

(see also remarks above, on A. cassis sensu Brodniewicz

(1965) from the Baltic). It is even three times the size of

Poag’s (1978) material from the Gulf Coast estuaries, the

largest known specimens of A. morenoi from the tropics,

P.BRONNIMANN AND J.E. WHITTAKER

moreover Poag’s specimens are very elongate and com-

pressed with the uniserial portion quite unlike that of the true

cassis.

The dimensions of the lectotype are: maximum height —

1600 um; maximum width — 785 um; maximum thickness —

360 um; thickness of planispiral portion — 125 pm.

4. Ammobaculites prostomum Hofker, 1932.

This species was described by Hofker (1932: 87-91, text—figs

14a-f, 15a—d) from the Ammontatura, a part of the Gulf of

Naples, with a depth of 150-200 m. The shapes of the

illustrated specimens, seen laterally, particularly the short

individuals (text-figs 14a and f), much resemble the small

specimens of Ammotium morenoi such as our Fig. 54. On

Hofker’s short specimens the sutures are not shown fully, but

on the larger specimens (text—-fig. 15e) they are, toward the

outer margin, at first outward slanting (not inward), then

parallel up to the end of the uniserial portion. In lateral

outline, these short specimens are near isomorphs of A.

morenoi. However, the aperture is not placed asymmetri-

cally, at the outer margin of the test as in Ammotium, but

symmetrically in respect to the shape of the final chamber.

For these reasons, Hofker’s species does not belong to

Ammotium. It is also a marine species and much resembles

the group pf forms described and illustrated by Hoglund

(1947, pl. 31, figs la-g) from Bjorkholmen, Gullmar Fjord,

from a depth of 30 m, under the name of Ammoscalaria

pseudospiralis (Williamson).

5. Ammoscalaria pseudospiralis sensu Héglund, 1947.

The genus Ammoscalaria was erected by Héglund (1947:

151-153) with Haplophragmium tenuimargo Brady (1884) as

type species. Into his new genus he also placed Proteonina

pseudospiralis Williamson, 1858. However, Ammoscalaria

pseudospiralis was described by Héglund (1947: 159-162, pl.

31, figs la—p) exclusively from material obtained in the

Gullmar Fjord, where it occurs commonly, and from the

Skagerak, not on the basis of Williamson’s material which

was not available to him. The chambers of the rectilinear

portion of this marine species are ‘irregularly rectangular in

lateral view’ and there are ‘no external sutures’. The oblong

aperture is in a symmetrical position in respect to the final

chamber and not asymmetric, as in Ammotium. We therefore

do not regard Héglund’s species as a synonym of pseudospira-

lis, although certain smaller specimens could be regarded as

isomorphs of Williamson’s taxon, particularly when seen in

lateral view (e.g. pl. 31, figs 1m,n). Rather, Héglund’s form

is most probably a junior synonym of Ammobaculites

(=Ammoscalaria) prostomum Hofker, 1932.

Figs 16-21

‘Haplophragmoides’ wilberti Anderson. Figs 16-18, Side,edge and view of other side (X115). BMNH no. 1911.11.1.5003; Figs

19-21, Side, edge and view of other side (160), respectively. BMNH no. ZF 5002. Specimen from Brénnimann sample BR146, Acupe,

Brazil, for comparison.

Figs 22-24 Trochammina? milletti sp.nov. Figs 22,23, Detail of aperture (1,700) and side view (X320), respectively. Holotype, BMNH no.

1911.11.1.1088; Fig. 24, Side view (X260). Paratype, BMNH no. 1955.11.1.1089.

Figs 25-27 Paratrochammina simplissima (Cushman & McCulloch). Spiral, edge and umbilical views (X170). BMNH no. 1955.11.1.1141.

All from the Millett Collection, Malay Archipelago, except where stated.

TAXONOMIC REVISION OF FORAMINIFERA IN MILLETT COLLECTION

116

Ammotium pseudocassis (Cushman & Brénnimann,

1948) Figs. 1:5,50; 53

1899 Haplophragmium cassis (Parker); Millett: 359 (pars);

pl. 5, figs Sa,b only (non Lituola cassis Parker, 1870).

1948b Ammobaculites pseudocassis Cushman & Bronni-

mann: 39, 40; pl. 7, figs 12a,b.

1983. Ammoscalaria pseudospiralis (Williamson); Haman:

72; pl. 5, fig. 5 (non Proteonina pseudospiralis Will-

iamson, 1858).

REMARKS. This species was illustrated by Millett (1899, pl. 5,

figs 5a,b; here reproduced as Fig. 1.5a,b) under the name of

Haplophragmium cassis (Parker). It is an upper fragment of

an elongate test consisting of three rounded (in transverse

section), hardly compressed chambers. Millett’s drawing

shows the inward and backward trending sutures and the

rounded aperture in marginal position. Within the concept of

this species, Millett also included specimens of Ammotium

directum (Cushman & Bronnimann)(Figs 1.4a,b, 31,736) and

Ammoastuta salsa Cushman & Bronnimann (Figs 1.6a,b, 35).

As already mentioned, the fragment of A. pseudocassis

illustrated by Millett in his pl. 5, figs Sa,b (see our Figs

1.5a,b) could not be found in his Malay collection.

For comparison, we have illustrated typical specimens in

lateral view of A. pseudocassis from the mangrove sediments

of Guaratiba, Brazil (see Zaninetti et al., 1977), one by SEM

(Fig. 50), the other by optical photography in immersion

(Fig. 53). In the latter specimen the early spiral is reduced to

two chambers, a relatively large proloculus of 50 um diam-

eter, and a larger deuteroloculus of about 75 um diameter.

The embryonic chambers are not enclosed by other spiral

chambers, as in A. pseudospirale (Williamson, 1858). The

total number of chambers, including embryonics, is eight.

Ammotium pseudocassis differs from A. pseudospirale by

the elongate, somewhat incurved test, the less compressed

and elongate chambers and the reduced initial spire. The final

chamber is usually the dominant one, making up about

one-third of the test. It narrows toward the oblong aperture

and extends on the inner side of the test toward the initial

spiral. The early coil, represented by a reduced spire, consists

of very few chambers only. A typical embryo consists of two

very thin-walled chambers, a large proloculus, about

40-60 um in diameter, and an equally large deuteroloculus.

The embryo may form all the initial portion of the test. We

have never found a microspheric specimen of A. pseudocassis

and where the taxon is frequent, A. pseudospirale is usually

absent. The aperture of the holotype of A. pseudocassis,

deposited in the U.S. National Museum of Natural History

(registration no. 56764), is not as circular as that shown in

Millett’s drawing, but distinctly oblong. As in Ammotium

cassis and A. morenoi, the aperture is situated at the apex of

the last chamber in a marginal position (see Cushman &

P.BRONNIMANN AND J.E. WHITTAKER

Bronnimann, 19485, pl. 7, fig. 12b). However, it seems that,

when the final chamber is hardly compressed, the aperture

may become rather centred and more rounded than slit-like,

but never completely circular.

DIMENSIONS. Fig. 50: Height of test — 480 um; maximum

width — 150 um; height of final chamber — 290 um; maxi-

mum diameter of aperture — 50 um.

Fig. 52: Height of test — 575 um; maximum width — 170

um; height of final chamber — 375 um; maximum diameter

of aperture — 75 um; thickness of wall (final chamber)

10 um.

ENVIRONMENT. The group of forms referred by Millett to H.

cassis (Parker) occur only at Station 9, Area 1, . . . ‘where

they are not uncommon’. They are all exclusively brackish

water species.

Ammotium directum (Cushman & Bronnimann, 1948b)

Figs 1.4,(?1.7),31,36,37,45—47

1899 Haplophragmium cassis (Parker): 359 (pars); pl. 5,

figs 4,?7 only (non Lituola cassis Parker, 1870).

1948b Ammobaculites directus Cushman & Bronnimann:

38; pl. 7, figs 3,4.

1956 Ammotium sp. D. Warren: 139; pl. 1, figs 19-21.

1957 Ammobaculites directus Cushman & Broénnimann;

Todd & Brénnimann: 23; pl. 2, fig. 6 only (non Fig.

Dy:

1988 | Ammotium casamancensis (sic) Debenay: 46,47; pl.

1, figs 1-3.

REMARKS. Under the name of H. cassis (Parker), Millett

(1899, pl. 5, figs 4a,b; here reproduced as Figs 1.4a,b)

illustrated a specimen of Ammotium directum (Cushman &

Bronnimann). Our identification is based on the overall

outline, the shape of the sutures, and the strong lateral

compression of the test. The specimen illustrated by SEM in

Fig. 31 is that very same specimen, viewed from the other

side.

The fragment shown in Millett’s fig. 7, which he compared

to... ‘a species of Reophax, with the plan of growth and

chevron-shaped chambers of Frondicularia’ may be that

illustrated by SEM in Figs 36,45,46, although the chevron-

shaped chambers are exaggerated, as they are in the drawing

of fig. 4a (compare with our Fig. 31). The aperture in both

specimens is slit-like, without an everted border, and is

situated at the apex of the final chamber. For comparative

purposes, a lateral view of a specimen of A. directum, from

the mangrove sediments of Acupe, Brazil (BMNH no. ZF

4999) is illustrated in Fig. 47; the asymmetrical sutures are

well exhibited.

The tests of Ammotium directum in the Millett Collection

are extremely fragile, in contrast to those found in Trinidad

Figs 28-30 Trematophragmoides bruneiensis Bronnimann & Keij. Side, edge and view of other side (X115). BMNH no. 1955.11.1.1136.

Figs 31, 36,37 Ammotium directum (Cushman & Brénnimann). Side views of three separate specimens (X185, 205 and 185, respectively).

BMNH nos. 1955.11.1.1118-1120.

Figs 32-34 Ammotium morenoi (Acosta). Side, edge and oblique apertural views (X250). BMNH no. 1955.11.1.1122.

Fig. 35. Ammoastuta salsa Cushman & Bronnimann. Side view (X200). BMNH no. 1955.11.1.1121.

Figs 38-41

Bay, Gulf of St. Lawrence, Canada.

Ammotium cassis (Parker). Apertural, oblique apertural, side and edge views (X45). Lectotype, BMNH no. ZF 4637, Gaspé

Figs 42-44 ‘Ammobaculites’ exiguus Cushman & Brénnimann. Fig. 42, Side view (X185). BMNH no. 1955.11.1.1058; Figs 43,44,

Oblique-apertural and edge views (X175). BMNH no. 1955.11.1.1057.

All from Millett Collection, Malay Archipelago, except where stated.

jaa

fe)

—

aa)

—

fo)

fe)

118

or Brazil. Cushman & Brénnimann (1948)) distinguished two

species of Ammotium with strong lateral compression,

namely A. directum and A. diversum. To these has to be

added a third, A. subdirectum Warren, 1956.

Ammotium directum, the more common species, as

described above, has strongly incurved sutures of an asym-

metrical type with a shorter outer or marginal branch and a

longer, inner branch, which slants toward the initial spire (see

Fig. 47). Occasionally, some sort of chevron pattern is

formed but rarely to the extremes indicated by Millett’s

drawing (pl. 5, fig. 4a; our Fig. 1.4a). A. directum is always

characterized by this asymmetrical type of suture. The test,

moreover, is strongly compressed laterally and the width of

the flattened chambers does not increase much in the course

of growth. The aperture is slit-like and situated at the apex of

the final chamber, more or less in a marginal position. The

initial spiral consists of several chambers.

A. diversum is less common than A. directum. The only

significant difference lies in the sutural shape, which in the

former in the final ontogenetic stage, is always more or less

horizontal, slightly incurved and in extremes, no longer

asymmetrical (see Cushman & Bronnimann, 1948), pl. 7, figs

5,6). Furthermore, in this species, the sutures of the early

uniserial portion are slanting inward toward the initial coil.

Occasionally, there is a suggestion that the two are linked by

transitional forms. Should the two be ultimately considered

synonymous then we would prefer to retain A. directum, as

this, although printed on the same page, was described first.

For the time being, however, both are retained.

A. subdirectum was described by Warren (1957, pl. 4, figs

6-8) from the marshes of the Buras-Scofield bayou region of

southeastern Louisiana. We have encountered it but rarely in

the mangrove sediments of Acupe, Brazil and Warren him-

self (1957: 33) mentioned that ... ‘specimens were rare

wherever found except in one of the polyhaline marsh

samples’. Two specimens from Acupe (BMNH nos ZF 5000

and 5001) are illustrated in Figs 48,49,51,52 for comparison

with the Malay species of Ammotium. A. subdirectum is a

many-chambered species characterized by the same asym-

metrical type of sutures as found in A. directum. In the final

growth stages the sutures may become more or less symmetri-

cal and arranged in a chevron-like pattern, as shown in

Millett’s drawing of the fragment illustrated in pl. 5, figs 7a,b;

Fig. 1.7a,b. Figs 36,45,46 could represent this specimen

which is part of either an A. subdirectum, or an A. directum

as discussed above. Normally, A. subdirectum is about twice

as long as A. directum and composed of more chambers. The

test is slightly incurved and the width of the chambers, seen

laterally, increases quite strongly towards the final chamber.

The aperture is a narrow oblong slit, asin A. directum and A.

diversum, and situated at the apex of the final chamber, more

of less in a marginal position (see Figs 48,49,51,52). In all

three species, the agglutinant is fine-grained and the surface

of the test usually appears smooth, occasionally even some-

what glossy.

DIMENSIONS OF FIGURED SPECIMENS (BMNH no.

1955.11.1.1118). Height of test — 290 um; maximum width of

final chamber — 73 um.

(BMNH no. 1955.11.1.1119). Height of fragment —

270 um; length of apertural slit — 45 um.

(BMNH no. 1955.11.1.1120). Height of test — 330 um.

ENVIRONMENT. This species was found only at Station 9

P.BRONNIMANN AND J.E. WHITTAKER

(Area 1). It is a typical brackish-water species.

Genus ARENOPARRELLA Andersen, 1951la

TYPE SPECIES. Trochammina inflata (Montagu) var. mexi-

cana Kornfeld, 1931.

Arenoparrella mexicana (Kornfeld, 1931)

Figs 1.3, 9-12

1899 Haplophragmium agglutinans d’Orbigny var. triper-

forata Millett: 358 (pars); pl. 5, figs 3a,b only; non

figs 2a,b.

1931 Trochammina inflata (Montagu) var. mexicana Korn-

feld: 86; pl. 13, figs Sa—c.

Arenoparrella mexicana (Kornfeld); Andersen: 31;

fig. la—c.

Arenoparrella mexicana (Kornfeld); Andersen; 96;

pl. 11, figs 4a—c.

1977 Arenoparrella mexicana (Kornfeld); Zaninetti et al. ;

pl. 2ahies 3:7.

REMARKS. One of Millett’s illustrated specimens (1899, pl. 5,

figs 3a,b; reproduced here as Figs 1.3a,b) of H. agglutinans

var. triperforata is, in fact, a typical specimen of Arenopar-

rella mexicana (Kornfeld). It is refigured here by SEM (Figs

9-12) and shows that the original drawing of the edge view in

particular, is very misleading. As discussed above under the

description of the lectotype of Acupeina triperforata (Millett),

the edge view of fig. 3b also suggests that there are only three

large everted apertures. The reality is an aperture consisting

of a vertical slit lined by slightly uplifted borders, in an

interiomarginal position, of about 50 um length and 8 um

width, and 12 small, irregularly arranged, rounded pores

above this primary aperture, of between 5 and 10 um diam-

eter, devoid of everted rims. Millett’s specimen (BMNH no.

1955.1.1.1075) is tilted so far forward in apertural view that

the primary vertical slit, so clearly visible in fig. 3a, might not

have been seen, but it is puzzling to understand why he

illustrated the apertural pores as he did. Millett’s material

from stations 5 and 9 quite clearly represents both Acupeina

triperforata and Arenoparrella mexicana, which is not surpris-

ing as they commonly occur together. The illustrated speci-

men of the latter is completely involute (hence the small axial

depression is closed). The final whorl consists of 4, axially

compressed chambers which gradually increase in size with

growth. In edge view the periphery is rounded. Umbilical and

spiral sutures are poorly defined and the agglutination is

rather fine-grained and produces a smooth surface.

195la

1951b

DIMENSIONS OF FIGURED SPECIMEN (BMNH no.

1955.11.1.1075). Maximum diameter — 290 um; minimum

diameter — 240 um; axial height (thickness) — 120 um.

ENVIRONMENT. See under Acupeina triperforata (p._ ).

Arenoparella mexicana is a typical tropical and subtropical

mangrove swamp species.

Genus HAPLOPHRAGMOIDES Cushman, 1910

TYPE SPECIES. Nonionina canariensis d Orbigny, 1839.

REMARKS. The wall structure of the type species is unknown;

we are not even sure of the apertural position, for that

matter. Although the wall of H. wilberti, the species in the

TAXONOMIC REVISION OF FORAMINIFERA IN MILLETT COLLECTION

Millett Collection, is of the Trochamminina-type (see below),

we prefer, in our treatment of this species, to use Haplo-

phragmoides in inverted commas, until more is known about

canariensis.

‘Haplophragmoides’ wilberti Andersen, 1953

Figs 1.12, 16-21

1899 Trochammina ochracea Williamson; Millett: 363, pl.

5, figs 12a—c (non Williamson, 1858).

1953. Haplophragmoides wilberti Andersen: 21, pl. 4, figs

7a,b.

1961 Haplophragmoides wilberti Andersen; Todd & Low:

162 pl. fig. 5.

1973 Haplophragmoides wilberti Andersen; Haynes:

27-30, pl. 2, fig. 1; pl. 29, fig. 7?; text-figs 5.3-7.

1977 Haplophragmoides wilberti Andersen; Zaninetti et

ae: ple Uy ties 12513.

1981 Trochammina sp., Cann & de Deckker: 668, pl. 2,

figs 1-19.

1983. Haplophragmoides wilberti Andersen; Haman: 71;

pl. 3, figs 14,15.

REMARKS. On re-examination, Millett’s (1899, pl. 5, figs

12a—c; reproduced here as Figs 1.12a—c) so-called Trocham-

mina ochracea proved to be a planispiral ‘Haplophrag-

moides’. It is re-illustrated by SEM in Figs 16-18 and it

(BMNH no 1955.11.1.5003) clearly shows the same collapse

features as the original drawings. In addition to this speci-

men, we have illustrated for comparative purposes (Figs

19-21), another, somewhat less deformed specimen, from

Acupe, Brazil (BMNH no. ZF 5002).

The coiling of Millett’s species is planispiral, virtually

involute, with 7 or 8 chambers in the final whorl. The

aperture is a single interiomarginal equatorial slit with a

broad everted border. The intercameral sutures are incurved,

occasionally sinuous. We are placing it into the widespread

brackish form, ‘H’. wilberti Andersen. Millett must have

regarded it as a Trochammina because of the incurved test

seen in edge view.

Cann & de Deckker (1981, pl. 2, figs 1-19) illustrated from

ephemeral lakes adjacent to the Coorong Lagoon, South

Australia, a series of haplophragmoid forms, in part

deformed, which they called Trochammina sp. They are very

similar to T. ochracea sensu Millett and we have also placed

them in ‘H’. wilberti.

Collapse features occur often in brackish foraminifera. The

overall consistency of the agglutinated phase, in particular its

_ thickness and cementation, seems to play a role. In the

deformed Millett material it appears that the agglutinated

phase is rather weakly developed. In non-deformed speci-

mens of ‘H’. wilberti, at our disposal, from brackish localities

in Nigeria and New Guinea, the wall structure was analysed

using high-resolution scanning electron microscopy of frac-

tured tests. It was found that the wall of these specimens is

made up of the organic phase (represented by thin inner and

outer sheets and material (‘glue’) between agglutinated ele-

ments), and the agglutinated phase. There were no perfora-

tions nor alveolar pseudopores present. This is the

characteristic Trochamminina-type wall. In these latter, non-

deformed specimens, the agglutinated phase appears to be

stronger, perhaps better cemented, than the Millett material

from the Malay Archipelago.

DIMENSIONS OF FIGURED SPECIMENS (MALAY SPECIMEN,

119

BMNH no. 1955.11.1.5003). Maximum diameter — 490 um;

axial height (thickness) — 125 um.

(BRAZILIAN SPECIMEN, BMNH no. ZF 5002). Maxi-

mum diameter — 340 um.

ENVIRONMENT. According to Millett (1899; 363) this species

. ‘has been observed only at Station 3’. It is a good

brackish water indicator and occurs in association with Trem-

atophragmoides bruneiensis at this locality.

Genus PARATROCHAMMINA Bronnimann, 1979

TYPE SPECIES. Paratrochammina madeirae Bronnimann,

1979

Paratrochammina simplissima (Cushman & McCulloch,

1948) Figs 1.9, 25-27

1899 Haplophragmium nanum Brady; Millett: 360; pl. 5,

figs 9a—c (non Brady, 1881).

1939 Trochammina pacifica Cushman var. simplex Cush-

man & McCulloch: 104; pl. 11, fig. 4 (non Friedburg,

1902).

1948 Trochammina pacifica Cushman var. simplissima

Cushman & McCulloch: 76 (nomen novum).

1956 Trochammina simplissima Cushman & McCulloch;

Bandy: 198; pl. 29, figs 14a—c.

1979 Paratrochammina simplissima (Cushman & McCul-

loch); Bronnimann: 10; figs 2,3,6A—J,8A—H (q.v. for

full synonymy).

REMARKS. Millett’s illustrated specimen, attributed to H.

nanum Brady (op.cit. pl. 5, figs 9a—c; reproduced here as Figs

1.9a—c), is a sinistrally coiled specimen with 5 chambers in the

final whorl. From the drawings it can be seen that the

chambers of the final whorl are strongly compressed in an

axial direction and the ultimate chamber is radially elongate.

The intercameral sutures are well defined and the agglutinant

of the spiral side appears to be distinctly coarser than that of

the umbilical side. The aperture, which is an essential generic

criterion, is only visible in edge view and its umbilical

extension, if any, cannot be seen in the drawing of the

umbilical side. We have searched the Millett Collection to

find this figured specimen but the closest to it is a dextrally

coiled individual (Figs 25-27), so it is possible that Millett’s

drawings could be reversed. Our figured specimen is

undoubtedly Paratrochammina simplissima (Cushman &

McCulloch). The single umbilical aperture is an elongate

interiomarginal slit in the final septum, which extends from

the surface of the first chamber of the final whorl onto that of

the penultimate chamber. Its length is about 120 um and it is

lined by a weakly uplifted border of agglutinated fragments.

The final whorl has 5 chambers, as in the original drawing,

but the ultimate chamber, perhaps, is radially not as elongate

as in Millett’s figure. The test consists of 10 chambers, the

coiling is rather tight and the axial depression (umbilicus) is

therefore virtually closed. The radial sutures are well defined

on both sides and the outline of the test is weakly lobate; the

periphery, as seen in edge view, being compressed but still

rounded. The spiral side is almost flat and the umbilical side

slightly concave. As in Millett’s illustrated specimen, ours is

also more coarsely agglutinated on the spiral side than

umbilically.

The marine, shallow water P. simplissima differs in all

120

pertinent features (size, chamber inflation and shape, aper-

ture, etc.) from Brady’s deep water species Haplophragmium

(=Trochammina) nanum which was __lectotypified,

redescribed and illustrated by Bronnimann & Whittaker

(1980: 177, figs 1-9). P. simplissima is highly variable in the

overall shape and outline of the test (see Bronnimann, 1979:

14, figs 6A—J), however it is usually less compressed axially

than Millett’s specimens.

DIMENSIONS OF FIGURED SPECIMEN (BMNH no.

1955.11.1.1141). Maximum diameter — 370 um; minimum

diameter — 280 um; axial height (thickness) — 90 um.

ENVIRONMENT. According to Millett (1899: 360), this species

. ‘is most abundant in Area 1’. It is a marginal marine

species and significantly, was not listed where true brackish

species such as Acupeina triperforata, Ammoastuta salsa,

Arenoparrella mexicana, etc. were recorded.

Genus TREMATOPHRAGMOIDES Bronnimann & Keij,

1986

TYPE SPECIES. Trematophragmoides bruneiensis Bronnimann

& Keij, 1986.

REMARKS. The genera Haplophragmoides, Cribrostomoides,

and Discammina are all superficially similar to Tremato-

phragmoides. Trematophragmoides Bronnimann & Keij is

slightly evolute and planispiral with 3 apertures per chamber:

a single primary equatorial interiomarginal aperture and one

on each side of the chamber, umbilically situated on the

suture and posteriorly directed. Haplophragmoides Cushman

(1910) is also planispiral but has only one aperture per

chamber. Cribrostomoides Cushman (1910) is usually invo-

lute, with streptospiral coiling initially, becoming planispiral

in the adult whorls; the aperture is a equatorial, single areal

slit (with lip) near the base of the septal face, becoming

subdivided into a linear series of openings in gerontic forms.

Discammina Lacroix (1932) is planispiral and slightly evolute,

has a low interiomarginal equatorial aperture and is said to

have an... ‘interior divided by thin straight organic parti-

tions, not corresponding to the original apertural face and not

always reflected at the surface’ (Loeblich & Tappan, 1987:

68).

Trematophragmoides bruneiensis Br6nnimann & Keij,

1986 Figs 1.8, 28-30

1899 Haplophragmium compressum Goés; Millett: 359; pl.

5, figs 8a—c (non Lituolina irregularis var. compressa

Goés, 1882).

1986 Trematophragmoides bruneiensis Bronnimann &

P.BRONNIMANN AND J.E. WHITTAKER

Keij: 16; pl. 1, fig. 1-10, pl. 2, figs 3-5, pl. 10, figs

1-3, text-fig. 1.

REMARKS. Broénnimann & Keij (1986) described from brack-

ish waters of Brunei, NW Borneo, a planispiral agglutinated

foraminifer with an interiomarginal and equatorial primary

aperture and two secondary lateral apertures per chamber.

The test shape is quite variable and the periphery, as seen in

edge view, can be broadly rounded or compressed, almost

subcarinate. Millett’s roughly agglutinated species (1899, pl.

5, figs 8a—c; reproduced here as Fig. 1.8a—c), attributed by

him to Goés’ species H. compressum, shows two lateral

Openings and a single equatorial primary opening per cham-

ber, and is undoubtedly a Trematophragmoides. Our SEM

illustrations (Figs 28-30) show the same specimen as that

drawn by Millett. As the early radial sutures are indistinct,

the total number of chambers cannot be determined with

certainty; the final whorl, however, contains 6 chambers. The

lateral secondary apertures are well exhibited in Fig. 30 and

the equatorial primary aperture in Fig. 29. Although the

number of chambers in the final whorl is less than in the types

of T. bruneiensis, other features agree well and there is no

doubt that the two are one and the same.

DIMENSIONS OF FIGURED SPECIMEN (BMNH no.

1955.11.1.1136). Maximum diameter — 470 um; maximum

thickness (final chamber) — 200 um.

ENVIRONMENT. This species occurs only at Station 3. Millett

(1899) does not offer any information about its association

with other species, but from a study of his collection it can be

seen to occur with ‘Haplophragmoides’ wilberti. From this

information, Station 3 must have been a brackish locality.

Genus TROCHAMMINA Parker & Jones, 1859

TYPE SPECIES. Nautilus inflatus Montagu, 1808.

Figs 1.10,22-24

Haplophragmium anceps Brady; Millett: 361, pl. 5,

figs 10a,b (non Brady, 1884).

DIAGNOsIs. Small conical, tightly-coiled Trochammina? with

three large subglobular chambers in the final whorl.

Trochammina? milletti sp.nov.

1899

NAME. In honour of Fortescue William Millett.

HO.LotyPe. BMNH no. 1955.11.1.1088. Illustrated in Figs

22,23. This may be the specimen figured by Millett (1899, pl.

5, figs 10a,b; reproduced here as Figs 1.10a,b). From Station

12, Area 1.

DESCRIPTION (HOLOTYPES). Test free, dextrally coiled coni-

cal trochospire with pointed initial portion. Final volution

Figs 45-47 Ammotium directum (Cushman & Bronnimann). Figs 45,46, Detail of aperture (975 and X280, respectively). BMNH no.

1955.11.1.1119; Fig. 47, Side view (X160). BMNH no. ZF 4999, mangrove sediments, Bronnimann sample 93, Acupe, Brazil

Figs 48, 49, 51,52 Ammotium subdirectum Warren. Figs 48,49, Side and edge views (X85). BMNH no. ZF 5000; Figs 51,52, Side and

oblique-apertural views (X85 and 125, respectively). BMNH no. ZF 5001. Both from mangrove sediments, Bronnimann sample 93, Acupe,

Brazil.

Figs 50,53 Ammotium pseudocassis (Cushman & Bronnimann). Side view (X160) and separate specimen in clearing oil (265).

Bronnimann Collection, mangrove sediments, Guaratiba, Brazil.

Fig. 54

Ammotium morenoi (Acosta). Side view (X205). Bronnimann Collection, mangrove sediments, Guaratiba, Brazil.

Fig. 55 Ammotium distinctum (Cushman & Broénnimann). Side view in clearing oil (330). Brénnimann Collection, sample 145, from

Acupe, Brazil.

All from Millett Collection, Malay Archipelago, except where stated.

TAXONOMIC REVISION OF FORAMINIFERA IN MILLETT COLLECTION

122

triserial, consisting of large subglobular chambers, somewhat

compressed in axial direction, making up major part of the

test. Coiling tight and axial depression (umbilicus) closed.

Sutures well defined. Single interiomarginal aperture a small

arch resting with its slightly upturned border completely on

surface of first chamber of final whorl. Agglutinant rather

coarse.

DIMENSIONS (HOLOTYPE). Height of test — 160 um; width

(umbilical diameter) — 150 um.

REMARKS. Millett (op.cit., pl. 5,, figs 10a,b) attributed this

small, rather fragile, conical form, to Haplophragmoides

anceps Brady. The aperture is a broadly rounded interiomar-

ginal arch, sitting with its border completely on the final

whorl and therefore the species should belong to Trocham-

mina (see Bronnimann ef al., 1983). Fig. 23 illustrates a

typical specimen from the Millett Collection but there are

extremes (Paratype BMNH no. 1955.11.1.1088; Fig. 24)

where the height of the trochospire and the umbilical diam-

eter are about the same or the former even appears to be

slightly larger. In 1983, Bronnimann et al. held great store by

the fact that in the Trochamminacea the umbilical diameter

was invariably greater than the length of the axis of coiling

(height of the trochospire), whereas in the Ataxophragmia-

cea the reverse was true. This is the first time we have found a

species, and apparently a single population, at the borderline

of the two groups. For this reason we have only tentatively

placed this interesting species in Trochammina.

The true H. anceps Brady, 1884 is the type species of

Globotextularia Eimer & Fickert, 1899. This is a robust, deep

water form, much larger than Millett’s species, with a very

high, often irregular coil, an open umbilicus and larger

aperture.

ENVIRONMENT. According to Millett (1899: 361), . . . ‘speci-

mens [of ‘“H. anceps‘| are numerous and well distributed’.

They are found at stations 5,11,12 (Area 1) and 27,28 (Area

2). It is associated with the agglutinating foraminifera Ammo-

baculites exiguus at stations 12 and 27 and rare Acupeina

triperforata/Arenoparrella mexicana at Station 5. The former

is found in both marginal marine and brackish localities,

whereas the latter are true brackish forms. It is therefore not

known for certain whether 7.? milletti is a marine or a

brackish species.

Genus TRUNCULOCAVUS gen.nov.

TYPE SPECIES. Trunculocavus durrandi sp.nov.

DIAGNOsIS. Test free, initially biserial, then abruptly unise-

rial. Biserial chambers subglobular, uniserial chambers with

circular transverse section. Wall agglutinated, of

Trochamminina-type. Aperture single, terminal, circular and

large, devoid of everted border.

NAME. Derived from the Latin: cavus, a hole or hollow, and

trunculus, tip or end.

REMARKS. Our new genus has the basic morphology of

Bigenerina d’Orbigny, 1826 (type species B. nodosaria d’Or-

bigny, 1826), but differs in the large circular aperture of the

final chamber of the uniserial stage, devoid of a border

structure. In contrast, the terminal aperture of B. nodosaria

is a small central porus with everted border. According to

Loeblich & Tappan (1987: 172), Bigenerina also has a perfo-

P.BRONNIMANN AND J.E. WHITTAKER

rate (‘canaliculate’) wall, whereas Trunculocavus has a

Trochamminina-type wall.

In the Millett material, there are well-preserved specimens

of Trunculocavus durrandi showing an organic structure

within the large rounded aperture. This structure is different

from the inner organic sheet (inner organic lining in the sense

of Bender, 1989: 278), which occurs along the inside of the

wall of the Trochamminina, because it is independent of the

agglutinated-organic wall proper. It is suggested that it repre-

sents the epidermal layer of the protoplasmic body of the

foraminifer. Therefore, we must distinguish between this type

of organic structure, as part of the protoplast, and the inner

organic sheet which covers the inside of the agglutinated wall

of the Trochamminina-type test (see Bronnimann & Whit-

taker, 1988), which, although has also been produced by the

protoplast, is not directly part of it.

This organic structure, or the epidermal layer of the

protoplasmic body, occurs inside the terminal aperture of the

test, either as a large rounded opening limited by a thickened

border (Fig. 7), or it closes the aperture of the test completely

and reveals 6 small perforations with tube-like extensions

(Fig. 1) along the apertural periphery. This organic structure

does not have a counterpart in the agglutinated-organic phase

of the wall, another reason for separating it nomenclatorally

from the inner organic sheet. In fossil specimens, the epider-

mal layer of the protoplast will naturally be absent, so it could

not be considered taxonomically. At present, therefore, it has

no standing in the systematic treatment of these agglutinated

foraminifera, which is based on test features alone. It should,

however, be remembered that this situation would have to be

modified once it becomes possible to take into consideration

the features of the living organism.

In a paper by Petrucci et al. (1983: 72-75), there is a

taxonomic appendix by Medioli, Scott & Petrucci. In this

appendix a new species, Polysaccammina hyperhalina, is

introduced which is of interest here because it shows organic

features similar to those described for 7. durrandi. P. hyper-

halina has a large circular terminal aperture with an irregu-

larly finished border, devoid of particular border structures.

Medioli et al. (1983: 72, pl. 21, figs 2,3,6,8) described the

aperture as invaginated ... ‘to form an inner, backward

pointing funnel’. Their pl. 1, figs la,2a show the large,

rounded aperture is closed on the inside, as in 7. durrandi, by

an organic structure having in its centre a small circular porus

with everted border. Also, as in 7. durrandi, this small

opening appears to be a different from the aperture of the test

and that it represents the epidermal layer of the protoplast,

with features which have no counterpart in those of the test

wall and which is different from the inner organic lining.

Figs 2.1,3-8

1900 Bigenerina digitata d’Orbigny var. Millett: 6, pl. 1,

figs la,b (non Bigenerina (Gemmulina) digitata d’Or-

bigny, 1826).

DIAGNOsIs. As for genus; Trunculocavus is presently mono-

typic.

NAME. In honour if A. Durrand FRMS, the collector of the

Malay Archipelago foraminifera described by Millett.

HOLOTYPE. BMNH no. 1955.11.1.187. Illustrated in Figs

3,4. From Station 9, Area 1.

Trunculocavus durrandi sp.nov.

DESCRIPTION (HOLOTYPE. Test free, small and elongate; ini-

TAXONOMIC REVISION OF FORAMINIFERA IN MILLETT COLLECTION 123

tially a subglobular protoconch, followed by 4 pairs of

subglobular, biserial chambers, then abruptly uniserial with 3

cylindrical chambers. Aperture large, terminal and central

without everted border; the inner organic sheet closing the

aperture, however, develops around its circumference, 6

minute pores with tubular borders. Agglutinated wall of

granular, but overall smooth appearance. Tubular organic

pores have no counterpart in agglutinated phase.

DIMENSIONS (HOLOTYPE). Height of test — 270 um; maxi-

mum width of test — 75 um; diameter of aperture — 35 um;

diameters of tubular pores around circumference of aperture

— 45 um.

PARATYPES. Two paratypes (BMNH nos. 1955.11.1.188,189)

are illustrated in Figs 5-8. In side view, they are as the

holotype with a short biserial stage followed by the uniserial

stage composed of 2 or 3 cylindrical chambers. Paratype

(BMNH no. 1955.11.1.188; Figs 6-8) shows an aperture

where the inner organic sheet does not close the opening. The

sheet itself has an opening, bordered by a thickened rim,

which is virtually of the same diameter as the rounded

terminal aperture of the agglutinated phase; there are no

minute pores as in the holotype.

DIMENSIONS (PARATYPES). (BMNH_ no. 1955.11.1.188)

Height of test — 230 um; maximum width — 90 um; diam-

eter of aperture — 40 um.

(BMNH no. 1955.11.1.189) Height of test — 240 um;

maximum width 75 um.

REMARKS. Millett’s actual figured specimen (1900, pl.1, figs

la,b; reproduced here as Figs 2.1a,b) could not be recognised

with certainty. Millett’s drawing, however, shows a specimen

with a rather indistinct biserial initial portion of 4 or 5 pairs of

chambers, then a 4 or S-chambered uniserial stage; the

uniserial chambers are cylindrical and the large terminal

rounded aperture is devoid of an everted border.

ENVIRONMENT. According to Millett (1900: 6), this species is

‘confined to Station 9, and the examples, although

minute, are moderately abundant’. From the same locality

Millett (1899: 358,359) also found Acupeina triperforata,

Arenoparrella mexicana, ‘Ammobaculites’ exiguus, Ammoas-

tuta salsa and Ammotium spp., all brackish, mangrove

sediment-dwelling species. It is therefore assumed that 7.

durrandi also lives in a brackish habitat.

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Bull. nat. Hist. Mus. Lond. (Zool.) 59(2): 125-170 Issued 25 November 1993

Foregut anatomy, feeding mechanisms,

relationships and classification of the

Conoidea (= Toxoglossa) (Gastropoda)

JOHN D. TAYLOR

Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD

YURI I. KANTOR

A.N. Severtzov Institute of Animal Evolutionary Morphology and Ecology, Russian Academy of

Sciences, Lenin Avenue 33, Moscow 117071

ALEXANDER V. SYSOEV

Institute of Parasitology, Russian Academy of Sciences, Lenin Avenue 33, Moscow 117071

CONTENTS

TE Gt HRN GEN ODE Mea ete ale IR aah nae ai alae ele ain sl niglata ara ic Maca ce seiesacrcins slDS Wsinaltn ds ad dda oiuaiisiadiaciladasine Splaaaree 125

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Functional morphology of the digestive system and feeding mechanisms in the Conoidea ......................00665 145

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Appendix 1. Description of shell characters used in phylogenetic amalysis. ................0ccececnceececneneceeeenenenes 162

NONE U CIN © LASSINCAIOMOM NCCOLU MEIC PCHOTA ..cccsscsnenctecsededscocnstcscstevcsteerscescasecscasoctscnsctcatudvens ceeds 163

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Synopsis. A survey of the anterior alimentary system of species from all the higher taxa of the highly diverse

/ gastropod superfamily Conoidea (including the Turridae, Terebridae, and Conidae) has revealed a great variety of

foregut structure. A series of anatomical characters of the rhynchodeum, proboscis, buccal mass, radular apparatus

and foregut glands has been defined and their distribution established amongst the various conoidean families and

sub-families. Twelve major types of foregut structure were recognised, which ranged from gastropods with a full set

of foregut organs and glands to others in which most of the structures including the radula, venom gland and

proboscis are absent. A set of these anatomical characters together with a few shell characters were used in a

cladistic analysis attempting to determine relationships amongst the conoidean higher taxa. A classification

incorporating the new anatomical data and based partly upon the phylogenetic analysis recognises 6 families and 13

subfamilies of Conoidea. New data suggest that the Pervicaciinae and Terebrinae share a common ancestor and

there is little evidence to justify familial separation of the Coninae. Some major foregut structures seem to have

evolved independently in different clades. Thus, hollow ‘hypodermic’ radular teeth have been derived indepen-

dently in a least five clades; the radular caecum and rhynchodeal introvert have evolved independently in two clades.

Several clades also show loss of major foregut structures such as the proboscis, venom gland and radular apparatus.

Finally, the 378 genera of Recent “Turridae’ are placed into the higher taxa recognised in the proposed classification.

eee

INTRODUCTION

The prosobranch gastropod superfamily Conoidea (=Toxo-

glossa), which includes the families Turridae, Conidae, Per-

vicaciidae and Terebridae, is extremely diverse, with as many

Fe 679 genera and 10,000 living and fossil species claimed for

the Turridae alone (Bouchet, 1990) and Conus with around

300 living species, is considered to be the most diverse genus

of marine animal (Kohn, 1990). Current classifications of

‘axa within the Conoidea are based almost entirely upon shell

characters, or upon a combination of shell and radular

characters (Turridae—Powell, 1966; McLean 1971; Kilburn,

1983, 1985, 1986, 1988; Terebridae—Bratcher & Cerno-

horsky, 1987). The Turridae are the most morphologically

disparate of the four families with seventeen subfamilies in

current use. However, most of these subfamilies are rather

poorly defined. Despite the biological interest in the venom

apparatus of the group, little is known of the relationships of

the Conoidea to other gastropods, of relationships between

the families of the Conoidea or of relationships within the

constituent families.

126

The Conoidea are considered to be monophyletic, because

the families share the common apomorphy of a venom

apparatus con-sisting of the venom gland and muscular bulb.

This is thought to have been lost in some taxa, such as some

highly-derived members of the Daphnellinae and Terebridae

(Kantor & Sysoev 1989; Taylor, 1990) and all Strictispirinae.

Compared with the number of living species and the

attention paid to the description of shells, particularly of

Conidae, there have been very few anatomical studies of

Conoidea. However, recently, a much wider range of species

from the Turridae, Terebridae and Pervicaciidae (Sysoev &

Kantor 1987, 1988, 1989; Kantor & Sysoev, 1989; Miller,

1989, 1990; Kantor, 1990; Taylor, 1990) have been investi-

gated anatomically. These studies illustrate the great variety

of foregut anatomy, particularly within the Turridae and

Terebridae. By comparison, the Conidae appear to have a

relatively uniform foregut anatomy (Marsh, 1971; Miller,

1989), although they have been surprisingly little studied.

Until recently, attempts to use anatomical characters in

determining relationships amongst conoideans were con-

strained either by the limited range of taxa that had been

studied or by the small number of characters used. For

example, an evolutionary scenario for the Conoidea based

upon characters of foregut anatomy was proposed by Sheri-

dan et al. (1973), but species were studied from only three out

of the seventeen turrid subfamilies. Additionally, Shimek &

Kohn (1981) used only radular characters to produce a

cladistic analysis of a wider range turrid taxa.

Another problem in comparing the different taxa studied

within the Conoidea, is that the nomenclature for the differ-

ent anatomical structures is inconsistent and very confused.

This has hampered the recognition of homologous structures

that may be shared between the different taxa.

In this paper we attempt a comparative review of the

anatomy and functional morphology of the conoidean

foregut. We have attempted to examine species from all the

currently-recognised subfamilies of Turridae, many species of

Terebridae, Pervicaciidae and a few species of Conus. Addi-

tionally, we have incorporated previously published studies

into our review and attempted to standardize the nomencla-

ture of the anatomical structures.

The overall objectives of the study are, firstly, to evalu-ate

the use of characters of foregut anatomy in determining

relationships among the Conoidea and secondly, to propose a

new classification of conoidean higher taxa which incorpo-

rates these anatomical characters. Foregut anatomy was

chosen as the focus for this study, because a few previous

studies (Sheridan et al. 1973; Kantor, 1990) had drawn

attention to the diversity and complexity of the digestive

system. As far as is known, other organ systems are similar to

other neogastropods.

MATERIAL AND METHODS

The material on which this study is based consists mainly of

longitudinal serial sections of the foreguts of a wide range of

gastropods from all of the currently recognised subfamilies of

Turridae, many Terebridae and Pervicaciidae and a few

species of Conidae (Table 1). Dissections were also made of

most of these species. Also indicated in Table 1 are species

for which we have used previously published anatomical

J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV

descriptions in our analysis. Additionally, radular prepara-

tions were made from a range of other species.

Critical-point dried preparations for scanning electron

microscopy were made of some anatomical structures and

some small species (methods in Taylor & Miller, 1989).

Radula preparations for both light and scanning microscopy

were made by standard methods.

FOREGUT ANATOMY

A generalized diagram of the conoidean foregut (Fig. 1)

shows the relative positions of the major structures.

rstm

Fig. 1 Composite diagram of the foregut of a hypothetical

conoidean gastropod showing the location of the major structures

discussed in the text. No single gastropod possesses all these

features. Abbreviations: as, anterior sphincter of buccal tube; bl,

buccal lips; bm, buccal mass; bt, buccal tube; is, intermediate

sphnicter of buccal tube; m, mouth; mb; muscular bulb; oe,

oesophagus; p, proboscis; rcoel, rhynchocoel; rs, radular sac; rsp,

rhynchostomal sphincter; rstm, rhynchostome; rw, rhynchodeal

wall; s, septum; sg, salivary gland; tm, transverse muscles of

rhynchodeal wall (shown in part only); vg, venom gland.

FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA

127

Table 1. List of species examined in this study. The classification in the list is traditional and follows Powell (1966), McLean (1971) and

Kilburn (1983-89). A new classification is given at the end of this paper. The prefix ‘a’ denotes species that were studied anatomically and

the prefix ‘r’ denotes species for which only the radula was examined. In most cases, animals were both dissected and serial sections made

of the anterior alimentary systems.

Pseudomelatominae

aPseudomelatoma penicillata (Carpenter, 1864). Punta San

Bartoleme, Mexico.

aHormospira maculosa (Sowerby, 1834). Sonora, Mexico

Drilliinae

aClavus unizonalis (Lamarck, 1822). Lizard I., Queensland,

Australia.

aClavus sp. (undescribed species). Guam.

aSplendrillia chathamensis Sysoev & Kantor, 1989. Chatham Rise,

South Pacific.

rDrillia cydia (Bartsch, 1943). British Virgin Islands.

rDrillia rosacea (Reeve, 1845). West Africa.

tIlmaclava unimaculata (Sowerby, 1834). Baja California, Mexico.

rSpirotropis monterosatoi (Locard, 1897). East Atlantic.

tCrassopleura maravignae (Bivona, 1838). Naples, Italy.

Clavatulinae

aToxoclionella tumida (Sowerby, 1870). South Africa.

aClionella sinuata (Born, 1778). Oudekraal, South Africa.

aClavatula caerulea (Weinkauff, 1875). Sierra Leone, West Africa.

aClavatula muricata (Lamarck, 1822). Dakar, Senegal.

Turrinae

rLophiotoma acuta (Perry, 1811). Lizard I., Queensland, Australia.

aGemmula deshayesi (Doumet, 1839). Hong Kong.

rGemmula kieneri (Doumet, 1840). Hong Kong.

aLophiotoma leucotropis (Adams & Reeve, 1850). Hong Kong.

aPolystira albida (Perry, 1811). Caribbean. Data from Leviten

(1970).

Cochlespirinae

aTurricula javana (Linnaeus, 1767). Hong Kong.

aTurricula nelliae spurius (Hedley, 1922). Hong Kong.

aAforia abyssalis Sysoev & Kantor, 1987. North-East Pacific.

aA foria lepta (Watson, 1881). South Pacific, nr New Zealand.

aA foria inoperculata Sysoev & Kantor, 1988. North-East Pacific.

alrenosyrinx hypomela (Dall, 1889). East Atlantic.

aAntiplanes sanctiioannis (Smith, 1875). Okhotsk Sea.

rAntiplanes vinosa (Dall, 1874). Sakhalin Bay, Okhotsk Sea.

Crassispirinae

rAustrodrillia angasi (Crosse, 1863). Sydney, Australia.

aFuna latisinuata (Smith, 1877). Hong Kong.

alnquisitor spp. Indian Ocean.

aVexitomina garrardi (Laseron, 1954). Sydney, Australia.

rPtychobela griffithi (Gray, 1834). Karachi.

Strictispirinae

aStrictispira paxillus (Reeve, 1845). British Virgin Islands.

rStrictispira stillmani Shasky, 1971. Panama.

rCleospira ochsneri (Hertlein & Strong, 1849). Galapagos Islands.

Zonulispirinae

aPilsbryspira nympha (Pilsbry & Lowe, 1932). Sonora, Mexico.

Borsoniinae including Mitrolumninae (fide Kilburn, 1986)

aLovellona atramentosa (Reeve, 1849). Guam.

aAnarithma metula (Hinds, 1843). Indian Ocean.

aBorsonia ochraea Thiele, 1925. Indian Ocean, nr Zanzibar 740m.

aMicanthapex parengonius (Dell, 1956). South Pacific, nr New

Zealand.

aTomopleura reevei (C.B. Adams, 1850). Indian Ocean.

aSuavodrillia kennicotti (Dall, 1871). Japan Sea.

aTropidoturris anaglypta Kilburn 1986. Southern Indian Ocean.

aTropidoturris fossata notialis Kilburn, 1986. South Africa.

aOphiodermella inermis (Hinds, 1843). Bremerton, Washington.

aOphiodermella ogurana (Yokoyama, 1922). Japan Sea.

Clathurellinae

aGlyphostoma candida (Hinds, 1843). Sonora, Mexico.

Mangeliinae

aMangelia brachystoma (Philippi, 1844). Galway, Ireland.

aMangelia nebula (Montagu, 1803). Galway, Ireland. Also data

from Sheridan et al. (1973) & Delaunois & Sheridan (1989).

aMangelia powisiana (Dautzenberg, 1887). Plymouth, England.

aEucithara stromboides (Reeve, 1846). Guam.

aHemilienardia malleti (Recluz, 1852). Guam.

aParamontana cf. rufozonata (Angas, 1877). Rottnest I., Western

Australia.

Oenopotinae

aOenopota levidensis (Dall, 1919). Washington. Data from Shimek

(1975)

tPropebela rugulata (Moller, 1866). White Sea.

Daphnellinae

aComarmondia gracilis (Montagu, 1803). Brittany, France. Data

from Sheridan et al. (1973)

aDaphnella reeveana (Deshayes, 1863). Guam.

aGymnobela emertoni (Verrill & Smith, 1884). Eastern Atlantic

Ocean.

aTeretiopsis levicarinatus Kantor & Sysoev, 1989. Eastern Atlantic

Ocean.

aA byssobela atoxica Kantor & Sysoev, 1986. Northern Pacific

Ocean.

aGymnobela latistriata Kantor & Sysoev, 1986. Northern Pacific

Ocean.

aGymnobela oculifera Kantor & Sysoev, 1986. Northern Pacific

Ocean.

aPontiothauma abyssicola Smith, 1895. Indian Ocean. Data from

Pace (1901).

aPontiothauma mirabile Smith, 1895. Indian Ocean. Data from Pace

(1901)

Conorbinae

aBenthofascis biconica (Hedley, 1903). Sydney, Australia.

aGenota mitraeformis (Woods, 1828). West Africa.

aGenota nicklesi Knudsen, 1952. West Africa.

Thatcheriinae

aThatcheria mirabilis Angas, 1877. North Western Australia.

Taraniinae

aTaranis moerchi (Malm, 1861). Sweden.

Conidae

aConus flavidus Lamarck, 1810. Queensland, Australia, Data from

Marsh (1971)

aConus ventricosus Gmelin, 1791. Tunisia.

Pervicaciidae

aPervicacia capensis (Smith, 1873). South Africa.

aPervicacia kieneri (Deshayes, 1859) Albany, Western Australia.

aPervicacia tristis (Deshayes, 1859). New Zealand.

aDuplicaria colorata Bratcher, 1988. Western Australia.

aDuplicaria duplicata (Linnaeus, 1758). Kenya.

aDuplicaria spectabilis (Hinds, 1844). Hong Kong.

a‘Terebra’ nassoides Hinds, 1844. Oman.

Terebridae

aHastula aciculina (Lamarck, 1822). Ghana.

aHastula bacillus (Deshayes, 1859). Phuket, Thailand.

aTerebra affinis Gray, 1834. Guam.

aTerebra babylonia Lamarck, 1822. Guam.

aTerebra gouldi Deshayes, 1857. Hawaii.

aTerebra maculata Linnaeus, 1758. Guam.

aTerebra subulata Linnaeus, 1767. Maldives.

128

Characters of the rhynchocoel

In all toxoglossans there is a permanent cavity in the anterior

part of the body called the rhynchodeal cavity or rhynchocoel

(Fig. 1). It contains the proboscis and is maintained even

when the proboscis is extended. The rhynchodeal cavity

opens to the exterior via the rhynchostome, which is situated

at the ventral margin of the head. The walls of the rhynchoc-

oel (rhynchodeum) are usually thick and muscular.

Rhynchostomal sphincter

This an annular, muscular sphincter which encircles the

mouth of the rhynchocoel (Fig. 1). It is present in most

species of Turridae, Terebridae, Pervicaciidae and Conidae,

but absent in the turrids Clavatula diadema and Tomopleura

violacea and the pervicaciids Pervicacia tristis, ‘Terebra’ nas-

soides, and ‘T.’ capen-sis. In these latter pervicaciids and

some turrids without a prominent sphincter, for example

Tomopleura, the anterior part of the rhynchodeum is very

muscular.

Position of rhynchostomal sphincter

In the normal condition, the sphincter is usually situated

around the rhynchostome, but in some turrids (for example in

Glyphostoma, Borsonia, Lophiotoma, Pontiothauma and

Thatcheria) it is situated more posteriorly. In Ophiodermella

inermis (but not O. ogurana) and Suavodrillia kennicotti the

moderately large, posteriorly situated, rhynchostomal sphinc-

ter is probably able to evert, forming a sort of ‘rhynchostomal

introvert’ but situated in the middle part of the rhynchocoel

(Fig. 2). The ability to evert is indicated by the presence of a

well-developed layer of longitudinal muscles underlying the

epithelium and by the existence of free space between the

sphincter and the longitudinal muscle layers. This structure

may demonstrate the possible origin of the true rhynchodeal

introvert (see below) or alternatively be an autapomorphy for

the species.

Rostrum

In the some fish-feeding species of Conus, the anterior part of

the rhynchocoel is elastic and can be greatly extended to

accomodate large food items during preliminary digestion.

This extensible feature, known as the rostrum, cannot be

inverted into the rhynchocoel.

Rhynchodeal introvert (= labial tube or

pseudoproboscis)

In this structure, the rhynchostomal lips are mobile and can

be retracted into the rhynchocoel by infolding, or extended as

a tube (Figs 3 & 4). The introvert is found in nearly all the

species which we and others have studied from the turrid

sub-family Daphnellinae, e.g. Philbertia linearis, P. leufroyi,

P. gracilis, Cenodagreutes, Daphnella reeveana (Smith, 1967;

Sheridan er al., 1973; unpublished observations), in Hemi-

lienardia malleti (Mangeliinae) and in all Terebridae and

Pervicaciidae (Miller, 1975, 1980; Taylor, 1990). We have not

seen an introvert in any other subfamily of Turridae (except

perhaps for Ophiodermella, see above), or in the Conidae. In

species of Daphnellinae the introvert is fairly short, but in

some terebrids, for example Terebra maculata, the introvert

J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV

0-5mm

Fig. 2 Ophiodermella inermis; longitudinal section of the anterior

rhynchodaeum showing the posteriorly-situated, rhynchostomal

sphincter located on an introvert-like structure. Abbreviations: in,

introvert; m, mouth; p, proboscis; r, rhynchostome; s, sphincter.

Fig. 3. Hemilienardia malleti; extended rhynchodeal introvert,

forming a pseudoproboscis in a relaxed, critical-point dried

specimen. Scale bar = 100 um.

is very long, and when retracted, lies coiled in the rhynchoc-

oel (Miller, 1970).

In those animals possessing a rhynchodeal introvert, the

outer and inner walls are joined by radial muscles (Fig. 5). In

Turridae, the possession of an introvert is associated with a

reduction in size or complete loss of the proboscis. However.

within the Terebridae, even those species with a well-

developed proboscis possess an introvert.

Epithelium of the rhynchodeum

In some Turridae, there is a distinct division in the character

of the epithelium lining the inner wall of the rhynchocoel. In

the anterior part of the cavity the epithelial cells are high and

FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA

“ey

ee SS

\\ yo > ait,

WPAN

aq!

oDecsag,

= once anne,

Fig. 4 Daphnella reeveana; A, longitudinal section through the

foregut; B, Enlargement of the mouth area showing the short

proboscis lying behind the septum. Abbreviations: bm, buccal

mass; cm, columellar muscle; con, circum-oral nerve ring; in,

rhynchodeal introvert; oe, oesophagus; ors, opening of radular

sac; Ovg, opening of venom gland; p, proboscis;s, rhynchostomal

sphincter; spt, septum; vg, venom gland.

glandular (Fig. 6C), but in the posterior half the epithelium is

low, cuticularized and similar in morphology to that of the

outer surface of the proboscis. This feature indicates that the

posterior part of the rhynchodeum can be extended outwards

when the proboscis is protruded through the rhynchostome.

We have observed this condition of the rhynchocoel epithe-

lium in Clavatula, and Clionella (Clavatulinae), Vexitomina

(Crassispirinae), Turricula nelliae spurius (Cochlespirinae),

Pilsbryspira nympha (Zonulispirinae). and Anarithma metula

(Borsoniinae).

In ‘lower’ turrids, excepting Vexitomina, this feature seems

to associated with those species in which the buccal mass lies

in a distal position within the proboscis (see below). Its

presence may be connected with the mechanism by which the

buccal mass is everted from the proboscis tip.

| Septum in rhynchodeum

A thin, slightly muscular septum, pierced by a circular orifice,

and dividing the rhynchodeal cavity into two parts is known in

Daphnella reeveana (Fig. 4), Philbertia purpurea (Sheridan et

al., 1973) and Terebra subulata (Taylor, 1990). The probos-

129

cis, when withdrawn, lies behind the septum, with the

retracted introvert lying to the anterior. A probably homolo-

gous septum is also found at the extreme posterior and

ventral end of the rhynchocoel in Thatcheria and Pontio-

thauma (Pace, 1901). A thin septum is also found in the

posterior part of the rhynchocoel in Pervicacia tristis (not

reported by Rudman (1969)) and in Duplicaria kieneri (Tay-

lor, unpublished).

The function of the septum is unknown, but it appears

better developed in species with a long proboscis and where

the proboscis withdraws behind the septum.

Accessory proboscis structure

This is an extensible muscular structure which arises from the

left hand wall of the rhynchocoel. It has been found only a

few species of Terebridae and Pervicaciidae. It is long and

branched in Hastula bacillus (Taylor & Miller, 1990), shorter

and club-like in Terebra affinis (Miller, 1971), ‘Hastula’

colorata and D. kieneri and a curved, club-shape in Terebra

imitatrix (Auffenberg & Lee, 1988). In H. bacillus the

accessory proboscis is covered in possible chemosensory

structures (Taylor & Miller (1990).

Snout gland

This is a subspherical gland which opens into the right-hand

posterior end of the rhynchocoel in a number of Conus

species (Marsh, 1971). The gland consists of folded glandular

epithelium (Fig. 7) and is surrounded by a muscular sheath of

circular muscles. From histochemical tests, Marsh (1971)

concluded that the gland secretes mucus. The gland has been

reported in 18 species of Conus, all but one of which are

known to be vermivorous (Marsh, 1971).

The proboscis and its structures

An extensible proboscis arising from the posterior of the

rhynchocoel is present in the Drilliinae (formerly Clavinae;

ICZN decision pending on further name change to Clavusi-

nae) and all the radulate turrids examined, excepting Gymno-

bela emertoni, where the radula is vestigial. A proboscis is

present in all species of Conus, in Hastula, and in other

radulate Terebridae, such as T. subulata, and T. babylonia

(Taylor, 1990). The distal opening to the proboscis forms the

true mouth as in all probosciferous gastropods. Shimek

(1975) referred to the opening of the buccal cavity as being

the true mouth.

A proboscis is absent in the radula-less Turridae such as

Teretiopsis, Taranis (Kantor & Sysoev, 1989), Philbertia

leufroyi boothi, P. linearis (Smith, 1967, Sheridan et al. , 1973)

and the radulate Gymnobela emertoni. A proboscis is also

absent in species of Duplicaria and Pervicacia, which are

radulate forms of the Pervicaciidae (Taylor, 1990), and in the

many species of Terebridae which lack a radula, such as

Terebra maculata, T. gouldi, T. dimidiata, and T. affinis

(Miller, 1970, 1975; Taylor, 1990).

In Duplicaria spectabilis and Gymnobela emertoni we have

observed a low cylinder of muscular tissue surrounding the

opening to the buccal cavity (Fig. 8) (Taylor (1990, Fig.2).

We think that this may represent the remnant of a much

reduced proboscis. A similar reduced structure found in

Cenodagreutes spp. and Philbertia linearis, was described by

Smith (1976) as the muscular sheath.

130

J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV

Fig. 5 Duplicaria spectabilis; relaxed, critical-point dried specimen. A, Section of the rhynchodeal wall showing the transverse muscles

joining the inner and outer walls. Scale bar = 100 um. B, Detail of junction of transverse muscles joining the inner wall of the

rhynchodaeum. Scale bar = 20 um.

Buccal tube

The buccal tube is that portion of the alimentary canal lying

between the buccal cavity and the true mouth, which is

situated at the distal end of the proboscis. The buccal tube is

present in all toxoglossans with a proboscis and is absent only

in those species where that organ is lost. It is very short in

Strictispira paxillus where the buccal mass lies at the extreme

anterior end of the proboscis.

In the Mangeliinae the epithelium of the buccal tube is very

thin (Fig. 9), but much thicker in species of other subfami-lies

such as the Drilliinae and Clavatulinae (Fig. 6). Shimek

(1975) refered to the buccal lips (see below) as the buccal

tube, and he called the true buccal tube, the inner proboscis

wall.

Buccal tube sphincters

In most toxoglossans, one or more annular sphincters may be

found in various positions within the proboscis.

a) Distal sphincter(s)

In most species with a proboscis, there is a distal sphincter

around the true mouth. Frequently, there is a second sphinc-

ter also near the proboscis tip, but located just to the

posterior of the first (Fig. 6). In ‘lower’ turrids such as the

Drilliinae Cochlespirinae and Clavatulinae, the sphincter(s)

grip the solid, radular teeth whilst they are held at the

proboscis tip (Sysoev & Kantor, 1989; Kantor & Taylor,

1991).

b) Intermediate sphincter

A small muscular sphincter, situated about half way along the

length of the proboscis is found in Splendrillia (Kantor &

Sysoev, 1989, fig. 3c). Species of Conus also have a sphincter

situated some distance posterior to the proboscis tip (Greene

& Kohn, 1989) which we classify as an intermediate sphinc-

ten:

c) Basal sphincter

A sphincter located near the base of the proboscis has been

described for Mangelia nebula (Sheridan et al., 1973).

Recently, Delaunois & Sheridan (1989) have illustrated a

section through the buccal area of M. nebula, showing a

single radular tooth held in the buccal tube. The tooth is

gripped at the anterior end by the buccal tube introvert (see

below), and the posterior end by the basal sphincter (Fig. 9).

Buccal tube introvert

This is a muscular, flap-like structure found towards the distal

end of the buccal tube of Mangelia nebula (Fig. 9) and called

a valve (valvule) by Sheridan et al. (1973). Eucithara strom-

boides has a longer, but apparently homologous structure

(Fig. 10). Delaunois & Sheridan (1989) showed that one of

the functions of this structure is to grip the radular tooth in

the buccal tube, but in Eucithara where the structure is very

long (Fig.10), it may possibly be used to transport teeth to the

proboscis tip.

Sen

FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA

rhs

Fig. 6 Clionella sinuata;A, longitudinal section through the

foregut; B, section of tip of proboscis showing sphincters; C,

section of portion of the inner wall of the rhynchodeum, showing

the differentiation in epithelium from that similar to the proboscis

wall, to that typical of the lining of the rhynchocoel.

Abbreviations: bm, buccal mass; bts, buccal tube sphincters; con,

circum-oral nerve ring; mb, muscular bulb; od, odontophore; oe,

oesophagus; p, proboscis; rhs, rhynchostomal sphincter; rs,

radular sac; sg, salivary gland; tec, tall epithelial cells; vg, venom

gland.

Sac-like enlargement of buccal tube

One other character associated with the gripping of marginal

teeth at the proboscis tip, is the presence of a sac-like

en-largement of the anterior or middle parts of the buccal

tube. It is found in different ‘lower’ turrids (Kantor & Taylor,

1991) as well as Mangelia nebula (Sheridan et al., 1973) and

_ Conidae (Conus catus (Greene & Kohn, 1989) and C. ventri-

cosus). Usually, the epithelium lining the enlargement is

formed of much taller cells than in the rest of the buccal tube.

These cells tightly surround the single radular teeth whilst

they are being held at the proboscis tip and may afford a

better grip. In Splendrillia chathamensis, Sysoev & Kantor

(1989) found the base of tooth adhering to a pad of epithelial

cells.

Protrusive lips of proboscis/ buccal tube

| In a few species, the inner lining of the outer lips of the

proboscis can be protruded. For example, in Turricula nelliae

spurius, the lips (Fig. 11) are densely covered by paddle or

| discocilia, which according to Haszprunar (1985) may indi-

cate the presence of chemosensory cells. Similar protrusible

lips are also found in Lophiotoma leucotropis and probably in

Aforia aulaca alaskana (Sysoev & Kantor, 1987).

In relaxed specimens of Mangelia powisiana, a sac consist-

ing of a single layer of cells is protruded from the proboscis

131

Fig. 7 Conus ventricosus; longitudinal section of the foregut

showing the proboscis retracted into the rhynchodeum.

Abbreviations: bm, buccal mass; bts, buccal tube sphincter; dasg,

duct of accessory salivary gland; fpw, fold of proboscis wall; ors,

opening of radular sac; ovg, opening of venom gland; p,

proboscis; rhs, rhynchostomal sphincter; sng, snout gland.

5mm

Fig. 8 Gymnobela emertoni; longitudinal section of the foregut

showing, the remnants of the proboscis, buccal lips and vestigial

radular sac. Abbreviations: bl, buccal lips; con, circum-oral nerve

ring; m, mouth; pr, reduced proboscis; rhs, rhynchostomal

sphincter; rm, radial muscles in rhynchodeal wall; rs, radular sac;

sg, salivary gland.

132

tip (Fig. 12). This sac is covered in granule-like structures

which are formed from single cells with large rounded nuclei.

The distinctive epithelial cells seen at the proboscis tip of

Mangelia nebula by Sheridan et al. (1973) may be the same

structure but in a more contracted position. The function of

this sac structure is not known.

Position of the buccal mass

Three conditions are known in the Conoidea;

a) Buccal mass situated at the base of the proboscis (Fig. 1)

For three reasons we consider this condition to be the

primitive state within the Conoidea. Firstly, a basal buccal

mass is found in species of the subfamily Drilliinae, which

with five teeth in each radula row, are considered to possess

the least-derived type of radula. Secondly, and also in the

Drilliinae, there is a muscular connection between the retrac-

tor muscle of the radular sac and the columellar muscle

(Kantor, 1990). This is a condition found in some meso- and

archaeogastropods, as for example in Littorina, Cryptonatica

and Tegula (Fretter & Graham, 1963; Kantor, unpublished

observations). In most other probosciform gastropods,

including those turrids where the buccal mass lies within the

proboscis, this connection is broken and the radula is con-

nected by muscles to the walls of the proboscis. Finally, the

basal buccal mass is a character-state shared amongst most of

the subfamilies of Turridae, along with the Terebridae,

Pervicaciidae and Conidae.

b) Buccal mass located within the proboscis

In Clavatula diadema (Clavatulinae), the buccal mass lies

within the proboscis, but in a proximal position (Kantor,

1990, fig. 8). In Clionella sinuata (Clavatulinae), Pilsbryspira

nympha (Zonulispirinae) and Funa latisinuata (Crassispiri-

nae), it lies more anteriorly, about half way along the

proboscis (Figs 6 & 14). In Strictispira paxillus (Strictispiri-

J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV

nae) (Fig. 13), Toxiclionella tumida (Clavatulinae) (Kantor,

1990 fig. 4), and Turricula nelliae spurius (Cochlespirinae)

(Taylor, 1985; Miller, 1990), the buccal mass lies in a distal

position near the tip of the proboscis.

The distally shifted position of the buccal mass in these few

turrids is a derived condition, being found only in some

species of the subfamilies Clavatulinae, Cochlespirinae

Zonulispirinae and Strictispirinae.

c) Buccal mass situated a long way to the posterior of the

proboscis base (Kantor, 1990, fig. 1).

This condition is found only in Hormospira (Pseudome-

latominae) and described by Kantor (1988).

Elongation of the oesophagus to the anterior of the

circum-oral nerve ring

In some turrids the oesophagus is elongated into a curved

loop between the base of the proboscis and the circum-oral

nerve ring (Fig. 14). This elongation is found in those turrids

with a buccal mass situated within the proboscis, and allows

forward movement of the buccal mass on protraction of the

proboscis. This condition is found in Clavatulinae, Stric-

tispirinae, Turricula nelliae spurius (Cochlespirinae), Cras-

sispitrinae such as Funa_ latisinuata, and _ Pilsbryspira

(Zonulispirinae).

Buccal lips (inner buccal tube)

These consist of muscular extensions of the anterior walls of

the buccal mass, which protrude as a tube into the lumen of

the buccal tube (Figs 1 & 9). In Oenopota levidensis where

the buccal lips are long (Shimek, 1975), they form a second

‘proboscis’ within the true proboscis. At full contraction of

the true proboscis, the tube formed by the buccal lips

protrudes through the mouth. Shimek (1975) called this

secondary ‘proboscis’ the buccal tube. Various developments

0-5mm

Fig. 9 Mangelia nebula; longitudinal section through the proboscis. A, with buccal lips protracted; B, radular tooth in proboscis and buccal

lips withdrawn into the buccal cavity. After Sheridan er al. (1973, fig. 7) & Delaunois & Sheridan (1989, plate II). Abbreviations: bc, buccal

cavity; bl, buccal lips; ds, distal sphincter of buccal tube; i, buccal tube introvert; m, mouth; ps, posterior sphincter of buccal tube; t,

radular tooth.

FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA

O-Imm

Fig. 10 Eucithara stromboides; longitudinal section through the

anterior end of the proboscis showing the buccal tube introvert.

Abbreviations: i, introvert; is, intermediate sphincter; m, mouth.

Fig. 11

Turricula nelliae spurius; extended proboscis, showing the

inner ring of the protrusive lips. Scale bar = 100 um.

of the buccal lips from a short tube to long proboscis-like

structures, are seen in species of the subfamily Mangeliinae.

Sections of Mangelia nebula (Sheridan et al. , 1973; Delaunois

& Sheridan, 1989) show that the buccal lips can be inverted

into the buccal cavity (Fig. 9b). In the genus Aforia (Cochle-

spirinae), some species have well developed buccal lips, but

in others they are absent (Sysoev & Kantor, 1987).

In some conoideans lacking a proboscis and radula (e.g.

Terebra gouldi (Miller, 1975)), the buccal lips are enlarged

and consist only of circular muscles. They have the appear-

ance of, and may be confused with, the true proboscis.

133

The buccal cavity and radular apparatus

From the true mouth, the buccal tube leads to a well-defined

chamber, the buccal cavity, which is surrounded by thick

walls of circular muscle. The radular diverticulum usually

opens ventrally into the buccal cavity. It consists of the

radular sac within which the radular teeth are formed, and in

less-derived turrids, an odontophore and sublingual pouch

(Fig. 15). The latter is the site where degeneration of the

radular teeth and ribbon occurs. The buccal sac is defined

(Shimek, 1976), as that part of the radular diverticulum that

lies between the buccal cavity and the entrance of the salivary

ducts.

In higher turrids without a radular membrane and odonto-

phore, the sublingual pouch is transformed into a caecum for

the storage of radular teeth prior to their use at the proboscis

tip.

Radula caecum (often called short arm of the radula

sac)

This is a diverticulum which branches off the anterior end of

the radular sac, in which detached radular teeth are stored

prior to their use at the proboscis tip (Fig. 15). We regard this

structure as a homologue of the sublingual pouch found in

other gastropods with a radular ribbon. A radular caecum is

present in higher turrids, for example the subfamilies Man-

geliidae, Daphnellinae, and Borsoniinae and also in Conidae

and some Terebridae.

Shimek (1976) showed that the caecum in Oenopota lev-

idensis is divided longitudinally by a septum. We have seen

this structure only in Micantapex parengonius (Borsoniinae).

Radular membrane

In general, the ‘lower’ turrids have a robust radular mem-

brane, whilst in ‘higher’ turrids, it is thin or absent. However,

even in ‘lower’ turrids, the strength of the membrane varies

considerably between taxa and we recognise only the pres-

ence or absence of the membrane as a functionally important

character.

A radular membrane is absent in the subfamilies Borsonii-

nae, Mangelinae, Daphnellinae, Conorbinae, Clathurelli-

nae, Taraniinae, Conidae and most Terebridae.

Odontophore

An odontophore with cartilages is present in many lower

turrids (Drilliinae, Pseudomelatominae, Strictispirinae, Clav-

atulinae, Turrinae, Cochlespirinae, Crassispirinae), the Per-

vicaciidae, and a few species of Hastula (Terebridae), but is

absent in higher turrids, Conidae and most other Terebridae.

If an odontophore is present, then the cartilages may be

either fused, or separated at the anterior end. If the cartilages

are separated, they are joined by a muscular connection. We

have seen fused odontophoral cartilages in Lophiotoma,

Pseudomelatominae, Splendrillia, Clavus sp., Inquisitor and

Funa spp., Toxiclionella and some Aforia species. Two

separate cartilages are usually present in species of Clavatuli-

nae (except Toxiclionella), Strictispira paxillus (Strictispiri-

nae) (Fig. 13). In Aforia lepta (Cochlespirinae), only the

muscle is present, over which the radular membrane bends

(Sysoev & Kantor, 1988).

134 J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV

Fig. 12 Mangelia powisiana; a, relaxed specimen showing sac-like structure at distal end of extended proboscis. Scale bar = 100 um. b,

detail of sac body with warty external surface. Scale bar = 100 um. c, section of the sac showing the thin epithelium with granule structures

produced by single cells with large nuclei. Scale bar = 50 um. d, detail of c. Scale bar = 10 um.

FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA

Radula

The radula has been by far the most studied of the organs of

the foregut and there are many published illustrations of

conoidean radular teeth (e.g. Powell, 1966; McLean, 1971;

James, 1980; Bandel, 1984; Bogdanov, 1990; Nybakken, 1990

and Taylor, 1990). Shimek & Kohn (1981) classified turrid

radulae into a number of functional groups and attempted a

cladistic analysis of radular characters. However, amongst the

‘lower’ turrids there is little evidence from direct observations

to support their functional categories. Indeed, recent evi-

dence shows that even in the least-derived radulae which

possess a strong radular ribbon, the marginal teeth can be

held singly at the proboscis tip in a stabbing position (Kantor

& Taylor, 1991).

A radula is present in most Turridae, all Conidae, possibly

all Pervicaciidae and some Terebridae. It is absent in some

species of Daphnellinae, Taraninae and many species of

Terebridae (Miller, 1970; Taylor, 1990). The phenomenon of

radula-loss in conoideans has recently been reviewed by

Kantor & Sysoev (1989).

For the purposes of the present analysis, we have

attempted to recognise different morphological types of

radula, without any functional interpretation.

The radula of the Drilliinae, which is usually regarded as

the least-derived condition within the Turridae, has five teeth

in each transverse row (Fig. 16a). These teeth are usually

refered to as central, lateral and marginal teeth respectively;

although there are different interpretations (Kantor, 1990;

Starobogatov, 1990). We consider the morphology of each of

these teeth in turn.

1. Central tooth

A central tooth is present in species of Drilliinae, Pseudome-

latominae, Turrinae, Clavatulinae, and Cochlespirinae. It can

135

be reduced and lost in some species of these subfamilies

except Pseudomelatominae. (i) In the Pseudomelatominae,

the central tooth is fairly robust and broad, with a large

curved central cusp and sometimes smaller cusps at either

edge (Fig. 17e & f). (ii) In the Drilliinae the central tooth is

robust, but small and narrow (Fig. 16b & d), usually with a

prominent central cusp and a number of subsidiary cusps. (iii)

In the Turrinae and Clavatulinae (Figs 18a—d, 19a & b), the

central tooth appears broad, but apart from a spine-like

central cusp is poorly defined. The central cusp appears

homologous with the central tooth of the Drilliinae, but the

insubstantial, lateral ‘wings’ may represent vestiges of lateral

teeth which have fused with the central tooth. Alternatively,

the whole tooth might be homologous with the central tooth

of the Pseudomelatominae, the central cusp remaining promi-

nent, but the lateral edges becoming less substantial. Study of

the ontogeny of the radula in these taxa might distinguish

between these alternative possibilities.

2. Lateral teeth

We recognise two types of lateral teeth. (1) In what is

considered to be the least-derived condition, most species of

Drilliinae have large, multicuspidate, comb-like, lateral teeth

(Fig. 16a,c,e). However, reduced teeth are found in some

drilliine species (Bandel, 1984, fig. 306). (ii) In Antiplanes

(Cochlespirinae), the radula folds along the middle of the

radular ribbon, suggesting that the poorly defined, plate-like

teeth are in fact laterals (Kantor, 1990; Kantor & Sysoev,

1991, figs 26-27, 30-32). These ‘teeth’ were not visible on

S.E.M. preparations. Similar, poorly defined, lateral ‘teeth’

are also present in optical preparations of Crassispira and

Crassiclava of the Crassispirinae (Maes, 1983 fig. 31 & 37, p.

322; Kilburn, 1988, p. 239).

In all other subfamilies of Turridae, Pervicaciidae, Tere-

bridae and Conidae, lateral teeth are absent.

Fig. 13 Strictispira paxillus; transverse section of the rhynchoel and the proboscis tip. a, mouth with distally-situated radula and virtually no

bucal tube. Scale bar = 100 um. b, section of the proboscis slightly to the posterior of (a) showing the two large odontophoral cartilages.

Scale bar = 100 um.

136 J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV

3. Marginal teeth

In most conoideans the marginal teeth are the principal

functional teeth. Although diverse in appearance, they can be

divided into three broad categories of solid, wishbone and

hollow. There may be several subdivisions of each category.

Teeth of the first category are represented by a single, flat,

distally acute plate. Wishbone teeth are characterised by two

plates connected to each other. Hollow teeth are distin-

guished by a cavity within the tooth.

a) Solid marginal teeth

We recognise four main categories of solid teeth. (i) Simple,

flat teeth, often with a simple, blunt barb (Figs 16a, f, Fig.

20a). This type of tooth is common in the Drilliinae. (i1)

Simple teeth as in (i), but with the lateral edges of the tooth

curved to form a channel or gutter. This type of tooth has

been recorded from Drillia cydia (Powell, 1966, fig 81; Maes,

1983, fig. 28). (ii) Simple, solid teeth, which are curved and

pointed (Fig. 17e). This type of tooth is found only in

Pseudomelatominae (Kantor, 1988) and the Pervicaciidae

(Taylor, 1990).(iv) Simple, awl-shaped teeth with a large base

a } ; ; and pointed tip and a spathulate process midway along the

Fig. 14 Funa latisinuata, anterior alimentary system. A, proboscis tooth (Figs 17a-d). This type of tooth has been found only in

with buccal mass in extended position; B, with buccal mass in the subfamily Strictispirinae.

remacied tad aug sowigeute eop obit ‘oesephapus b) Wishbone teeth (sometimes called duplex teeth)

situated to the anterior of the nerve ring. Modified from an Ta. this type of dentition, the amaeainal toctneeeee te

unpublished drawing by J. Miller. Abbreviations: bm, buccal de 5 g E

mass; bt, buccal tube; con, circum-oral nerve ring; mb, muscular parts; pee sree ies pee: ee = is eee

bulb: oll Gesaphagealiloop ve venombelant! limb. Published illustrations suggest a great variety of form in

wishbone teeth, but S.E.M. observations show that some of

this variety results from artifacts produced by the transpar-

ency of light microscopy and by different positions of teeth

(often with displaced tooth parts) in preparations.

We recognise four basic types of wishbone teeth:

(i) Broad, slightly curved teeth, sometimes with a blunt

barb (Fig. 20 b-d). The lateral edges of the teeth are

thickened, with a thin accessory limb attached to the main

tooth at the anterior and posterior ends. This type of tooth is

common in some Crassispirinae such as /nquisitor, Paradrillia

and Funa, where the size and shape of the accessory limb

varies considerably between species (Kilburn, 1988). Because

the main limb is similar to the marginal teeth of the Drillii-

nae, we suggest this as the least-derived type of wishbone

tooth. (ii) The teeth of this type are robust, short and curved,

sometimes with a knife-like cutting edge on the main limb

and a large accessory limb (Figs 18a,c; 19a,d). Teeth of this

type are found in species of Turrinae, Clavatulinae, and

Cochlespirinae. (iii) Teeth that may be modified wishbone

teeth have been illustrated for Ptychobela nodulosa and

P.suturalis by Kilburn (1989, figs 17-19). The teeth are

awl-shaped without barbs, with apparently two nearly equi-

size limbs joined to form a central channel. An S.E.M. study

of these teeth is needed to claify their morphology. (iv) In the

radula of Ptychobela griffithi the teeth appear to be robust

and solid with a simple barb (Fig. 22a), but they may in fact

Fig. 15 Diagrammatic section through the radular sac. A, in

turrids possessing an odontophore; B, turrids lacking an

odontophore, but with a radula caecum. Abbreviations: bs,

buccal sac; od, odontophore; rs, radular sac; rc, radula caecum;

slp, sublingual pouch; t, radular teeth. Buccal sac is that portion

of the radular sac lying between the entrance of the salivary ducts

and the buccal cavity.

Fig. 16 Radulae of Drilliinae. a, half radula row of Clavus sp. from Guam showing blade-like marginal teeth, comb-like lateral teeth and the

small central tooth. Scale bar = 50 um. b, central tooth of Clavus unizonalis. Scale bar = 5 um. c, central and part of lateral teeth of

Spirotropis monterosatoi. Scale bar = 20 um. d, central tooth of S. monterosatoi. Scale bar = 10um. e, single lateral tooth of S.

monterosatoi. Scale bar = 20 um. f, marginal teeth of S. monterosatoi. Scale bar = 20 um.

Fig. 17 Radulae of Strictispirinae and Pseudomelatominae. a, radula of Strictispira paxillus. Scale bar = 50 um. b, marginal teeth of

Strictispira stillmani. Scale bar = 50 um. c, radula of Cleospira ochsneri. Scale bar = 50 um. d, marginal teeth of Strictispira paxillus seen

from side. Scale bar = 50 um. e, radula of Pseudomelatoma penicillata. Scale bar = 100 um. f, central tooth of P. penicillata seen from side.

Scale bar = 10 um. (see p. 138)

FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA

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FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA

Fig. 18 Radulae of Clavatulinae and Cochlesprinae. a, Clionella sinuata; wishbone marginal and small central teeth. Scale bar = 50 um. b.

Clionella sinuata small central tooth. Scale bar = 10 wm. c. Turricula nelliae spurius, radula with wishbone marginal teeth and central tooth

with spine-like cusp and lateral flanges. Scale bar = SO um. d. T. nelliae spurius central tooth. Scale bar = 5 um.

J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV

Fig. 19 Wishbone teeth of Turrinae and Cochlespirinae. a. radula of Gemmula deshayesi. Scale bar = 50 um. b. marginal tooth of Gemmula

deshayesi Scale bar = 20 um. c. marginal tooth of Lophiotoma acuta Scale bar = 10 um. d. marginal tooth of Antiplanes sanctiioannis.

Scale bar = 20 um..

FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA

be formed from two pieces as in Ptychobela suturalis (see

above). Lack of material precluded further study of this and

the type iii wishbone teeth.

c) Hollow teeth

There is a great diversity of detailed variation in the form of

hollow marginal teeth even within a single genus (see for

example, James (1980) and Nybakken (1990) for Conus and

Bogdanov (1990) for Oenopota). However, for the purposes

of this analysis we recognise only five main types of hollow

teeth. (i) Teeth of this type are long, slender, and enrolled,

141

with a small base. The base is not differentiated morphologi-

cally and is not solid. The distal end of the tooth may be

simple, or more or less, elaborately barbed (Figs 22e,g).

There is an opening near the distal tip and a second opening

placed more or less terminally at the proximal end. The shaft

of Hastula hectica is perforated by holes (Taylor, 1990, fig. 2).

For some Conus species, Nybakken (1990) has shown that

during ontogeny, the hollow, rolled teeth develop from open,

guttered forms and become progressively more elaborately

barbed. Hollow teeth of Type i are found in species of

Fig. 20. Radulae of Clavinae and Crassispirinae. a. marginal tooth of Drillia rosacea. Scale bar = 50 um. b. Funa latisinuata; blade-like

marginal teeth with thin accessory limb. Scale bar = 50 um. c. Vexitomina garrardi; part of blade-like marginal tooth with accessory limb

(arrowed). Scale bar = 10 um. d. enlargement of (d) showing accessory limb. Scale bar = 10 um.

142

Borsoniinae, Clathurellinae, Toxiclionella (Clavatulinae),

Conidae, and Terebridae (ii) Hollow teeth of this second type

are often short with a large, solid base (Fig. 23). The tooth

cavity opens laterally between the shaft and the base. There

are frequently side projections around the base (hilted dagger

form of Powell 1966), often with a large irregular solid ‘root’

projecting from the base (Fig. 23e,f). These teeth are often

only partially enrolled. Barbs may be present. Marincovich

(1973) records rows of holes in the teeth of Agathotoma

ordinaria (Mangeliinae). Teeth of Type ii are found in the

subfamilies Mangeliinae, Oenopotinae (Bogdanov, 1990, figs

407-438), Thatcheriinae, and the radulate Daphnellinae. (111)

Teeth of this type are partially enrolled at the base, but solid

and blade-like in the distal part (Fig. 22b). This type of tooth

is presently known only from Hastula bacillus (Taylor &

Miller, 1990). It may represent a transitional form between

the solid teeth found in the Pervicaciidae and the hollow teeth

of the Terebridae. (iv) This type of tooth is loosely enrolled

to form a central channel, with a simple barb at the tip. The

tooth was first described in detail from Jmaclava unimaculata

(Clavinae) by Shimek & Kohn (1981 fig. 7). Imaclava other-

wise has comb-like lateral teeth as in typical Clavinae. Similar

teeth are present in other species of Imaclava (McLean, 1971,

fig. 7). (v) Enrolled teeth with a complex appearance are seen

in Pilsbryspira nympha (Zonulispirinae) (Fig. 21). Although

these are hollow teeth with a small barb, the shaft is complex

and appears to be formed by partial enrolling of two units

(Fig. 21b). The tooth may be derived by the enrolling of the

elongate wishbone teeth typical of the Crassispirinae. (vi)

Vestigial teeth, semi-enrolled, with a gutter along the tooth.

Teeth of this type are considered by Bogdanov (1990) as

J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV

derived from the distal part of the shaft of Type ii teeth. This

type of tooth is found in Propebela turricula and P. harpularia

(Oenopotinae) (Bogdanov, 1990, figs 41, 433).

Glands of the foregut

Salivary glands

Salivary glands are present in most turrids, Conus and the

radulate species of Terebridae and Pervicaciidae. In most

species a pair of glands is present, but these may be fused

together. The salivary ducts always open into either side of

the buccal sac (Fig. 1). In Turricula nelliae spurius, which has

a distal buccal mass, the salivary glands are contained within

the proboscis and attached to the oesophagus (Miller, 1990).

In most conoideans, the salivary glands are acinous, but in

the Mangeliinae, Thatcheriinae, Daphnellinae and Hae-

dropleura septangularis (Crassispirinae) the glands consist of

long, convoluted, single tubes (Sheridan ef al., 1973; own

observations).

Turrids without a radula also lack salivary glands, but in

the Terebridae, glands are present in some radula-less forms,

such as Terebra gouldi and T. maculata (Miller, 1970, 1975).

Accessory salivary glands

These are known in a few species of Turridae, some Conidae

(Marsh, 1971; Schultz, 1983) and Terebridae (Taylor &

Miller, 1990; Taylor, 1990). They have a similar histology to

the accessory salivary glands found in other neogastropod

families such as the Muricidae (Andrews, 1991). Further-

Fig. 21. Enrolled teeth of Pilsbryspira nympha. a. several adjacent marginal teeth. Scale bar = 25 um. b. detail of base of tooth showing

double structure (arrow) suggesting that tooth may be formed by the enrolling of wishbone teeth. Scale bar = 5 um.

Fig. 22 Single marginal teeth from Turridae and Terebridae. a. Ptychobela griffithi. Scale bar = 10 um. b. Hastula bacillus. Scale bar =

5 um. c. Glyphostoma candida Scale bar = 50 um. d. enlargement of the tip of the G. candida tooth. Scale bar = 10 um. e. Genota

mitraeformis. Scale bar = 20 um. f. Terebra babylonia. Scale bar = 20 um. g. Conus ventricosus Scale bar = 20 um.

viii

FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA

J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV

FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA

more, the ducts from the accessory glands open near the tip

of the buccal tube, which is the homologous position to that

found in other neogastropods.

Within the Turridae, we have observed accessory salivary

glands in only two subfamilies: the Borsoniinae (Scrinium

neozelanicum, Borsonia ochracea, and Micantapex parengo-

nius) and Cochlespirinae (Aforia hypomela, A. kupriyanovi,

A. abyssalis). In the Terebridae, we have seen accessory

glands in Hastula bacillus, Terebra babylonia, T. funiculata

and T. subulata (Taylor, 1990). Usually, only a single gland is

found, but two glands are present in Terebra subulata.

Venom apparatus (venom gland and muscular bulb)

The long, tubular, and convoluted venom gland is the most

conspicuous organ of the conoidean foregut. It always passes

through the nerve ring and always opens into the buccal

cavity immediately posterior to the opening of the radular sac

(Figs 1 & 7). The venom gland is present in most conoideans,

except the radulate Strictispira (Maes, 1983); Gymnobela

tincta, which has a vestigial radula; the radula-less turrids

from the subfamilies Daphnellinae and Taraninae (Smith

1967; Sheridan et al., 1973; Kantor & Sysoev, 1989), the

radula-less Terebridae (Miller 1975; Taylor, 1990) and the

radulate Pervicaciidae (Taylor, 1990).

In some species, the histology of the venom gland changes

in the anterior portion of its length, after its passage through

the nerve ring. The posterior portion is packed with venom

granules (Fig. 24), but the anterior portion is duct-like and

ciliated (e.g. Clavatula, Clionella, Turricula, Lophiotoma and

Pilsbryspira). This change in histology is usually correlated

with the elongation of that part of the oesophagus lying

between the nerve ring and buccal mass. In other conoideans,

venom granules are present all the way along the length of the

gland, sometimes even into the buccal cavity.

Extensive studies have been made of the composition and

pharmacology of the venom in a few Conus species (review

by Oliviera et al., 1990). The composition of the venom is

very complex and the results from these studies have a

potential utility in phylogenetic analysis. However, no com-

parable studies yet exist for the Turridae and Terebridae.

Muscular bulb

The muscular bulb (Figs 1 & 6) lies at the posterior end of the

venom gland and is present in all those species possessing the

gland. Differences between taxa are observed both in the

number, orientation and relative thickness of the various

muscular layers forming the wall of the bulb. The usual

condition is of an outer, circular-muscle layer, a thin, middle

connective tissue layer, with an inner longitudinal layer. We

have, however, observed other configurations of the muscle

layers. For example in Mangelia species and Eucithara, the

outer muscular layer is very thin, but the inner layer very

thick. Daphnella reeveana has only a single, thin muscle

layer, whilst Conus textile has four distinct alternating circular

and longitudinal muscle layers, three of them lying inside the

connective tissue layer.

Additionally, Ponder (1970) mentions that he has observed

glandular cells in the epithelium lining the muscular bulb in

145

Lucerapex (Turrinae) and Maoritomella albula (Borsonii-

nae). We have not observed the glandular cells in any turrid

we have examined.

Summary of foregut anatomy

From the foregoing discussion, it is clear that there is a great

variety of foregut anatomy present within the Conoidea and

considerable variation may be present even within species of

one subfamily. As a summary, twelve of the main types of

foregut configuration are shown diagramatically in Figs 25 &

26. It should be emphasized that only a relatively small

number of conoidean species have been investigated ana-

tomically and it is likely that further types of foregut remain

undiscovered. Nevertheless, there are several anatomical

characters which define the Conoidea and are present in most

representatives (and in all the least derived groups). These

are:-

1. The presence of a venom gland.

2. The buccal mass located at the base of the proboscis.

3. The proboscis formed by the elongation of the buccal

tube.

4. The presence of a permanent rhynchodeum.

5. The tendency for the loss of central and lateral teeth from

the primary five toothed radular row.

FUNCTIONAL MORPHOLOGY OF THE

DIGESTIVE SYSTEM AND FEEDING

MECHANISMS IN TOXOGLOSSA

As has been outlined in the previous section, the morphology

of the digestive system of Conoidea and especially that of the

Turridae, is highly varied. These variations in morphology

probably reflect differences in feeding behaviour and diet.

Apart from Conus, conoidean diets are still very poorly

known. Indeed, for in excess of 4000 living species of

Turridae, feeding information is available for less than 30

species (reviewed by Miller, 1989). These data, derived

mainly from gut content analysis, show that turrids feed

mainly on errant and sedentary polychaetes and more rarely

on other phyla such as sipunculans, nemerteans, and mol-

luscs. Very few direct observations of the feeding process in

the Turridae have been made (Pearce, 1966; Shimek, 1883a,

b, c; Shimek & Kohn 1980; Miller, 1990). Because of this lack

of information, our conclusions concerning the feeding

mechanisms of Turridae are based upon analysis of the

morphology of the digestive tract and by comparison with

species whose feeding mechanism is known.

Our classifications of feeding mechanisms is based upon

the following characters listed in order of priority: the

presence/absence of venom apparatus (used for immobilizing

or killing the prey); the mode of radula function ( which may

be used solely as a whole organ, as a whole organ with

simultaneous use of separate teeth, or as separate teeth only

at the proboscis tip); position of the buccal mass (either basal

or shifted anteriorly towards the proboscis tip). We recognize

Fig. 23 Hypodermic-type marginal teeth with a large solid bases. a. Paramontana sp. Scale bar = 2 wm. b. Propebela rugulata. Scale bar =

10 um. c. & d. Thatcheria mirabilis Scale bars = 20 um. e.& f. Mangelia powisiana. Scale bars = 5 um G. Eucithara stromboides. Scale bar

= 10 um.

J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV

Fig. 24 Venom gland of Clavus sp. Guam. a. section through critical-point dried venom gland showing venom granules. Scale bar = 10 um

b. enlargement of single venom granule. Scale bar = 1 um.

five main and several sub-types of feeding mechanism. Some

of these have already been described (Kantor & Sysoev,

1990; Kantor, 1990), but are here partially revised and

corrected.

I. Venom gland present

Feeding mechanism Type 1

The first functional type of digestive system and feeding

mechanism, that in which the radula is used only as a whole

organ in conjunction with the venom apparatus is found

among species of Pseudomelatominae and in Toxiclionella

tumida (Clavatulinae) and can be subdivided into two sub-

types.

The first sub-type is characteristic of the Pseudomelatomi-

nae, an endemic subfamily from western central America,

which includes 3 genera and several species (McLean in

Keen, 1971). The anatomy of two species Pseudomelatoma

penicillata and Hormospira maculosa indicates the isolated

position of the group among Conoidea (Kantor, 1988). This is

particularly clear, from the radular morphology, which con-

sists of a large and well developed central tooth, flanked by

large, scythe-like, but solid, marginal teeth.

The buccal mass is situated either at the proboscis base and

far ahead the nerve ring in Pseudomelatoma penicillata, or in

front of the nerve ring and distant from the proboscis base in

Hormospira maculosa. The anterior part of the digestive tract

forms a long curve, either by the elongation of that part of the

oesophagus between the nerve ring and the buccal mass (P.

penicillata), or by the elongation of the posterior part of the

buccal tube (H. maculosa).

Both species have a well-developed venom gland and

although the diet of Pseudomelatominae is unknown, the

presence of the large venom gland indicates the predatory

mode of feeding. The gastropods also have a muscular

proboscis with a wide oral opening but without a sphincter.

The absence of the oral sphincter, which is usually used for

holding single radular teeth at the proboscis tip (Kantor &

Taylor, 1990), coupled with the curved form of the marginal

teeth, indicate that the gastropods do not use separate teeth

for stabbing the prey. Kantor (1988) supposed that prey

capture occurs with the aid of the proboscis tip and is

facilitated by the wide and highly extensible oral opening. If

this is so, then envenomation of the prey should occur within

the anterior part of the proboscis. This facilitates the trans-

port of prey into the buccal cavity, by the peristaltic move-

ments of well-developed circular muscles of the buccal tube.

However, the presence of the elongated part of the

oesophagus between the buccal mass and nerve ring in P.

penicillata may indicate another mode of prey capture. In

some turrids (e.g. Funa latisinuata, Fig. 14), the presence of

such an elongation of the oesophagus is connected with the

ability to evert the buccal mass, with the radula, through the

proboscis and mouth. It is possible, that P. penicillata can

evert the buccal mass through the mouth and use the radula

directly in prey capture. Envenomation would in this case

occur through the damage to the prey made by the radular

teeth. Also the very large odontophore (the largest of all the

turrids studied) suggests that the radula may also tear the

prey.

The morphology of Hormospira differs from that of

Pseudomelatoma, in that the curve is formed by the posterior

part of the buccal tube and elongated buccal mass. The

FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA

Fig. 25 Diagram (with Fig. 26) summarizing some of the major

types of foregut morphology found amongst the Conoidea, with

radulae, where present, illustrated alongside. Not to scale. A.

Clavus unizonalis; B. Clionella sinuata; C. Turricula nelliae

spurius; D. Mangelia nebula; E. Ophiodermella inermis; F.

Daphnella reeveana. Abbreviations: asg, accessory salivary

glands; sg, salivary glands; rs, radular sac; vg, venom gland; black

dots are sphincters.

radular sac is located far behind the base of the proboscis.

Therefore, it is doubtful that the buccal mass can be everted

through the mouth opening. This species probably catches

prey using the proboscis tip. Envenomation could occur

either by the squirting of venom through the mouth, when the

proboscis is in contact with the prey, or in the anterior part of

the proboscis, when the prey is partly swallowed. In either

case the radula is not used to envenomate the prey and is

either used for further transportation in the oesophagus of for

partial tearing of prey tissue.

The second sub-type is found in Toxiclionella tumida and

differs from the first in that the buccal mass is located near

the proboscis tip (Kantor, 1990, fig. 4), which has no distal

sphincter. This species is characterized by a radula formed of

hollow, and barbed marginal teeth (Kilburn, 1985, fig. 14),

which are attached all along their length to the radular

membrane. The hollow radular teeth are similar in morphol-

ogy to those of higher conoideans. The gastropod has a long

venom gland and in the posterior part of the proboscis there

is a single salivary gland with paired ducts. The radular teeth

are sufficiently long, that during protraction of the odonto-

147

Fig. 26 Further types of foregut morphology found in the

Conoidea. G. Gymnobela emertoni; H. Philbertia linearis; 1.

Conus ventricosus; J. Duplicaria spectabilis; K. Terebra subulata,

L. Terebra maculata.

phore, the tips would protrude through the oral opening, and

thereby stab the prey.

A comparable mechanism may occur in Turricula nelliae

spurius (Taylor, 1985), which has the buccal mass located in a

similar distal position in the proboscis to that of 7. tumida,

and during feeding can protrude the odontophore through

the mouth opening (Miller, 1990). But T. nelliae possesses a

sphincter in the anterior part of the buccal tube, and this

feature usually correlates with the use of separate marginal

teeth for stabbing (Kantor & Taylor, 1991).

In conclusion, we suggest that a similar type of feeding

mechanism evolved independently in Pseudomelatoma and

Toxiclionella. In the former, the primitive character of the

radula suggests that the feeding mechanism is primary;

whilst in Toxiclionella it is probably a secondary feeding

mode when compared with other members of .the subfam-

ily. It is possible that with the shift of the buccal mass to the

proboscis tip, Toxiclionella lost the mechanism of stabbing

the prey with single marginal teeth and instéad protrudes

the radula through the mouth and uses the hotlow:teeth

which remain firmly anchored to the radular mémbrane. *

148

Feeding mechanism Type 2

The second feeding mechanism is typical of the majority of

‘lower’ turrids and the terebrid Hastula bacillus, which pos-

sess a well developed radular membrane and lack a radular

caecum. The characteristic feature of this mechanism is the

use of separate marginal teeth at the proboscis tip for

stabbing the prey, whilst the radula is also used as a whole

organ for different purposes (Sysoev & Kantor, 1986, 1989).

The use of single marginal teeth at the proboscis tip by

turrids having radulas with well developed subradular mem-

branes has been demonstrated in representatives of all 7

subfamilies of ‘lower’ Turridae (excepting the Pseudome-

latominae) and also the terebrid Hastula bacillus (Kantor &

Taylor, 1991).

According to the position of the buccal mass this type may

be divided into two sub-types. Gastropods of the first sub-

type have the buccal mass situated at the proboscis base.

These include species of Drilliinae, Cochlespirinae, Turrinae

and many Crassispirinae. In these gastropods, the solid or

wishbone marginal teeth, which become detached from the

membrane during its degeneration in the sublingual pouch,

are used at the proboscis tip for stabbing the prey. It should

be noted, that separate teeth were not found in the sublingual

pouch, therefore it does not serve for the storage of teeth.

Meanwhile, the radula as a whole organ probably has a

different function within the buccal cavity. This is most likely

for the transport of food from the cavity to the oesophagus.

Some evidence for this comes from the observations of Maes

(1981), who noted the presence of intact sipunculans in the

posterior part of the oesophagus of Drillia cydia (Drillinae).

Although at first sight, it might be thought that the large,

pectinate, lateral teeth found in this species might serve for

tearing or rasping the prey.

A characteristic feature of the proboscis is the presence of

the sac-like enlargement of the anterior part of the buccal

tube and a well-developed, distal sphincter(s). Gastropods of

this group lack a radular caecum, so they can use only teeth

which are sporadically detached from the membrane. Either

the marginal teeth are not used in every feeding act, or, the

teeth are held at the proboscis tip for a long time. That is,

from the moment of their detachment from the subradular

membrane to the next feeding act. We have found teeth at

the proboscis tip in sections of ‘lower’ turrids much more

frequently, than in the ‘higher’ turrids. Moreover, in Splen-

drillia chathamensis, in addition to the normal buccal sphinc-

ters of the buccal tube, teeth are attached by their base to a

‘mat’ of epithelial cells in the enlargement of the buccal tube

(Kantor, 1990, fig. 3). Such a mechanism of tooth fixation

confirms the long-term presence of the tooth at the proboscis

tip. Thus, the enlargement of the anterior part of the buccal

tube, could be considered as a functional analogue of the

radular caecum.

The use of marginal teeth at the proboscis tip, in turrids

with a well-developed radular membrane, explains how hol-

low, marginal teeth might have evolved independently in

different groups possessing the radular membrane and odon-

tophore. For example, Jmaclava (Drillinae) (Shimek &

Kohn, 1981), has hollow teeth and most probably uses these

at the proboscis tip for stabbing the prey in a manner similar

to that of higher Conoidea.

The second feeding sub-type is seen in Funa latisinuata

(Crassispirinae), which feeds upon nemerteans. From dissec-

tion of relaxed animals, Miller (1989, fig 6f) showed that in

J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV

the everted position, the buccal mass with the radula is

protruded through the mouth opening (Fig. 14a). In sections

of animals with a contracted proboscis, the buccal mass lies

towards the base. It is known that this species uses the

marginal teeth at the proboscis tip (Kantor & Taylor, 1991).

Thus, the mode of feeding may be reconstructed as follows.

After stabbing the prey, the gastropod everts the buccal

mass, with the walls of the buccal tube, through the mouth

opening and picks up the prey with the protruded radula.

With retraction of the buccal mass, the prey is pulled into the

proboscis. Correlated with this feeding mechanism, is the

elongation of the anterior oesophagus between the buccal

mass and the circum-oral nerve ring. During protraction of

the buccal mass, the oesophagus should be pulled through the

nerve ring. But, as the nerve ring in Conoidea is highly

concentrated, and usually tightly attached to the oesophagus,

the only possibility is the elongation of the oesophagus itself

anterior to the nerve ring, forming a loop, which is straight-

ened during eversion of the buccal mass (Fig. 14b).

In addition to Funa latisinuata, this elongation of the

oesophagus between the buccal mass and the nerve ring has

been found in species from several different subfamilies of

Turridae—Pseudomelatominae, all Clavatulinae, Pilsbryspira

nympha (Zonulispirinae), Vexitomina (Crassispirinae), Tur-

ricula nelliae spurius (Cochlespirinae), the radulate terebrids,

Hastula bacillus, and Pervicacia tristis (Pervicaciidae). It is

likely, that the turrid species at least have a feeding mecha-

nism similar to that of F. latisinuata. The elongation of the

anterior oesophagus is usually associated with the permanent

shifting of the buccal mass towards the distal end of the

proboscis. This is well demonstrated in the Clavatulinae and

probably facilitates the eversion of the buccal mass through

the mouth.

In all species possessing an elongated oesophagus (except

Pseudomelatoma), there is a change in the histology of the

ante-rior part of the venom gland after its passage through

the nerve ring. However, such a change occurs in two species

(Lophiotoma leucotropis and Inquisitor sp.) which lack the

elongated oesophagus. The anterior part of the gland is

ciliated and duct-like, with no secretory granules. This indi-

cates, that the differentiation of the gland is connected with

the elongation of the oesophagus and thus, the latter is a

secondary feature.

Feeding mechanism Type 3

The majority of Conoidea possess the third type of feeding

mechanism, in which separate marginal teeth are used at the

proboscis tip for stabbing prey, and the radula not used as a

whole organ.

The very specialized radular morphology is the most

remarkable and well-known feature of the toxoglossan diges-

tive system. It is characterized by a marked tendency towards

a reduction in the strength of the subradular membrane,

leading to its complete absence in many species of Turridae,

the majority of Terebridae and all Conidae. Species without a

subradular membrane, have a radula consisting only of

complex, hollow, marginal teeth. They are known for the

highly specialized feeding mechanism, in which individual

teeth are used at the proboscis tip for stabbing and killing

prey with secretions of peptide neurotoxins produced by the

venom gland (Oliviera et al. 1990).

Despite the similarities with the previous feeding mecha-

nism, those ‘higher’ conoideans with hollow teeth and no

FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA

radular membrane are extremely diverse compared with the

‘lower’ conoideans. Moreover, this relative diversity has

steadily increased throughout the Cenozoic (Sysoev, 1991).

This suggests that higher conoideans may possess some

adaptive advantages. In our opinion these advantages lie in

the features of the morphology of the radular diverticulum.

The higher Conoidea lack a subradular membrane, and the

radular diverticulum is divided into two different parts; the

radular sac and radular caecum (also known as long and short

arms). The caecum serves for the storage of the fully-formed,

marginal teeth. Many teeth can be stored; for example, in a

specimen of Mitromorpha (Mitrolumna) sp. there were 106

teeth in the radular sac compared with 64 in the caecum

(Kantor & Sysoev, 1990). Species of higher Conoidea can

probably use several teeth in each feeding act. For example,

observations on the feeding of Conus textile showed that up to

17 teeth can be used in the same attack (Schoenberg, 1981).

By contrast, in lower turrids, there is no caecum and probably

no more than a single tooth can be used in each feeding act.

Predatory attacks by higher Conoidea are thus likely to be

more successful, and the mechanism of prey capture probably

more efficient. This may explain the relative success of the

higher Conoidea.

The feeding and diets of gastropods of this functional type

are well known (Oliviera et al. 1990) and it is unnecessary to

describe the process in detail. Only the most important

morphological features should be noted. These are the vesti-

gial, or completely reduced, radular membrane; the absence

of an odontophore; the presence of a radular caecum where

the fully-formed marginal teeth are stored, and a well-

developed oral sphincter for gripping the teeth. The radula is

represented by hollow, marginal teeth. The tooth ligament

(long flexible stalk attached to the tooth base) is probably the

rudiment of the radular membrane (Fig. 23c). Also the

gastropods of this group often have enlarged rhynchostomal

lips. In some species, the lips are able to invert (i.e. to form

an introvert or pseudoproboscis) and this is also used in prey

capture. It should be noted, that some vermivorous species of

Conus (Marsh, 1970) and the fish-feeding C. geographus

(Johnson & Stablum, 1971) do not stab they prey in every

feeding act. This is possibly an initial stage of transition to

feeding mechanism type 5.

Usually, teeth are gripped at the proboscis tip by the buccal

tube sphincter, but in some turrids the buccal tube introvert

(valvule of Sheridan et al., 1973) is involved (Fig. 9). This

structure has been reported so far in Mangelia nebula

(Sheridan et al., 1973; Delaunois & Sheridan, 1989) and in

Eucithara stromboides (Fig. 10). It is also possible, that the

buccal tube introvert can be everted through the mouth

| opening and have a role in holding the prey.

After envenomation, the prey may be held by the tooth

itself, as occurs in many vermivorous species of Conus

| (Kohn, 1959), or with the mouth. The buccal lips may play a

| role in the transport of prey to the buccal cavity. These are

highly protrusive in many Mangeliinae, and at least in M.

, nebula (Fig. 9) can be retracted into the buccal cavity

| (Delaunois & Sheridan, 1989). A similar possibility was

_ described for Oenopota by Bogdanov (1990), who suggested

_ that the buccal lips and the proboscis itself might be inverted

into the buccal cavity.

149

II. Venom gland absent

Feeding mechanism Type 4

Gastropods of this group have a radula with a well-developed

radular membrane and a proboscis may be either present or

reduced. According to the position of the buccal mass they

can be divided into two sub-types.

Conoideans of the first sub-type which at present includes

only Strictispira and probably Cleospira, have the buccal mass

located at the tip of a well-developed proboscis (Fig. 13). The

buccal mass and radular apparatus are large, with two large

odontophoral cartilages and massive odontophoral and pro-

boscis retractor muscles. The radula has a strong membrane

with two rows of solid, awl-shaped, marginal teeth. The

buccal tube is very short and there are no oral sphincters.

Apart from the record of polychaete setae in two individuals

of Strictispira paxillus (Maes, 1983), nothing is known of the

habits of this group.

The terminal position of the buccal mass on the muscular

proboscis, the short buccal tube and the massive radular

apparatus, suggest that when the gastropod is feeding the

radula is protracted out of the extended proboscis tip. The

solid teeth and absence of venom apparatus suggest that the

radula is involved in biting and tearing rather than stabbing.

The feeding mechanism is thus probably more similar to

other neogastropods such as the Buccinidae rather than to

other conoideans.

Conoideans of the second sub-type differ from these of the

first one in possessing a basal buccal mass. The radula is

well-developed, whilst the proboscis is either absent or highly

reduced, and a rhynchostomal introvert is usually present.

This feeding mode is found in the Pervicaciidae. The diet of

this family is largely unknown, except for ‘Terebra’ nassoides

which feeds on capitellid polychaetes (Taylor, 1990).

In the Pervicaciidae, the absence of a proboscis means that

the rhynchodeal introvert becomes the main organ of prey

capture, as occurs in some proboscis-less terebrids such as T.

gouldi (Miller, 1975). Prey are presumably pulled into the

rhynchocoel by the introvert. In Duplicaria spectabilis there

are large muscular buccal lips and probably a protrusive

odontophore (Taylor, 1990, fig. 7). However, in Pervicacia

tristis and Duplicaria kieneri there is a septum with a narrow

aperture dividing the rhynchocoel and it is very unlikely that

the odontophore can be protruded through the septum.

Although we have no direct evidence, it is possible that the

septum functions to hold prey during swallowing and perhaps

early digestion.

Feeding mechanism Type 5

Finally, there are many conoideans which lack a radula,

venom and salivary glands. Gastropods of this group include

some Daphnellinae, Taraninae and some Terebridae. In

addition to the absence of foregut glands and radula, a

characteristic feature of these species is the very reduced size

or complete absence of the proboscis. Radula-less Conoidea

either have well-developed, rhynchostomal lips or a large

rhynchostomal introvert, as for example, in the Terebridae

(Miller, 1975) or Philbertia linearis (Sheridan et al. , 1973). It

is possible, that a rhynchostomal introvert is also present in

Teretiopsis, although all sectioned specimens have it in the

extended position and it was overlooked during the original

description (Kantor & Sysoev, 1989). In some turrids, such as

150

Cenodagreutes (Smith, 1967) and Abyssobela atoxica (Kantor

& Sysoev, 1986), which lack the rhynchodeal introvert, there

is a vast rhynchocoel and well-developed cavity between the

rhynchodaeum and body walls. The walls of this cavity are

connected by numerous transverse muscles. Both the intro-

vert and cavity are lacking in the genus Taranis (Taraninae).

A feeding mechanism for radula-less species is known for

some terebrids (Miller, 1970, 1975). Thus, Terebra gouldi

which has a relatively short introvert feeds upon the enterop-

neust Ptychodera flava, and Terebra maculata with a long

introvert feeds on polychaetes. Prey are caught with the aid

of the introvert. Turrids lacking the introvert, but with the

cavity between the rhynchodaeum and the body walls, prob-

ably engulf prey by contraction of the radial muscles in the

wall. This would cause negative pressure and an increase in

the inner volume of the rhynchocoel.

The origin of the radula-less feeding mechanism can be

easily envisaged. It is known, that in some Conus species

hypodermic envenomation is not necessary in each feeding

attack (Kohn, 1959; Marsh, 1970; Sanders & Wolfson, 1961).

It is probable that some Turridae and Terebridae, especially

those with well-developed rhynchostomal lips or introvert,

also feed without stabbing the prey with radular teeth. Thus,

Daphnella reeveana, which possesses a venom gland, has a

very short proboscis and is probably unable to hold a tooth at

its tip (Fig. 4). As stabbing of the prey becomes unnecessary,

the proboscis, venom gland and radula disappear. An inter-

mediate stage is found in Gymnobela emertoni, in which the

proboscis and venom gland have disappeared, but there is

still a very short and reduced radular sac, opening to the

outer side of the buccal lip (Fig. 8).

RELATIONSHIPS OF THE CONOIDEA

Monophyly of the Conoidea

There has been much discussion concerning the relationships

of the Conoidea to other prosobranch gastropods; some

considering them to be part of a monophyletic group with

other neogastropods (Ponder, 1973; Taylor & Morris, 1988),

whilst others suggest an origin entirely independent of the

neogastropods (Sheridan et al. 1973; Shimek & Kohn, 1981;

Kantor, 1990).

In this section we briefly review some of the evidence for

the relationships of the Conoidea with other prosobranchs.

Some of this evidence has been discussed in some detail by

Kantor (1990) and only the principal arguments are presented

here.

The location of the buccal mass at the base of the proboscis

as found in most conoideans, is different from the situation

seen in most neogastropods, where the buccal mass is found

at the distal end of the proboscis. The proboscis in most

conoideans is formed by the elongation of the buccal tube,

whilst in neogastropods it originates from the elongation of

the anterior oesophagus (Ponder, 1973). However, a basal

buccal mass is now known for the neogastropod Benthobia

(Pseudolividae) which also exhibits a number of other primi-

tive characters, and in Amalda (Olividae) (Kantor, 1991).

Additionally, in Benthobia, the radular retractor muscle

passes through the nerve ring and is connected to the

columellar muscle (Kantor, 1991 fig. 15a). This condition is

seen species of the turrid subfamily Drilliinae, and in most

J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV

lower caenogastropods, but is absent in probosciform caeno-

gastropods.

A key autapomorphy of the Conoidea is the possession of

the venom apparatus, comprising the venom gland and

muscular bulb. There has been much discussion concerning

the homology of this gland. But, Ponder (1970; 1973)

showed, that in the neogastropod family Marginellidae a long

coiled gland, similar in general appearance to the conoidean

venom gland is formed by the stripping off of glandular folds

from the oesophagus. In some marginellids the gland termi-

nates at the posterior in a muscular bulb which is homologous

with the gland of Leiblein. The venom gland of conoideans

may have been derived in a similar way and is probably

homologous with the glandular folds of the oesophagus and

the gland of Leiblein in other neogastropods.

The possession of tubular, accessory salivary glands is also

considered to be an apomorphy of the Neogastropoda (Pon-

der, 1973). These glands are patchily distributed amongst

conoideans, but are known in some Turridae, Conidae and

Terebridae. Both the histology of the glands (Schultz, 1983;

Andrews, 1991) and the position of the opening of the ducts,

confirms their homology in the Conoidea and in other

neogastropods. The primitive Benthobia also has a large

accessory salivary gland (Kantor, 1991).

A radula with five teeth in each row, as is found in the

turrid subfamily Drilliinae, has been considered as evidence

for a separate origin of the Conoidea and Neogastropoda, the

latter normally have three or less teeth in each row. (Shimek

& Kohn, 1981). However, it is now known that some Olivella

and Nassariidae have five teeth in each row (Bandel, 1984;

Kantor, 1991). All this suggests is that the common ancestor

of the Conoidea and the other neogastropods possessed five

or more teeth in each row.

In conclusion, conoideans share a number of characters

with the neogastropods which suggest a common ancestry.

Nevertheless, the evidence both from the position of the

buccal mass and the formation of the proboscis, suggests an

early divergence of the two groups. An evolutionary scheme

for the derivation of the conoidean intraembolic proboscis

from the acrembolic type, typical of many mesogastropods,

has been developed by Kantor (1990). His arguments cor-

roborate and elaborate Ponder’s (1973) hypothesis that the

Conoidea diverged from the other neogastropods before the

formation of the proboscis. Ontogenetic studies of proboscis

and foregut development in the Conoidea and other neogas-

tropods might provide corroborative evidence.

Relationships within the Conoidea

Phylogenetic analysis

We attempted to determine relationships within the

Conoidea using cladistic analysis of many of the foregut

characters described in the first part of this paper, combined

with a few shell characters.

Taxa used

We have included 40 species in the analysis, with at least one

from all the currently-recognised, subfamilies. In a few cases

we have used previously published work. The species studied

represent only a small proportion of living species from any of

the subfamilies. Some of these subfamilies are very diverse

and morphologically disparate and our sample is certainly

FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA

151

Table 2. Characters and character states of the foregut and shell used in cladistic analysis. See text for details of foregut characters.

* denotes characters where the states were treated as unordered.

Foregut characters

1. Rhynchodeal introvert

*2. Rhynchodeal sphincter

3. Accessory proboscis structure

4. Proboscis

5. Transverse muscles in rhynchodeum wall

6. Epithelium of posterior rhynchodeal

wall continuous with proboscis wall

Sphincter at distal end of buccal tube

Sphincter in middle of buccal tube

9. Sphincter at base of buccal tube

10. Buccal tube introvert (‘valvule’)

11. __ Protrusive lips of buccal tube

12. Position of buccal mass

13. Connection of radular retractors

to columellar muscle

14. Extensible buccal lips

15. Septum dividing anterior and

posterior areas of the rhynchocoel

16. Elongation of oesophagous between

buccal mass and nerve ring

17. Salivary glands

18. Salivary ducts

19. Type of salivary gland

20. Accessory salivary glands

21. Radula

22. Radular caecum

*23. Central tooth

24. Lateral teeth

25. Marginal teeth

*26. Type of solid radular teeth

*27. ‘Type of wishbone teeth

*28. Type of hollow teeth

29. Venom gland

*30. Connective tissue layer of muscular bulb

*31. Muscle layers of muscular bulb

32. Odontophore

33. | Odontophoral cartilages

Shell and opercular characters

*34. Shell form

*35. Number of protoconch whorls

*36. Sculpture of the protoconch

*37. — Siphonal canal

*38. Position of the anal sinus

39. Presence of apertural ornament

(teeth on the outer lip)

*40. Number of the teleoconch whorls

*41. Development of subsutural ramp

42. Operculum

43. Position of opercular nucleus

inadequate. Although anatomical data are available for many

terebrids (Taylor, 1990 and unpublished), most of these were

eventually excluded from the analysis for the following rea-

son. Many of the morphological trends in the Terebrinae,

involve partial to total loss of the foregut organs (Taylor,

1990); thus many of the characters used in the cladistic

analysis were recorded as missing. In our earlier attempts at

cladistic analysis, terebrid species tended to appear in rather

disparate positions on the cladograms. Consequently, we

have used only three species to represent the Terebrinae and

0 — absent, 1 — present

0 — present anterior, 1 — present posterior, 2 — absent

0 — absent, 1 — present

0 — present, 1 — absent

0 — absent, 1 — present

0 — absent, 1 — one sphincter, 2 — two sphincters

0 — absent, 1 — present

0 — basal, 1 — distally shifted

0 — present, 1 — absent

0 — absent, 1 — present

() — absent, 1 — present

0 — absent, 1 — present

0 — two/one glands present, 1 — glands absent

0 — two ducts present, 1 — one duct present

0 — acinous, | — tubular

0 — two/one glands present, 1 — glands absent

0 — present, 1 — absent

0 — absent, 1 — present

0 — robust muriciform, 1 — narrow 2 — broad with central spine

0 — comb-like, 1 — absent

0 — solid, 1 — wishbone, 2 — hollow, 3 — absent

0 — flat, 1 — curved-pointed, 2 — semi-enrolled (Hastula bacillus)

0 — large blade, small accessory limb, 1 — short knife type, equilimbed

0 — large base, 1 — thin small base

0 — present, 1 — present with changed histology in anterior portion, 2

— absent

0 — present, 1 — absent

0 — more or less equal, 1 — outer layer thin, 2 — single layer only

0 — present, 1 — absent

0 — not fused, 1 — fused

0 — fusiform, 1 — coniform, 2 — turreted, 3 — terebriform, 4 —

rounded

0 — less than two, | — more than two

0 — absent or very weak, 1 — present

0 — not differentiated, 1 — moderate, 2 — long

0 — sutural, 1 — shoulder, 2 — peripheral, 3 weak or absent

0 — absent, 1 — present

0 — less than 4, 1 — from 4 to 8, 2 — more than 9

0 — absent, 1 — present

0 — present, 1 — absent

( — terminal, 1 — mediolateral

Pervicaciinae, the taxa being the least-derived known for

each group.

Characters

We used 43 characters, coded as 101 states in the analysis. Of

these, 35 characters concerned foregut anatomy and a further

eight, the shell or operculum. The characters and their states

are listed in Table 2. Full discussion of the anatomical

characters will be found in the section of this paper concern-

152

ing foregut anatomy. Additionally, brief descriptions of the

shell characters used are given in Appendix 1.

Outgroup

The relationships of the Conoidea to other Neogastropoda

are very unclear and there is no obvious sister group. In our

various analyses we used two outgroups. The first is Bentho-

bia the most primitive non-coniodean neogastropod known

(Kantor, 1991). This gastropod has a buccal mass situated at

the base of the proboscis, a muscular connection between the

radular retractors and columellar muscles, and a full set of

glands connected with the oesophagous. Additionally, we

used as a second outgroup a hypothetical ancestral taxon

consisting of the underived states, where known, of all the

characters used in the analysis.

Method

The data were analysed using version 3.0 of the PAUP

program (Swofford, 1991). Characters 2, 7, 23, 25, 26, 27, 28,

29, 30, 31, 33, 34, 35, 36, 37, 38, 41, 42 were treated as

unordered. The matrix of taxa and character states is shown

in Table 3.

Table 3.

abbreviations.

Ancestor 0

Benthobi 0

PseudomP 0

StrictiP 0

ClavusUn 0

SplendrC 0

ClavatuD 0

ClavatuC 0

ClionelS 0

ToxicliT 0

LophiotL 0

PolystiA 0

TurricuN 0

AforiaAb 0

FunaLati 0

VexitomG 0

PilsbryN 0

MicantaP 0

BorsoniO 0

TomopleV 0

TropidoF 0

OphiodeIl 0

AnarithM 0

GlyphosC 0

EucithaS 0

MangeliN 0

MangeliP 0

OenopotL 0

PhilberP 1

PhilberL 1

DaphnelIR_ 1

GymnobeE 1

TeretioL 1

AbyssobA 0

Benthofa 0

GenotaNi 0

ThatcheM 0

TaranisM 0

ConusVen 0

PervicaT i

HastulaB 1

DuplicaC 1

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FPOoOrOrcOoOrRrPrRPrRP COCCCCCoCoCoCOoOCcCOCCCCocoocooooococoocoocoocoocoeoo

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PS} T SES) |S BS) (SY) SOS pS) (Sy SSS) SSS) SSS SS) Sy Soa] &] See. ] yn SIONS

PKS) (SS) RS) (Sy aS (SS (SSS) Se) SS) SONS SI SSSI SSS SOS)

SS) SS (Soa Sy Sea SS Se STS IS SHS SS SO SOS SS SS aS Soya] Sy SS)

ooc*aqooocooocd oo SoCo So Sooo Sco Cor oO Sc oo So So So oo >

Le ce ce ec ce ce ce ce ce ce ce re cc cc ee ce ee

LN YN aS i Jif Yi S| SN I fe) (es) (=>) ) OY) i) (SS SS NS) (SIS SS SSS) SH)

SIS SO SSO) SI S| SS aS Sea) SB SS) S/S] &) SS SSS OS SI SY SSS SY SSS SS SS]

Sy) SSS SI SSS) | eS) (SS SS SN SS SSS) SSS SSI Vea aaa) (St)

STO) TS) SS) i a (SS SSS () (CS) (SS) (SS) (SS) SS a aS Se =)

qooocoqc”ocqcnooorwnvrocorroorococooo Orr RB VRP OOrFrPrP FP OCOFrFE oO

qoqoooocooocoocooocoeoocoeoooeoooocorocooorcorrrocorcoo

ooroeocoeocoeooocooocnoooooocoocooocoOrPrRP KF Or CORP RP Re RFP OOrR FP eK oO

oOooo7yrFr oot VRE RSP DP OrRrFP RFP OOOO; oCoooocooooooooooodcdoes

Lt ct ct ec ce ee ee ee a aa)

J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV

Results of phylogenetic analysis

Although we have used many new anatomical characters, the

cladistic analysis gave rather disappointing results. The taxa

of ‘lower conoideans’ especially, were rather poorly resolved

with major branches supported by rather few weak charac-

ters. Additionally, small adjustments to the data set produced

rather large changes in tree topography and the number of

alternative trees generated.

Despite these limitations we thought it worthwhile to

present the results of our analysis, which is the first for the

Conoidea to use anatomical characters. Future work will

extend on the character set shown in Table 3 and hopefully

improve the resolution of the analysis.

A heuristic search with the matrix shown in Table 3 and

with Ancestor as outgroup, produced over 900 equally parsi-

monious trees (189 steps; consistency index 0.296;

homoplasy index 0.704). A 50% majority rule consensus

cladogram derived from these trees is shown in Fig. 27.

Despite the large number of trees generated, most of the

trees are very similar to each other and most of the branches

are supported in 75—100% of the trees. Autapomorphies of

the internal nodes are listed in Table 4.

The least-derived group are the two species of Drilliidae,

Matrix of taxa and character states used in the analysis. See Table 2 for further details of characters and caption to Fig. 27 for taxon

SS Sri Oise FS So SoS So SoS SiS oor oo oS ooo oS So So Soo or > >

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RD RD V VY VV VV VV VV VV VV VY VV VV VV VV VV VV VV VV VOOR RP Ro

POO ES OO OOO OO ORDO OMEN EC DO EEC DODO DOM NCE NC MERC NC EENC Ml an anil Wl lO Ee lO CEC EC EC)

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WBWHWrFARNOCOHFH HHH HAHA HONHROHR HE ROCCOCHE HCOOH HERE HY

OR OF CORR YVR RP RP RP ROR RP RP RP RP OOrRCORFR OVOOCOCOCOoOCcCooocooOorHOor

SOoocorrooy RP rr FP PrP rPrPrPoCoOoOCoOorcoooco yOrFcocococooocoo-+,

SOCOCOR RP RP RP NFR OR RP EP RP RP ORF OR POR RP OR REP NNNNKEF ORF OCCOFR OCF

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Seooo7yry VOM VY VY VVONVVV VY VOOCOCOCOCOROCOORRPHRE RH OOCOCSO

PONONOOONNNONCTOCTOCCOCCOCCOCOCOORPRP RP OR OKF ORF OOCONONO

WWWWR OF OOOO OW WF RRR RD RRR DR RR BB DNR RRP ee eee Dm WD

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NMIVVR VORrRP VY VMOOVOOCOCC OF OF OR ROR II VI V VR UV VV YY

FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA

Table 4. Synapomorphies for interior nodes. Nodes numbered as

in Fig. 27.

Node Synapomorphies (Character: state)

1 7(1), 20(1), 35(0), 37(0), 38(1)

2 13(1), 23(2), 24(1), 26(1)

Si 1(1), 3(1), 34(3), 38(3), 40(2)

4 4(1), 14(1), 15(1), 29(1)

5 25(1), 37(1)

6 6(1), 16(1), 41(1)

7 7(0), 23(0), 25(0)

8 12(1)

9 33(1), 43(1)

10 7(2), 29(1)

11 12(1)

12 34(0), 40(0)

13 14(1), 27(0), 43(0)

14 7(1), 35(1)

16 34(0), 37(2)

17 2(1), 38(2), 40(2)

18 22(1), 25(2), 32(1)

20 20(0)

21 2(1), 34(0)

22 2(1), 14(1), 28(0)

23 18(1)

24 14(0), 35(1), 39(1), 42(1)

75) 7(1), 35(1)

26 8(1), 34(0)

27 20(0), 39(0)

28 19(1), 36(1)

29 28(0), 42(1)

30 10(1), 11(1)

31 6(1), 7(1), 14(1), 310)

32 30(1), 31(2), 38(0)

33 1(1), 14(1), 21(1), 391)

34 38(3)

35, 4(1)

36 5(1), 29(2), 39(0), 41(1)

Bi) 17(1), 40(0)

38 1(0), 35(0)

which are the only conoideans possessing five teeth in each

radular row. They also retain the connection of the radular

retractor muscle to the columellar muscle. Their distinctive

apomorphy is the possession of large, comb-like lateral teeth.

We have studied only three species in this group (the third

species identical to Clavus unizonalis) which are very similar

to each other. However, we note the very different hollow,

enrolled ‘hypodermic-style’ marginal teeth of IJmaclava

(Shimek & Kohn, 1981) and the possible ‘wishbone’ margin-

als of Drillia roseola (McLean, 1971). Anatomical studies of

these taxa are needed to determine their status.

All other conoideans are separated from the Drilliidae at

Node 2 by the loss of the radular retractor/columellar muscle

connection, by the loss of the lateral teeth and possession of

curved pointed marginal teeth. None of the non-drilliid taxa

that we have included in the cladistic analysis possess lateral

teeth, although what appear to be vestigial lateral teeth are

seen for example in Antiplanes (Kantor & Sysoev, 1991) and

a few other species. Also, it is possible that the broad central

teeth seen in Cochlespirinae may be formed by fusion of

lateral teeth. Another apomorphy at this node is the posses-

sion of a broad central tooth with a spine-like central cusp.

153

Node 3 separates the Terebridae, with five apomorphies

including the possession of a rhynchodeal introvert and the

accessory proboscis structure. The Pervicaciinae (Node 4) are

separated from Hastula (representing the Terebrinae) by the

loss of the proboscis, the presence of extensible buccal lips, a

septum in the rhynchocoel (although this is present in some

Terebrinae) and the loss of the venom gland.

Node 5 separates all other conoideans with two apomor-

phies namely the presence of wishbone marginal teeth and a

moderately long siphonal canal. The latter is a weak charac-

ter and although we consider the fomer to be a strong

character, some taxa in Clade 6 have solid teeth which PAUP

considers a reversal from the wishbone condition.

Clade 6 comprises taxa with the epithelium of the posterior

part of the rhynchodeum continuous with that of the probos-

cis and with an elongated loop of oesophagus anterior to the

nerve ring.

Clade 7 includes two taxa with solid marginal teeth and no

buccal tube sphincter and Tovxiclionella which has hollow

teeth. PAUP treats the solid teeth as a reversal, but we think

that this is unlikely. However, it is possible that the ‘flanges’

on the teeth of Strictispirinae may be modifications of a

second limb on the tooth. Toxiclionella and Strictispira are

grouped together at Node 8, because both have a buccal mass

situated at the distal end of the proboscis. However, Toxi-

clionella shares many characters with the Clavatulinae

(including the medio-lateral nucleus of the operculum), but

has a very different radula with hollow and barbed marginal

teeth firmly attached to the radular ribbon located in the

distal buccal mass. Although Toxiclionella tumida lacks a

central tooth, a clavatuline type central is known in T. elstoni

(Kilburn, 1985). Turricula nelliae (Node 12) shares many

apomorphies with clavatuline species and should be trans-

fered from the Cochlespirinae to the Clavatulinae.

PAUP suggests that Funa and Vexitomina (Crassispirinae)

and Pilsbryspira (Zonulispirinae) are derived from the Clav-

atulinae. They share a number of characters, but Funa and

Vexitomina have distinctive wishbone teeth with one broad

flat limb and a small, thin, subsidiary limb. Pilsbryspira has

enrolled marginal teeth and a distal buccal mass. This type of

tooth could be derived by enrollment of the crassispirine type

of wishbone tooth. Both groups have an operculum with a

terminal nucleus which PAUP treats as a reversal from the

medio-lateral nucleus of the Clavatulinae.

Lophiotoma and Polystira (Turrinae) (Node 16) have a

peripheral anal sinus and a posteriorly situated rhynchodeal

sphincter. Aforia has an accessory salivary gland and PAUP

treats this appearance as a reversal, the glands having already

been lost between the outgroup and the first node. However,

it is highly unlikely that these glands are regained once lost.

Accessory glands have a very patchy distribution amongst the

Conoidea (Conus, Benthofascis and some Clathurellinae) and

apart from their occurrence in some terebrids, Aforia is the

only ‘lower’ conoidean in which we have seen the glands. The

distribution of this character should become clearer as more

species are examined. Maybe significant, is the fact that

Aforia is the only other conoidean in which the multidigitate

osphradial leaflet typical of Conus has been found (Sysoev &

Kantor, 1988 fig. 2J).

From Node 18 onwards are all the so-called ‘higher’

conoideans, which in all our analyses form a monophyletic

group. The apomorphies which define this node are the

presence of a radula caecum for storage of detached radular

teeth, hollow, enrolled marginal teeth, loss of the radular

154

15 19

28 30

J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV

Ancestor

Clavus U "|

Splendrillia C

Hastula B

Pervicacia T

Drilllidae

| Terebridae

ii Duplicaria C

Pseudomelatomidae

Clavatulinae

Strictispiridae

Pseudomelatoma

Toxiclionella T

8 Strictispira P

Clavatula D

Clionella C

Clavulata S

12 Turricula N

Clavatulinae

Funa L

Crassispirinae

Vexitomina G p

14 Pilsbryspira N = =——— Zonulispirinae

Aforia A Cochlespirinae

Lophiotoma L

17 Polystira A | Turrinae

Micantapex P

Borsonia O ] Clathurellinae

21 Ophiodermella

Oenopota L Oenopotinae

Tropidoturris F

Anarithma M | Clathurellinae

24 Glyphostoma C

Genota WN ;

Benthofascis | Conorpinae

Zl Conus V Coninae

Tomopleura V Clathurellinae

Eucithara S

Mangelia N | Mangeliinae

31 Mangelia P

Thatcheria M

Philbertia P

Philbertia L

Daphnella R Daphnellinae

Gymnobela E

Teretiopsis L

Abyssobela A

38 Taranis M Taraninae

Fig. 27 Majority-rule (50%) consensus tree. Autapomorphies for each node given in Table 4. Higher taxa names at the top of branches

reflect our new classification. Taxon abbreviations in order top to bottom on the tree: Clavus U = Clavus unizonalis, Splendrillia C =

Spendrillia chathamensis, Hastula B =Hastula bacillus, Duplicaria C = Duplicaria colorata, Pseudomelatoma P = Pseudomelatoma

penicillatus , Toxiclionella T = Toxiclionella tumida, Strictispira P = Strictispira paxillus, Clavatula D = Clavatula diadema, Clionella S =

Clionella sinuata, Clavatula C = Clavatula caerulea, Turricula N = Turricula nelliae, Funa L = Funa latisinuata, Vexitomina G =

Vexitomina garrardi, Pilsbryspira N = Pilsbryspira nympha, Aforia A = Aforia abyssalis, Lophiotoma L = Lophiotoma leucotropis,

Polystira A = Polystira albida, Micantapex P = Micantapex parengonius, Borsonia O = Borsonia ochraea, Ophiodermella I =

Ophiodermella inermis, Oenopota L = Oenopota levidensis, Tropidoturris F = Tropidoturris fossata, Anarithma M = Anarithma metula,

Glyphostoma C = Glyphostoma candida, Genota N = Genota nicklesi, Benthofascis = Benthofascis biconica, Conus V = Conus

ventricosus, Tomopleura V = Tomopleura reevei, Eucithara S = Eucithara stromboides, Mangelia N = Mangelia nebula, Mangelia P =

Mangelia powisiana, Thatcheria M = Thatcheria mirabilis, Philbertia P = Philbertia purpurea, Philbertia L = Philbertia linearis, Daphnella

R = Daphnella reeveana, Gymnobela E = Gymnobela emertoni, Teretiopsis L = Teretiopsis levicarinatus, Abyssobela A = Abyssobela

atoxica, Taranis M = Taranis moerchi.

ribbon and loss of the odontophore.

Clade 19 is made up of various taxa formerly included in

the Borsoniinae and Clathurellinae with the addition of

Oenopota (Oenopotinae). The apomorphies defining the

nodes are very unsatisfactory with many reversals. More

characters need to be analysed in these taxa to achieve better

resolution.

Borsonia and Ophiodermella (Node 21) have posteriorly

situated rhynchodeal sphincters, and fusiform shells. The

taxa in the other clade (Node 22) have extensible buccal lips

and hollow radular teeth with large bases. Although the

Oenopotinae have been previously thought to have close

affinities with the Mangeliinae, they do have acinous salivary

glands, rather than the tubular type associated with the latter

subfamily.

A clade comprising Anarithma and Glyphostoma is defined

(Node 24) by three characters; a posteriorly situated rhyn-

chodeal sphincter, a single salivary duct and apertural orna-

FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA

ment. Glyphostoma has long slender radular teeth and has

been separated in the family Clathurellinae (McLean, 1971).

Anarithma has been classified in the Diptychomitrinae (=

Mitrolumninae), but Kilburn (1986) could see no significant

differences from the Borsoniinae.

Taxa normally classified in the Borsoniinae (Ophioder-

mella, Borsonia, Tomopleura, Micantapex, Tropidoturris and

Anarithma) do not form a monophyletic group in any of our

analyses. For this reason, in the classification derived form

this study we are leaving these taxa, along with Glyphostoma

and others in informal groupings within the subfamily

Clathurellinae.

Benthofascis (Conorbinae) and Conus (Coninae) (Node

27) share a number of characters. They lack an anterior

sphincter to the buccal tube, but have have an intermediate

sphincter instead. Both have accessory salivary glands and

retain an operculum. Additionally, both genera show resorp-

tion of the inner shell whorls. Although Genota (Node 26) is

usually classified in the Conorbinae, it lacks an operculum.

Taxa from Node 28 onwards have tubular salivary glands

and most have sculptured protoconchs. The Mangeliinae

(Node 30), represented by Eucithara and Mangelia, are a

well-defined group with the distinctive buccal tube introvert,

and protrusive lips of the buccal tube. Taxa from Node 32

have a muscular bulb made up of only one muscle layer and

lacking the connective tissue layer, with additionally, an anal

sinus located at the suture. Thatcheria (Node 32) has many

characters in common with the Daphnellinae and until many

more daphnellines have been examined anatomically it can

be classified with them. However a great range of foregut

anatomy is found in the Daphnellinae and it may be that the

group is paraphyletic. At the extreme end of the tree (Node

36) are taxa which have lost many foregut characters such as

radula, proboscis and glands. Taranis has been classified in a

separate subfamily Taraninae (Kantor & Sysoev, 1989), but

it has so few characters that its relationships are obscure. It

may be a highly derived daphnelline.

Conclusions

Our studies have shown that several major autapomorphies

associated with the Conoidea have developed independently

in separate clades. Also there has been parallel loss of foregut

structures. Some of the more important of these are briefly

discussed below.

Hollow, enrolled ‘hypodermic style’ radular teeth are con-

sidered a distinctive feature of the conoidean feeding mecha-

nism. Our analysis shows that hollow teeth have been

independently derived at least five times in the evolution of

the Conoidea. In /maclava the hollow marginal teeth seem to

have developed from the enrolling of the flattened drilliine-

type of marginal teeth. In Toxiclionella, the hollow teeth

were derived from wishbone teeth similar to those of Clav-

atula or maybe from solid teeth like those of Pseudome-

latoma. Hollow teeth are found in many Terebridae and are

thought to have been derived from solid teeth via semi-

enrolled intermediate forms such as found in Hastula bacillus.

The enrolled teeth of Pilsbryspira (Zonulispirinae) may have

been derived by enrolling of the crassispirine type of wish-

bone tooth. The hollow teeth of the higher conoideans such

Clathurellinae, Coninae, Mangeliinae and Daphnellinae in

| all their various forms may represent another separate deriva-

| tion. The radular caecum found in some Terebridae was

derived independently of that found in the higher turrids

155

(Clathurellinae, Oenopotinae, Mangeliinae, Daphnellinae)

and Coninae.

The rhynchodeal introvert found in some Daphnellinae, is

also found in all Terebridae (including pervicaciines). If our

ideas concerning the relationships of the Terebridae are

correct, then the structure was evolved independently in the

two groups.

A buccal mass situated at the base of the proboscis is

considered to be a diagnostic character of the Conoidea

(Ponder, 1973). However, in Turricula nelliae the buccal

mass was shown to be located at the distal end of the

proboscis (Taylor, 1985; Miller, 1990). We now know that a

distally-shifted buccal mass seems to be common feature of

the Clavatulinae and is found also in Pilsbryspira

(Zonulispirinae) and Strictispira (Strictispirinae) which lacks

the venom apparatus.

One surprising trend seen in at least four clades is the loss

of the venom apparatus. In the Daphnellinae, Taraninae and

some Terebrinae this is associated with the loss of the

proboscis and radular apparatus. Pervicaciinae have a well

developed radula apparatus but no proboscis or venom gland.

By contrast, Strictispira which also lacks the venom gland, has

a proboscis, a distally-located buccal mass and a robust radula

apparatus.

Relationships and status of Terebrinae and

Pervicaciinae

Some controversy concerns the status of the Terebrinae and

Pervicaciinae. Rudman (1969) and Taylor (1990) suggested

an independent origin for the two groups. However, anatomi-

cal studies of more species is revealing some shared apomor-

phies which suggest a common origin.

Although both subfamilies possess elongate multi-whorled

shells there are large anatomical differences between the two

groups. The family Pervicaciidae was orginally proposed by

Rudman (1969) for Pervicacia tristis, a terebriform species

with no proboscis and venom apparatus, but with an odonto-

phore and a radula with a strong membrane and two sickle-

shaped, solid teeth in each row. It is now known, that many

more ‘terebrids’ (Duplicaria species and others) share these

characters and should be included in the family (Taylor,

1990). Other characters of pervicaciids include a rhynchodeal

introvert and a septum in some species.

Most of the radulate Terebrinae s.s. possess hollow and

barbed, radular teeth, similar to those seen in Conus and the

Clathurellinae. However, some Hastula species possess an

odontophore and Hastula bacillus has partially-solid teeth

(Taylor & Miller, 1989). This discovery demonstrates that the

Terebridae must be derived from a lower conoidean with an

odontophore and radular ribbon, rather than from some

group such as the Clathurellinae, which have lost these

structures.

The accessory proboscis structure is an unusual organ

found in some Terebrinae, and is known from Hastula

bacillus, H. aciculina, H. imitatrix, H. raphanula, Terebra

affinis and T. pertusa (Miller 1971, Taylor , 1990; Auffenberg

& Lee, 1988; Taylor, unpub.). Some terebrines, for example

Terebra subulata, also possess a septum dividing the rhyn-

chocoel (Miller, 1971; Taylor 1990). We have found an

accessory proboscis structure in the western Australian spe-

cies Duplicaria kieneri, and Duplicaria colorata (recently

described as a Hastula by Bratcher (1988)), which otherwise

156

have an anatomy similar to Pervicacia.

Although the pervicaciines and terebrines apparently differ

considerably in foregut anatomy, they share a a number of

characters which suggest a common origin (Table 5). The

idea that the Terebrinae and Pervicaciinae were derived

separately (Rudman, 1969; Taylor, 1990) is rejected. Charac-

ters in common between the two groups are: the elongate

multi-whorled shell, the rhynchodeal introvert, and in some

species the rhynchodeal septum and accessory proboscis

structure. Thus, we propose that the common ancestor of the

combined Pervicaciinae and Terebrinae clade would have

possessed a rhynchodeal introvert, a proboscis, an odonto-

phore, a radula with two solid, sickle-shaped, marginal teeth

in each row, a venom gland, a pair of acinous salivary glands,

a pair of accessory salivary glands, an accessory proboscis

structure and a rhynchodeal septum.

Species in the Pervicaciinae clade have lost the proboscis,

venom gland and accessory salivary glands. In the Terebrinae

clade, the solid radular teeth were transformed into semi-

enrolled and then hollow teeth. The odontophore was also

progressively lost. Species with hollow teeth have developed

a radular caecum. Other, more-derived terebrines and possi-

bly pervicaciines, have lost virtually all the foregut structures,

with the rhynchodeal introvert becoming the main feeding

organ (Taylor, 1990).

Because the radula with solid, sickle-shaped marginal teeth

and well developed odontophore, is regarded as one of the

least-derived for the Conoidea, we regard the Pervicaciinae/

Terebrinae clade as an early branch from the rest of the

Conoidea. If our hypothesis of relationships is correct, then

the hollow, barbed teeth, the radular caecum, the rhyn-

chodeal introvert, and rhynchodeal septum of the terebrids,

have been derived independently of those similar structures

found in the Daphnellinae and Clathurellinae.

Status of Conidae

Despite the distinctive shell form and high species diversity of

the group, we have little anatomical evidence to support the

separation of Conus at family-level from other higher turrids.

We propose only sub-family status for the group. Every

anatomical character-state of the conine foregut is shared

with species of Clathurellinae and Conorbinae. Some Conus

species possess a snout gland in the rhynchocoel, but this

Table 5. Comparison of character states between Pervicaciinae and

J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV

organ has been little studied. Conus species also have a

distinctive osphradium with the multidigitate leaflets (Taylor

& Miller, 1989). However, the detailed structure of the

osphradium has been studied in only a few species of Tur-

ridae, but at least in some Aforia species (Cochlespirinae)

there are similar digitate osphradial leaflets (Sysoev & Kan-

tor, 1988). The resorption of the inner shell whorls has been

used as a diagnostic character of conines (Kohn, 1990), but

the occurrence of this feature has been little studied in other

conoideans, although it is present in Benthofascis (Conorbi-

nae).

CLASSIFICATION OF CONOIDEA

Introduction

Although many of the subfamilial names (as well as apparent

synonyms) currently-used within the Turridae were intro-

duced in the 19th or early 20th century, no detailed and

well-documented classification was developed in these earlier

works. Most authors based their classifications exclusively on

shell characters, although Stimpson (1865) used radula data

and Fischer (1887) divided the Conoidea into four subfamilies

solely by opercular characters. The rather detailed classifica-

tion of Casey (1904) who recognised eight tribes within the

Turridae (Donovaniini are not conoideans), was based on

both opercular and shell characters.

Thiele (1931) classified turrids into three subfamilies con-

tained within the family Conidae, with the Terebridae as a

separate family. Diagnoses of the turrid subfamilies mainly

consisted of combinations of such characters as ‘opercu-

late-inoperculate’ and ‘toxoglossate—nontoxoglossate denti-

tion’. This was the first classification where the taxonomic

difference between toxoglossate and nontoxoglossate radulae

was definitely indicated. An elaboration of this classification

was developed by Wenz (1938) who recognised five subfami-

lies of Turridae as well as the Conidae and Terebridae.

The classification of Powell (1942, 1966) provided a great

stimulus to conoidean taxonomy, and is used, with modi-

fications, by almost all authors concerned with Turridae.

Powell recognized nine subfamilies which were based prima-

rily on shell characters, although radular and opercular

Terebrinae.

Character Pervicaciinae Terebrinae

Shell shape Multiwhorled Multiwhorled

Radular teeth Solid sickle-shaped If present, usually hollow enrolled marginals

Odontophore Present Present in some Hastula species

Radular caecum Absent Present in hollow-toothed forms

Venom gland Absent Present in all with radula & proboscis

absent in others

Proboscis Absent Present in all radulate forms

Salivary glands Present Present in many species

Accessory salivary glands Absent Present in some species

Rhynchodeal introvert Present Present

Rhynchodeal septum

Accessory proboscis structure

Eyes Absent in all?

Operculum Present

Present in some

Present

FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA

features were also used. Powell believed that the hypodermic

toxoglossate dentition could develop independently in differ-

ent lineages and, more importantly, that the appearance of

toxoglossate radula was not a significant reason for separating

groups at the subfamilial level. As a result, he classified some

taxa having quite different radular types (including both solid

and hollow marginal teeth) within a single subfamily.

Morrison (1966) followed Thiele in recognizing a funda-

mental difference between groups with solid (= nontoxoglos-

sate) and hollow (= toxoglossate) marginal teeth. He

suggested a separation at the family level using the families

Turridae (with subfamilies Drilliinae, Clavatulinae and

‘Lophiotominae or Crassispirinae’), Mangeliidae and

‘Pseudomelatominae’.

The subfamily classification of Powell was considerably

revised by McLean (1971), who adhered strictly to the

principle of grouping together genera with the same type of

radula. He also added six subfamilies to Powell’s classifica-

tion; three of these being described as new (Clathurellinae

H. & A. Adams, erroneously). Several subfamilies were

recognised (or retained after Powell) on shell characters,

but which share the same radular type, and some of these

seem to be rather poorly documented. However, McLean’s

classification which includes 15 subfamilies is at present the

most detailed and well developed.

In a continuing series of papers concerning South African

Turridae, Kilburn (1983, 1985, 1986, 1988), adopted a prag-

matic approach (Kilburn, 1983 p.550 ‘ ... any practical

subdivision is better than none .. . ’), and revised to some

extent the composition of subfamilies which he studied. He

also synonymized the Diptychomitrinae (= Mitrolumninae =

Mitromorphinae) with the Borsoniinae.

Bogdanov (1986, 1987, 1990) described a new subfamily

Oenopotinae separating the operculate Oenopota and its

relatives from the Mangeliinae. Additionally, the subfam-

ily Taraninae was recently re-instated (Kantor & Sysoev,

1989).

Some nomenclatural changes in the names and authorships

of several subfamilies were made by Cernohorsky (1972,

1985, 1987), and Ponder and Waren (1988).

A different viewpoint was taken by Bouchet and Warén

(1980) in their study of North Atlantic deep-sea Turridae.

They avoided the use of any subfamilial divisions, considering

the present classification of Turridae to be artificial and based

mainly on (p. 5) ‘ . . . more or less randomly selected shell

characters’.

At present there is no completely agreed classification of

Turridae, nor is there any agreement on which are the

taxonomically important characters. The existing variants of

turrid classification are based almost exclusively on shell,

radular and opercular features.

The Terebridae have similarly been classified mainly on

shell characters. H. & A. Adams (1853) and Cossmann

(1896) divided the Terebridae into two subfamilies, includ-

ing the Pusionellinae as the second subfamily. Pusionella is

now known to belong to the turrid subfamily Clavatulinae.

A separate family, the Pervicaciidae, was proposed by

Rudman (1969) for Pervicacia tristis. However, Bratcher &

Cernohorsky (1987) included Pervicacia and similar forms

in the Terebridae. Taylor (1990) confirmed the distinctive-

ness of Pervicacia, and showed that many other terebrids

should be included in the family Pervicaciidae.

The Conidae have long been considered as a fairly homo-

geneous group, the main problems have concerned the limits

157

of the family and whether taxa such as Cryptoconus, Conor-

bis and Genota should be included. Cossmann (1896) for

example, included them in the subfamily Conorbinae within

the Conidae, whilst Powell (1966) includes this subfamily in

the Turridae.

New classification proposed

As a result of our analysis of foregut characters throughout all

the conoidean higher taxa we propose a new classification of

the superfamily. This classification represents a rather con-

servative compromise position. Although in principle the

classification should be based upon the results of the phyloge-

netic analysis, we were constrained by the rather poor

resolution obtained with our data set. Moreover, only a

rather small subset of conoidean species have been examined

in any detail. Information from taxa not included in the

cladistic analysis (mainly radular characters) has also been

used in constructing the classification. An example of the

problem is the family Turridae, which comprises the four

subfamilies with wishbone marginal teeth, plus the

Zonulispirinae. The cladistic analysis suggests two different

clades for these subfamilies. This is certainly possible, but the

branches are supported by rather few, and perhaps weak

apomorphies. Despite the deficiencies this is the first compre-

hensive classification of the Conoidea which includes ana-

tomical characters. Below we give descriptions of shell,

radula and foregut characters for each of the higher taxa that

we recognise. Some of the taxa have only provisional status.

For example, the subfamily Clathurellinae has been divided

up into five informal groups; it may well be polyphyletic, but

we have insufficient evidence to resolve the situation. Simi-

larly, we are uncertain of the status of the Conorbinae and

Taraninae.

Summary of proposed classification

Superfamily Conoidea

Family Drilliidae (ICZN pending)

Family Terebridae

Subfamily Pervicaciinae

Terebrinae

Family Pseudomelatomidae

Family Strictispiridae

Family Turridae

Subfamily Clavatulinae

Crassispirinae

Zonulispirinae

Cochlespirinae

Turrinae

Family Conidae

Subfamily Clathurellinae

Coninae

Conorbinae ?

Oenopotinae

Mangeliinae

Daphnellinae

Taraninae ?

158

DIAGNOSES OF HIGHER TAXA

Family Drilliidae Morrison, 1966 (ICZN pending)

Shell of small to medium size (usually 15-25 mm, up to 50

mm), claviform (with a more or less high spire, and a

relatively short, truncated base). Anterior canal indistinct,

short or moderately elongate. Anal sinus on the shoulder,

rather deep, often sub-tubular when a parietal tubercle is

present. Sculpture usually well developed. Protoconch pauci-

or multispiral, smooth or, sometimes, carinate (from the

second whorl or, rarely, from the beginning). Operculum

with terminal nucleus.

RADULA. With strong radular membrane, five teeth in each

row, with in some species the complete loss of the central

tooth and reduction of the laterals. Rachidian tooth small,

with a prominent central cusp and, often, smaller lateral

denticles. Lateral teeth are typically broad and curved,

comb-like, with many small cusps the outermost being

smaller. Marginal teeth have a variable morphology from

simple and flat, sometimes with a weak accessory limb, to

enrolled. In at least one species (Jmaclava unimaculata),

marginal teeth are hollow and enrolled, whilst the radula as a

whole is similar to that of other drilliids.

FOREGUT. Proboscis moderately long, with one or two distal

sphincters and sometimes a mid-buccal tube sphincter. Buccal

mass at base of proboscis, odontophore well-developed,

cartilages either separated or fused. Two acinous salivary

glands with two ducts. No accessory salivary glands. Venom

gland with uniform histology along its length. Retractor

muscle of the radular sac passes through the nerve ring and

joins the columellar muscle.

REMARKS. The anatomy and radula are known for only a

very few species of Drilliidae. This prevents us from introduc-

ing any subfamilial classification of this possibly complex

family.

Family Terebridae Morch, 1852

Elongate, multiwhorled shells, with small quadrate to trian-

gular apertures. Siphonal canal short. Anal sinus not visible.

Shell ornament of low axial ribs and grooves, spiral grooves,

a few species with tubercles, shells often smooth and pol-

ished. Protoconch of 1.5—5 whorls. Operculum rounded with

terminal nucleus. Radula with solid sickle-shaped teeth,

hollow harpon-like teeth or absent. Rhynchodeal introvert

present. Accessory proboscis structure and rhynchodeal sep-

tum present in some species. Proboscis present or absent.

Odontophore present in some species. Radular caecum

present in some. Acinous salivary glands present. Accessory

salivary glands present in some species. Venom gland present

or absent.

Subfamily Pervicaciinae Rudman, 1969

Shells medium to large, elongate, multiwhorled, anterior

canal short, ornament low axial ribs, spiral grooves, often

with a subsutural groove. Aperture quadrate. Operculum

rounded with terminal nucleus.

RADULA. With strong radula ribbon, two rows of sickle-

shaped solid marginal teeth.

J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV

FOREGUT. Rhynchodeal introvert. Rhynchodeal septum and

accessory proboscis structure present in some species. Pro-

boscis absent. Extensible buccal lips present in some species.

Odontophore with two cartilages. Two acinous salivary

glands and ducts. Venom gland and accessory salivary glands

absent.

Subfamily Terebrinae Morch, 1852

Shells medium to large, elongate, multiwhorled. Small quad-

rate to triangular aperture. Short siphonal canal. Shells often

smooth and polished. Shell ornament of low axial and spiral

ribs and grooves.

RADULA. Where present, long, hollow marginal teeth with

narrow bases, barbed or unbarbed. Hastula bacillus has

semi-enrolled teeth with a distal solid blade. Many species

have no radula.

FOREGUT. Rhynchodeal introvert present. Rhynchodeal sep-

tum and accessory proboscis structure present in some spe-

cies. Proboscis long, medium or absent. Odontophore with

cartilages present in some Hastula species. Radula caecum

present in many radulate species. Acinous salivary glands

with two ducts usually present. Accessory salivary glands

present in some species. Venom gland present or absent in

radula-less species.

Family Pseudomelatomidae Morrison, 1966

Shells of medium to large size (35-77 mm), fusiform. Ante-

rior canal moderately elongate. Anal sinus on the shoulder.

Protoconch smooth. Operculum with terminal or subcentral

nucleus. Egg capsules dome-shaped, with an operculum.

RADULA. With strong radular membrane; three teeth in each

radular row. Rachidian is large and rectangular with a large,

curved and pointed, central cusp and smaller lateral cusps.

Marginal teeth are solid, simple and curved.

FOREGUT. Proboscis very long, no anterior buccal tube

sphincter; buccal mass basal or posterior of the proboscis

base. Oesophagus elongated between the buccal mass and

nerve ring in Pseudomelatoma. Odontophore very large with

fused cartilages. Acinous salivary glands, paired in

Pseudomelatoma, but unpaired with a single duct in Hormo-

spira. No accessory salivary glands. Venom gland with uni-

form histology.

Family Strictispiridae McLean, 1971

Shell of rather small size (usually 15-20 mm), claviform.

Anterior canal short or indistinct. Sculpture well developed.

Deep subtubular sinus is situated on the concave shoulder

and bordered with well developed parietal callus. Protoconch

smooth, multispiral. Operculum with terminal nucleus.

RADULA. with strong radular membrane; 2 teeth in each

row, central and lateral teeth absent (latter maybe diapha-

nously on optical preparations). Marginal teeth solid, awl-

shaped, with pointed tips, a broad base and a mid-tooth

flange .

FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA

FOREGUT. Proboscis short; buccal mass located near the

proboscis tip, odontophore very large and muscular with

separate cartilages. Acinous salivary glands small and paired,

no accessory salivary gland, no venom apparatus.

REMARKS. This small family possesses unique radular teeth

and anatomy, but study of further material is necessary.

Family Turridae H. & A. Adams, 1853

Radula always with a membrane with either 3 radular teeth in

a row (central being small or weak), 4 (central lost, laterals

diaphanous) or with only marginals. Marginal teeth usually

wishbone type, rarely enrolled and hollow. Odontophore

always present. Radular sac not subdivided into short and

long arms. Venom gland always present. Salivary glands

always acinous. Accessory salivary gland either present or

absent. Operculum present.

Subfamily Clavatulinae Gray, 1853

Shell medium-sized (usually 15-30 mm, maximum 60 mm),

variable in form. Anterior canal moderately long, sometimes

short or trun-cated. Whorls usually adpressed below the

suture. Anal sinus located on the shoulder slope, rather deep

but sometimes indistinct. Protoconch smooth, of 1.5-3

whorls. Axial sculpture predominates or the sculpture is

subobsolete and the shell surface is glossy. Operculum ovate,

with medio-lateral nucleus. Egg capsules lens-shaped, verti-

cally orientated, without an operculum. Capsules attached to

the substratum by a stalk on the edge.

RADULA. Strong radular membrane with 3 to 2 teeth in each

row. Central tooth with large, very thin, inconspicuous, basal

plate and centrally thickened area with a single cusp. Central

tooth sometimes absent (Toxiclionella s.s.). Lateral teeth

absent. Marginal teeth usually robust wishbone type; hollow

harpoon-shaped and barbed in Toxiclionella.

FOREGUT. Epithelium of posterior rhynchocoel not glandu-

lar and continuous with proboscis. Moderately long proboscis

with 1 or 2 anterior buccal tube sphincters. Protrusive lips of

the buccal tube may be present (Turricula). Buccal mass

distal except Clavatula diadema in which it is basal but lies

within the proboscis. Odontophore medium to small in size,

cartilages unfused (except in Toxiclionella). Salivary glands

acinous, usually paired. Single salivary duct in Clavatula

caerulea. Single accessory salivary gland in Toxiclionella.

Anterior venom gland ciliated. Oesophagus elongated

between buccal mass and nerve ring.

REMARKS. Some species in this subfamily possess hollow

‘toxoglossate’ radular teeth associated with strong radular

| membrane, sometimes, with central teeth. The anatomy and

_ conchological characters of ‘toxoglossate’ clavatulines are,

| however, quite similar to those of ‘nontoxoglossate’ ones.

_ Thus at present we do not consider the appearance of

hollow teeth in Toxiclionella to be a taxonomic character of

subfamilial importance and therefore follow Kilburn (1986)

| in classifying Toxiclionella with other clavatulines.

The genus Turricula Schumacher, 1817 appears very simi-

lar to clavatulines in both radular characters and anatomy

(the distal buccal mass, ciliated anterior venom gland,

159

elongated oesophagus between the buccal mass and nerve

ring). Moreover, it is also similar to clavatulines in shell

characters and in its operculum with mediolateral nucleus.

On the other hand, Turricula differs in both shell and

anatomical characters from those of other ‘Turriculinae’.

Thus we transfer this genus, as well as Makiyamaia which has

similar characters, to the subfamily Clavatulinae.

Subfamily Crassispirinae Morrison, 1966

Shell of medium to small size (usually 10-20 mm, sometimes

up to 70 mm), claviform to fusiform. Anterior canal usually

short. Anal sinus on the whorl shoulder, parietal callus above

the sinus often well developed. Spiral and axial sculpture

often strong. Protoconch usually paucispiral, initially smooth,

later sometimes with axial (rarely spiral) folds. Operculum

with terminal nucleus.

RADULA. Strong radular membrane and 4, 3 or 2 teeth in

each row. Central tooth when present (Turridrupa) is thin,

quadrate and unicuspate, lateral teeth usually absent but

weak and vestigial in Crassispira and Crassiclava. Marginal

teeth, robust wishbone type or long flat teeth with a slender

accessory limb. Ptychobela has hollow teeth formed from two

components.

FOREGUT. Proboscis moderately long with two anterior buc-

cal tube sphincters. Epithelium of posterior rhynchocoel

continuous with proboscis (Funa latisinuata). Buccal mass

situated at the proboscis base in its contracted state. Odonto-

phore medium to small, with fused cartilages. Salivary glands

acinous, fused, ducts paired. Anterior venom gland ciliated in

some species. Oesophagus elongated behind buccal mass in

some species.

REMARKS. This most large and diverse subfamily of Turridae

is defined chiefly on shell and radular characters (i.e. rather

small claviform shells with wishbone radular teeth). Data on

the anatomy of its representatives are still unsufficient to

decide certainly whether the subfamily is of mono- or poly-

phyletic origin.

Subfamily Cochlespirinae Powell, 1942

Shell of medium to large size (usually 20-40 mm, up to 100

mm), narrow to broadly fusiform or pagodiform. Anterior

canal moderately elongate, rarely short or very long. Sculp-

ture variously developed, often with smooth shoulder, and

usually with rather short axial ribs below the shoulder, and

spiral riblets. Anal sinus usually deep, situated on the shoul-

der (sometimes on its lower part). Protoconch usually multi-

spiral, smooth or, sometimes, initially smooth and carinated

or spirally or axially lirate on subsequent whorls. Operculum

with terminal nucleus.

RADULA. Strong radular membrane, with three, four? (see

discussion of radula p. 135) or two teeth in each row. Central

tooth weak, unicuspid or absent. Marginal teeth of robust

wishbone type.

FOREGUT. Proboscis usualy long, with one or two anterior

buccal tube sphincters. Buccal mass basal, muscular buccal

lips may be present or absent. Odontophore small, cartilages

4, 2 or absent, fused or separate. Salivary glands acinous,

160

paired or fused. Single accessory salivary gland in Aforia.

REMARKS. Since the type-genus of the subfamily Turriculi-

nae, Turricula Schumacher, 1817, is transferred to the Clav-

atulinae (see above), the next available name for this group is

Cochlespirinae Powell, 1942.

Subfamily Zonulispirinae McLean, 1971

Shells rather small (15-25 mm), claviform. Anterior canal

usually short, sometimes moderately long. Predominantly

spiral scuplture, well developed. Protoconch multispiral, ini-

tially with smooth whorls, then with oblique axial riblets.

Anal sinus on the shoulder, often sub-tubular, with well

developed parietal callus. Operculum with terminal nucleus.

RADULA. With strong membrane and marginal teeth in each

row. Teeth semi-enrolled, to rolled, hollow teeth with narrow

base. Tips may be barbed or unbarbed.

FOREGUT. Proboscis long, with a single distal buccal tube

sphincter. Buccal mass distal. Odontophore small with two

unfused cartilages. Buccal lips present. Salivary glands fused.

Anterior of venom gland ciliated. Oesophagus elongated

between the buccal mass and nerve ring.

Subfamily Turrinae H. & A. Adams, 1853

Shell usually of medium to large size (up to 110 mm),

fusiform. Anterior canal elongated and narrow, rarely trun-

cated. Anal sinus on the whorl periphery. Axial sculpture

weak or absent. Protoconch smooth in its initial part, subse-

quent whorls axially costate; paucispiral protoconchs smooth.

Operculum with terminal nucleus. Egg capsules dome-

shaped, operculate.

RADULA. Strong radular membrane, 2-3 teeth in each row.

Central tooth either well-developed, small or absent, quad-

rate to rectangular with a strong central cusp. Lateral teeth

absent. Marginal teeth of robust wishbone type.

FOREGUT. Proboscis moderately long, rhynchostomal

sphincter posterior, a single distal buccal tube sphincter,

protrusive lips of buccal tube present. Buccal mass basal.

Odontophore small with fused cartilages. Salivary glands

paired. No accessory salivary glands. Anterior part of venom

gland ciliated.

Family Conidae Fleming, 1822

Radula consisting of hollow marginal teeth only. Radular

membrane absent. Radular diverticulum divided into short

and long arms. Odontophore absent. Radula and venom

gland may be absent. Salivary glands acinous or tubular.

Accessory salivary gland either present or absent. Operculum

either present or absent.

Subfamily Clathurellinae H. & A. Adams, 1858

Shell small to rather large, fusiform to biconic. Anterior canal

short or indistinct to moderately elongate. Sculpture pre-

dominantly spiral in most genera. Anal sinus deep to very

J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV

shallow, on the shoulder slope or on the periphery. Col-

umella with or without pleats. Protoconch usually paucispiral,

smooth, sometimes carinate or weakly spirally ribbed, rarely

axially costate on its last whorl. Operculum with terminal

nucleus present, vestigial or absent.

RADULA. Awl- or harpoon-shaped marginal teeth, without

(very rarely with) solid base, tooth cavity opens terminally at

the proximal end in vast majority of species.

FOREGUT. Proboscis short to long, 1 or 2 anterior buccal

tube sphincters, buccal mass basal. Short buccal lips in

Tropidoturris. Odontophore absent, radular caecum present

— divided by septum in Bathytoma (Micantapex). Salivary

glands tubular in Borsonia, acinous in others, paired, single

or absent. Single accessory salivary gland present in some

species. Venom gland with uniform histology. No elongation

of oesophagus.

REMARKS. This subfamily comprises species classified by

other workers in the subfamilies Borsoninae and Clathurelli-

nae. Being very variable in both anatomical and shell charac-

ters, the subfamily may be of polyphyletic origin. More

species need to be studied anatomically before any satisfac-

tory classification can be attempted. The subfamily is defined

mainly by the character of the radular teeth. Several groups

of genera can be isolated within Clathurellinae according to

shell characters.

‘Clathurellid’ group is characterized by medium-sized

shells (usually 10-25, up to 40 mm), with a moderately

elongate siphonal canal, and a well developed, often cancel-

late sculpture. Columella without pleats, but both inner and

outer lips may be denticulated; anal sinus deep located on the

shoulder. Protoconch usually multispiral, last whorls with a

pronounced medial carination and, sometimes, weak axial

lamellae on the lower half. A distinctive feature of this group

is densely granulated shell surface of most genera (except of

one subgenus of Glyphostoma and, probably, Nannodiella).

Operculum absent. Radular teeth long and slender, slightly

curved, without a solid base.

‘Bathytomid’ group. Shell of medium to rather large size

(usually 20-30, up to 70 mm), more or less biconic. Sculpture

usually well developed, entirely spiral, ribs often gemmulated

by growth lines; typically there is a peripheral tuberculated

flange. Anal sinus rather deep, located on the whorl periph-

ery. Columellar pleats strong to obsolete. Protoconch of

1.5-3 whorls, smooth or minutely papillated. Operculum with

terminal nucleus. Radular teeth either long, with more or less

terminal opening, or short, with large cylindrical solid base

and lateral opening.

‘Borsoniid’ group. Shell of rather small to medium size

(usually 15-25, up to 62 mm), fusiform. Anterior canal

moderately elongate, sometimes long. Both spiral and axial

sculpture may be present. Columellar pleats weak or absent.

Anal sinus on the shoulder slope. Protoconch of 1-2 smooth

whorls. Operculum fully developed, small or absent. Radular

teeth long, without solid base, open terminally, or, rarely,

short, with large cylindrical base, open laterally. Egg capsules

dome-shaped, with an operculum.

‘Mitromorphid’ group. Shell small (usually 4-8, up to 17

mm), biconic and ‘mitriform’. Anterior canal very short or

indistinct. Aperture narrow, columella with or without teeth,

outer lip usually denticulated, anal sinus shallow and subsu-

tural. Sculpture predominantly or entirely spiral. Protoconch

of 1.5-2 smooth whorls. Operculum absent. Radular teeth

FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA

rather short, of ‘candle flame’ shape, open terminally.

‘Tomopleurid’ group. Shell rather small to medium sized

(6-7 to 37 mm), claviform, with flattened whorls. Anterior

canal short. Anal sinus on the shoulder or just below it,

moderately deep. Columellar pleats absent. Sculpture

entirely spiral (except often raised growth lines), consisting of

well developed ribs or heavy keels. Protoconch pauci- or

multispiral. In the former case it is smooth or with minute

spiral striae or papillae, sometimes carinated; in the latter

case first 1-3 whorls with the same sculpture, later ones with

axial ribs and, sometimes, minute spiral striae. Operculum

with terminal or eccentric nucleus, sometimes absent. Radu-

lar teeth short or long and slender, without solid base, open

terminally.

Subfamily Conorbinae De Gregorio, 1890

Shell of medium size (up to 40 mm), biconic. Anterior canal

short, aperture long and narrow. Sculpture entirely spiral

except the growth lines. Anal sinus on the shoulder or almost

sutural, relatively deep. Protoconch multispiral, smooth or

spirally striated on later whorls. Operculum present or,

absent in Conorbis.

RADULA. Hollow, marginal teeth with barbed tips and nar-

row bases (Conorbis, Thiele, 192 fig 460; Benthofascis,

Powell, 1966, fig. 125).

FOREGUT. These observations are based on Benthofascis.

Rhynchostomal sphincter posteriorly situated. Proboscis

moderately long, not folded telescopically as in Conus. Distal

sphincter of buccal tube absent, intermediate sphincter

present. Middle part of buccal tube lined with glandular

epithelium. Single acinous salivary gland with two ducts.

Single accessory salivary gland. Venom gland with uniform

histology, muscular bulb with two muscular layers. No snout

gland.

REMARKS. The status of this subfamily is uncertain due to

lack of any anatomical information on Conorbis. We have

excluded Genota on the basis of shell morphology and the

absence of the operculum.

Subfamily Coninae Fleming, 1822

Shell of medium to large size (usually 30-50 mm, up to more

than 120 mm), biconic to conic. The inner shell walls are

partially resorbed. Anterior canal short, aperture usually

narrow, parallel-sided. Sculpture entirely spiral, usually weak

or obsolete, sometimes tubercules on the shoulder. Anal

sinus on the upper shoulder or almost sutural, shallow to

relatively deep, occupying a rather narrow zone. Protoconch

multispiral, smooth or spirally striated. Operculum small,

with terminal nucleus, rarelyabsent. Egg capsules, bilaterally

flattened, vasiform, arranged in clusters.

RADULA. Radular teeth harpoon-shaped, barbed or

| unbarbed on the tips, without solid base, usually open

terminally (rarely laterally) at the base.

FOREGUT. Proboscis moderately short and folded in con-

tracted state. Rhynchostome lacks definite sphincter and

rhynchodaeum can be greatly expanded to form a rostrum in

fish-feeding species. Radial muscles lie in rhynchodeal wall.

161

Snout gland present in many species. Distal buccal tube

sphincter absent, intermediate sphincter present. Middle part

of buccal tube is lined with glandular epithelium. Buccal mass

basal. Single acinous salivary gland with one or two ducts.

Single accessory salivary gland. Venom gland of uniform

histology, muscular bulb often with many muscular layers.

Subfamily Oenopotinae Bogdanov, 1987

Shell of small to medium size (usually 10-15, up to 30 mm),

oval to fusiform. Anterior canal rather short. Both spiral and

axial sculpture well developed. Anal sinus on the shoulder,

shallow, and often indistinct. Protoconch paucispiral, pre-

dominantly (sometimes entirely) spirally sculptured. Opercu-

lum with terminal nucleus present, vestigial, or rarely absent.

Egg capsules dome-shaped, with an operculum.

RADULA. Radular teeth with rounded or cylindrical solid

base and hollow shaft, sometimes with barbed tip; rarely

teeth vestigial; tooth cavity opens laterally between the shaft

and the base.

FOREGUT. Proboscis either long, or short and folded in

contracted state. Distal sphincter present or absent. Buccal

lips large, may be inverted into the buccal cavity. Buccal mass

basal. Salivary glands paired, acinous, although shown as

tubular (probably erroneously) in Oe0enopota levidensis

Shimek (1975). Venom gland of uniform histology. Muscular

bulb with a thin outer muscular layer.

REMARKS. Species of this group were previously treated as

Mangeliinae, but were isolated as a subfamily primarily on

the basis of the presence of an operculum and a spirally

sculptured protoconch (Bogdanov, 1987, 1990). None of

these features are presently considered as being of subfamilial

importance. However, one more character was revealed in

our study, the structure of the salivary glands, which distin-

guished Oenopotinae from the Mangeliinae. We provision-

ally retain the subfamilial rank of Oenopotinae until the

systematic importance of this character becomes certain.

Subfamily Mangeliinae Fischer, 1884

Shell small (usually 5-12 mm, up to 20 mm), ovate to

fusiform. Anterior canal rather short. Both spiral and axial

sculpture well developed. Anal sinus on the shoulder, shallow

to rather deep, sometimes subtubular. Outer lip usually with

terminal varix, sometimes denticulate. Protoconch smooth or

variously sculptured. Operculum absent. Egg capsules dome-

shaped, with an operculum.

RADULA. Radular teeth hollow with a solid base, sometimes

with a semi-enrolled shaft; tooth canal opens laterally.

FOREGUT. Proboscis moderately long, with a single or no

distal sphincter, intermediate and posterior sphincters some-

times present. Buccal tube introvert (‘valvule’) present. Dis-

tal lips of buccal tube can be inverted. Buccal lips large and

can be introverted into the buccal cavity. Buccal mass basal.

Salivary glands paired and tubular, accessory salivary glands

absent. Venom gland of uniform histology; muscular bulb

usually with a thin outer muscle layer.

162

Subfamily Daphnellinae Deshayes, 1863

Small to moderately large shells (usually 5-15 mm, deep-sea

species larger, up to 95 mm). Anal sinus sutural, shaped as a

reversed-L, or on the upper shoulder and varying in depth.

Sculpture variable, usually cancellate or with predominant

spirals, and often with a smooth shoulder. Protoconch usually

multispiral, rarely paucispiral, typically diagonally cancel-

lated, although some genera have spiral or axial ribbing.

Operculum absent. Egg capsules dome-shaped operculate.

RADULA. Radular teeth with large solid base and barbed or

unbarbed tips, tooth cavity opens laterally at the base.

Radula absent in some species.

FOREGUT. Rhynchodeal introvert present in many species.

Rhynchodeal septum present in some species. Proboscis

usually short, often absent. Buccal mass basal. Radula appa-

ratus absent in many species, vestigial in Gymnobela emer-

toni. Radial muscles present in the rhynchodeal wall in

radula- and proboscis-less species. Buccal lips well devel-

oped, can be intverted into the buccal cavity. Salivary glands

paired tubular or absent. Accessory salkivary glands absent.

Venom apparatus absent in many species. In Daphnella

reeveana the anterior part of venom gland is ciliated. Muscu-

lar bulb can be single layered.

REMARKS. Although Thatcheria is sometimes classified in a

separate subfamily Thatcherinae, we failed to find any ana-

tomical or shell characters which would justify separation

from the Daphnellinae.

Subfamily Taraninae Casey, 1904

Shell very small (up to 6 mm), ovate-fusiform. Anterior canal

rather short. Sculpture well developed. Anal sinus very broad

and shallow, situated on the shoulder or immediately below

it. Protoconch paucispiral, finely spirally striated, or with

spirally aligned granules. Operculum and radula absent.

FOREGUT. Rhynchostomal sphincter absent, no _ radial

muscles in rhynchodeal wall. Proboscis absent. Buccal mass

undefined. Salivary glands absent. Venom apparatus absent.

REMARKS. This monotypic radula-less subfamily was rein-

stated (Kantor & Sysoev, 1989) because it differs in shell

characters from any other turrids lacking a radula. However,

the very simplified morphology makes the evaluation of the

status of the subfamily difficult. For the present we conserve

the subfamily, but are unsure of its status.

ACKNOWLEDGEMENTS. For gifts and loan of material without which

this study could not have been completed, we are extremely grateful

to J.H. McLean, the late V. Maes, R.N. Kilburn, F.E.Wells, I.

Loch, R. L. Shimek, P. Bouchet, S. Gofas, B. Morton, W. F.

Ponder, G. Rosenberg, and B. Marshall. Other material was

obtained from the Institute of Oceanology, Academy of Sciences,

Moscow; Zoological Institute, Academy of Sciences, St Petersburg,

Zoological Museum, Moscow State University and we are very

grateful to the curators L.I. Moscalev, A.N. Mironov, B.I Sirenko

and D.L. Ivanov for their kind permission to use this material. David

Cooper expertly prepared many of the serial sections of gastropod

foreguts. Some unpublished material was made available by John

Miller and A.I Medinskaya. The optical photomicrographs were

made by Peter York. Andrew Smith and David Reid patiently

J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV

advised on problems of phylogenetic analysis. Yuri Kantor and John

Taylor are grateful to the Royal Society, London and the Russian

Academy of Sciences, Moscow for grants which enabled them work

in London and Moscow respectively. Yuri Kantor and Alexander

Sysoev gratefully acknowledge a grant from the Soros Foundation

and the Russian Academy of Natural Sciences.

APPENDIX 1

Features of the shell

Shell characters are still important for the systematics of

Conoidea, and thus should be included in the analysis.

However, there is probably no shell character which is

diagnostic of any single group. Moreover, there has been no

analysis of the adaptive or evolutionary significance of these

shell features. Nevertheless, a few shell characters appear to

be useful for the separation of clades.

Shell shape

This character which is concerned with overall shell shape is

the most subjective. We recognise five basic shell shapes:

1, fusiform shell; 2, cone-shaped shell; 3, turreted shell;

4, terebriform shell; 5, a large group of ‘intermediate’ states,

‘biconic-fusiform’, ‘ovate-biconical’, ‘ovate-fusiform’, ‘clavi-

form’, etc. characterized by rounded outlines of the shell,

which is more or less oval in its general profile.

Number of protoconch whorls

Two types of protoconch can be recognised; the paucispiral

and multispiral. These types of the protoconch were into-

duced into turrid systematics by Powell (1942, 1966) and they

are widely used in taxonomy. Generally, this subdivision

coincides with that between planktotrophic and non-

planktotrophic modes of larval development, although there

are many exceptions to the rule among turrids (Bouchet,

1990). The character is considered as being of little phyloge-

netic importance (Bouchet, 1990), but a predominance of a

single type of the protoconch can be noted in some taxa. For

instance, most Daphnellinae and Conidae have multispiral

protoconchs, whilst the paucispiral type is a typical of the

Oenopotinae (Bogdanov, 1990). Protoconchs with 1-2 whorls

are here considered as paucispiral, and these with two or

more whorls as multispiral (Bouchet, 1990).

Sculpture of the protoconch

The pattern of protoconch sculpture has been widely used in

conoidean taxonomy since Powell (1942, 1966). Turrids have

a very wide variety of protoconch sculpture and at present,

we are unable to classify them into clearly defined types.

Thus we recognize only two major states of the character;

firstly protoconchs lacking or with only weakly defined sculp-

ture and secondly, protoconchs with well developed sculp-

ture. Some higher taxa may be characterized by the presence

or absence of protoconch sculpture. For example, the closely-

related Turricula and Clavatula usually possess a smooth

protoconch, whilst in the Turrinae it is usually axially costate.

The only type of the protoconch sculpture characteristic of a

single subfamily is the ‘diagonally cancellated’ form found

FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA

among species of Daphnellinae, although this is not present

in all species.

Length of siphonal canal

To define groups of shells with different lengths of the

anterior siphonal canal we used parameter Rsl (relative

siphonal length) of Harasewych (1981). As a result, we

recognise three states of the character; a long siphonal canal

(Rsl more than 0.39; up to 0.48 in the species studied),

moderate canal (Rsl 0.21 to 0.34), short canal (RslI less than

0.20) or not differentiated from the apertural canal.

Position of the anal sinus

The anal (labial) sinus is a characteristic feature of Turridae

and its position on the shell whorls is widely used for

characterizing species and higher taxa (Powell, 1942, 1966;

McLean, 1971). We follow Powell (1966) in recognizing 4

types of sinus position; sutural (the deepest point of the sinus

is situated near the suture), subsutural (on the whorl shoul-

der), peripheral, and poorly pronounced (or very slight).

Most turrids have a subsutural sinus; a peripheral sinus is

characteristic for all Turrinae and some Clathurellinae

(Bathytoma and related genera); a sutural sinus is common

among the Daphnellinae. A weak, almost imperceptible sinus

occurs occasionally in many subfamilies. It should be empha-

sized that sinus types are recognized by growth lines, since

the form of sinus at the outer lip of a mature shell may not be

the same as that of the immature gastropod.

Operculum

The presence of an operculum is obviously the primitive state

of the character. All ‘lower’ conoideans have a well devel-

oped operculum. Among ‘higher’ conoideans, the operculum

is absent in almost all Daphnellinae and Mangeliinae, but

retained in the Oenopotinae and Conidae. In Clathurellinae

(incorporating Borsontinae), the operculum may be present,

vestigial or absent, even in apparently closely-related genera

(McLean, 1971).

Position of opercular nucleus

The opercular nucleus is usually situated in a terminal posi-

tion at the tip of the operculum, but in the Clavatulinae and

Turricula it is located medio-laterally.

Presence of apertural armament

The aperture of conoidean shells may be without ornament

on the outer lip or columella, or they may bear weak to strong

denticles, plications and folds. Armed apertures are found in

the subfamilies Mangeliinae, Clathurellinae and Daphnelli-

nae, and mostly amongst tropical shallow-water species.

Number of teleoconch shell whorls

We recognize three types of shells by this character. 1. shells

with a small number of whorls (4 and less); 2. with an

intermediate number of whorls (5 to 8); 3. with many whorls

(9 and more)

163

Presence of well developed subsutural ramp

A subsutural ramp, (a morphologically distinct, often flat-

tened part of the whorl profile immediately below the suture)

may be either absent, or pronounced, in many subfamilies of

Turridae. Usually, this character is clearly shown by a change

in both spiral and axial sculpture in this region of the whorl.

APPENDIX 2

Genus-group taxa of recent Turridae S.L.

(Compiled by A.V. Sysoev)

The list presented below is of Recent taxa of the genus-group

of Turridae s.l. distributed in respect to the classification

adopted in the present paper. Since all the data concerning

genera described before 1966 were given in Powell’s (1966)

monograph, type-species and bibliographic citations are

included only for genera and subgenera described after 1966.

Synonymy is also given only when it differs from that adopted

by Powell.

The classification used is to a great extent conservative; we

avoid the description of new taxa and radical changes in the

existing classification. As a result, some genera are of

‘unclear’ taxonomic position and cannot be assigned, despite

anatomical information, to any existing subfamily (Toxico-

chlespira, for example). Some other genera (such as Genota)

are only provisionally included into a certain subfamily.

There are 337 valid Recent genera and subgenera.

Family DRILLIIDAE Morrison, 1966. ICZN pending

Agladrillia Woodring, 1928

Eumetadrillia Woodring, 1928

Bellaspira Conrad, 1868

Calliclava McLean, 1971

Veliger 14(1): 117

Cymatosyrinx palmeri Dall, 1919

Cerodrillia Bartsch & Rehder, 1939

Lissodrillia Bartsch & Rehder, 1939

Viridrillia Bartsch, 1943

Clavus Montfort, 1810

Plagiostropha Melvill, 1927

Cymatosyrinx Dall, 1889

Drillia Gray, 1838

Clathrodrillia Dall, 1918

Elaeocyma Dall, 1918

Globidrillia Woodring, 1928

Horaiclavus Oyama, 1954

Anguloclavus Shuto, 1983

Mem. Fac. Sci. Kyushu Univ., ser.D (Geol.) 25(1): 9-10

Mangilia multicostata Schepman, 1913

Cytharoclavus Kuroda & Oyama in Kuroda, Habe & Oyama, 1971

The sea shells of Sagami Bay: 213

Pleurotoma (Mangilia) filicincta Smith, 1882

Imaclava Bartsch, 1944

Tredalea Oliver, 1915

Kylix Dall, 1919

Leptadrillia Woodring, 1928

164

Neodrillia Bartsch, 1943

Orrmaesia Kilburn, 1988

Ann. Natal Mus. 29(1): 201-202

Orrmaesia dorsicosta Kilburn, 1988

Splendrillia Hedley, 1922

Hauturua Powell, 1942

Spirotropis G.O.Sars, 1878

Syntomodrillia Woodring, 1928

Tylotiella Habe, 1958

?Acinodrillia Kilburn, 1988

Ann. Natal Mus. 29(1): 223

Acinodrillia viscum Kilburn, 1988 (s.d. Kilburn, 1988,

Ann. Natal Mus. 29(2): 557)

? Douglassia Bartsch, 1934

? Fenimorea Bartsch, 1934

? Paracuneus Laseron, 1954

Family PEUDOMELATOMIDAE Morrison, 1966

Hormospira Berry, 1958

Pseudomelatoma Dall, 1918

(=Laevitectum Dall, 1919)

Tiariturris Berry, 1958

Family STRICTISPIRINAE McLean, 1971

Cleospira McLean, 1971

Veliger 14(1): 125

Monilispira ochsneri Hertlein & Strong, 1949

Strictispira McLean, 1971

Veliger 14(1): 125

Crassispira ericana Hertlein & Strong, 1951

Family TURRIDAE H. & A.Adams, 1853

Subfamily CLAVATULINAE Gray, 1853

Benthoclionella Kilburn, 1974

Ann. Natal Mus. 22(1): 214

Benthoclionella jenneri Kilburn, 1974

Clavatula Lamarck, 1801

Clionella Gray, 1847

Makiyamaia Kuroda in MacNeil, 1960

Perrona Schumacher, 1817

Pusionella Gray, 1847

Scaevatula Gofas, 1989

Arch. Molluskenk. 120(1/3): 16

Scaevatula pelisserpentis Gofas, 1989

Toxiclionella Powell, 1966

Caliendrula Kilburn, 1985

Ann. Natal Mus. 26(2): 442-443

Latiaxis? elstoni Barnard, 1962

Turricula Schumacher, 1817

? Makiyamaia Kuroda in MacNeil, 1960

Subfamily CRASSISPIRINAE Morrison, 1966

Aoteadrillia Powell, 1942

Austrodrillia Hedley, 1918

Regidrillia Powell, 1942

J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV

Belalora Powell, 1951

Buchema Corea, 1934

Calcatodrillia Kilburn, 1988

Ann. Natal Mus. 29(1): 290-291

Calcatodrillia chamaeleon Kilburn, 1988

Carinodrillia Dall, 1919

Carinapex Dall, 1924

Ceritoturris Dall, 1924

Conorbela Powell, 1951

Conticosta Laseron, 1954

Crassiclava McLean, 1971

Veliger 14(1): 121

Pleurotoma turricula Sowerby, 1834

Crassispira Swainson, 1840

Burchia Bartsch, 1944

Crassispirella Bartsch & Rehder, 1939

Dallspira Bartsch, 1950

Gibbaspira McLean, 1971

Veliger 14(1): 122

Pleurotoma rudis Sowerby, 1834

Glossispira McLean, 1971

Veliger 14(1): 121

Pleurotoma harfordiana Reeve, 1843

Monilispira Bartsch & Rehder, 1939

Striospira Bartsch, 1950

(= Adanaclava Bartsch, 1950)

Doxospira McLean, 1971

Veliger 14(1): 124

Doxospira hertleini Shasky, 1971

Epideira Hedley, 1918

(=Epidirona Iredale, 1931)

Funa Kilburn, 1988

Ann. Natal Mus. 29(1): 267-268

Drillia laterculoides Barnard, 1958

Haedropleura Bucquoy, Dautzenberg & Dollfus, 1883

Hindsiclava Hertlein & Strong, 1955

(= Turrigemma Berry, 1958)

Inodrillia Bartsch, 1943

Inquisitor Hedley, 1918

Kurilohadalia Sysoev & Kantor, 1986

Zoologicheskij Zhurnal 65(10): 1462-1463

Kurilohadalia elongata Sysoev & Kantor, 1986

Lioglyphostoma Woodring, 1928

Maesiella McLean, 1971

Veliger 14(1): 123

Maesiella maesae McLean & Poorman, 1971

Mauidrillia Powell, 1942

Miraclathurella Woodring, 1928

Naskia Sysoev & Ivanov, 1985

Zoologicheskij zhurnal 64(2): 196-197

Naskia axiplicata Sysoev & Ivanov, 1985

Naudedrillia Kilburn, 1988

Ann. Natal Mus. 29(1): 276-278

Naudedrillia nealyoungi Kilburn, 1988

Nquma Kilburn, 1988

FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA 165

Ann. Natal Mus. 29(1): 247 Knefastia Dall, 1919

Pl t iS by, 1886

"Sialic Leucosyrinx Dall, 1889

Plicisyrinx Sysoev & Kantor, 1986 Sibogasyrinx Powell, 1969

Zoologicheskij Zhurnal 65(10): 1465-1466 Indo-Pacific Moll. 2(10): 343

Plicisyrinx decapitata Sysoev & Kantor, 1986 Surcula pyramidalis Schepman, 1913

Psittacodrillia Kilburn, 1988 Marshallena Allan, 1927

Ann. Natal Mus.: 29(1): 253

Pleurotoma bairstowi Sowerby, 1886

Ptychobela Thiele, 1925

Megasurcula Casey, 1904

Nihonia MacNeil, 1960

Turridrupa Hedley, 1922 Paracomitas Powell, 1942

?Paradrillia Makiyama, 1940 Paves E iy bod

(= Vexitomina Powell, 1942) Pyrgospira McLean, 1971

Coronacomitas Shuto, 1983 Veliger 14(1): 119

Mem. Fac. Sci. Kyushu Univ., ser.D (Geol.) 25(1): 1-2 Pleurotoma obeliscus Reeve, 1843

Paradrillia (Coronacomitas) gemmata Shuto, 1983

Rhodopetoma Bartsch, 1944

? Pseudexomilus Powell, 1944

Schepmania Shuto, 1970 Venus 29(2): 37-38 Surcula variabilis Schep-

Subfamily ZONULISPIRINAE McLean, 1971 man, 1913

Compsodrillia Woodring, 1928 ?Micropleurotoma Thiele, 1929

Mammillaedrillia Kuroda & Oyama in Kuroda, Habe & Oyama,

1971 Subfamily TURRINAE H. & A.Adams, 1853 (1840)

The sea shells of Sagami Bay: 208

Compsodrillia (Mammillaedrillia) mammillata OBIE TA RES Med

Kuroda & Oyama in Kuroda, Habe & Oyama, 1971 Decollidrillia Habe & Ito, 1965

Pilsbryspira Bartsch, 1950 Epidirella Iredale, 1931

Nymphispira McLean, 1971

Veliger 14(1): 126 Fusiturris Thiele, 1929

Crassispira nymphia Pilsbry & Lowe, 1932 Gemmula Weinkauff, 1875

Zonulispira Bartsch, 1950 Pinguigemmula MacNeil, 1960

Ptychosyrinx Thiele, 1925

Subfamily COCHLESPIRINAE Powell, 1942

Lophiotoma Casey, 1904

Abyssocomitas Sysoev & Kantor, 1986 (= Lophioturris Powell, 1964)

Zoologicheskij Zhurnal 65(10): 1461-1462 Unedogemmula MacNeil, 1960

Abyssocomitas kurilokamchatica Sysoev & Kantor, 1986 Xenuroturris Iredale, 1929

Aforia Dall, 1889 Lucerapex Iredale, 1936

Abyssaforia Sysoev & Kantor, 1987

Veliger 30(2): 117 Polystira Woodring, 1928

Aforia (Abyssaforia) abyssalis Sysoev & Kantor, 1987

Dallaforia Sysoev & Kantor, 1987

Veliger 30(2): 115-116

Trenosyrinx? crebristriata Dall, 1908 ’ ;

Steiraxis Dall, 1895 Family CONIDAE Fleming, 1822

Turris Roeding, 1798

Annulaturris Powell, 1966

eas Powell. 1942 Subfamily CLATHURELLINAE H. & A.Adams, 1858

Antimelatoma Powell, 1942 ‘bathytomid’ group of genera

Antiplanes Dall, 1902 Bathytoma Harris & Burrows, 1891

(= Rectiplanes Bartsch, 1944) Miciuiance been len Z28

Parabathytoma Shuto, 1961

Apiotoma Cossmann, 1889 Riuguhdrillia Oyama, 1951

Carinoturris Bartsch, 1944 Paraborsonia Pilsbry, 1922

Clavosurcula Schepman, 1913 ‘borsoniid’ group of genera

Cochlespira Conrad, 1865 Asthenotoma Harris & Burrows, 1891

(=Ancistrosyrinx Dall, 1881)

(=Pagodosyrinx Shuto, 1969 HOE beth Nis

Mem. Fac. Sci. Kyushu Univ., ser.D (Geol.) 19(1): 190-191 BOIL Lee

Pleurotoma (Ancistrosyrinx) travancorica granulata Smith,1904) The Veliger 14(1): 126-127

Leucosyrinx erosina Dall, 1908

Comitas Finlay, 1926 Borsonia Bellardi, 1839

Sheskcr, : Boettgeriola Wenz, 1943

Fusiturricula Woodring, 1928

Fusisyrinx Bartsch, 1934 Cordieria Rouault, 1848

166

Cruziturricula Marks, 1951

Ophiodermella Bartsch, 1944

Tropidoturris Kilburn, 1986

Ann. Natal Mus. 27(2): 645-646

Pleurotoma scitecostata Sowerby, 1903

Typhlomangelia G.O.Sars, 1878

Typhlosyrinx Thiele, 1925

? Darbya Bartsch, 1934

‘clathurellid’ group of genera

Clathurella Carpenter, 1857

Comarmondia Monterosato, 1884

Corinnaeturris Bouchet & Waren, 1980

J. Moll. Stud., suppl.8: 77

Pleurotoma leucomata Dall, 1881

Crockerella Hertlein & Strong, 1951

Glyphostoma Gabb, 1872

Glyphostomopsis Bartsch, 1934

Euglyphostoma Woodring, 1970

Prof. pap. U.S. Geol. Survey 306—D: 401

Glyphostoma partefilosa Dall, 1919

Nannodiella Dall, 1919

Strombinoturris Hertlein & Strong, 1951

?Etrema Hedley, 1918

Etremopa Oyama, 1953

Etremopsis Powell, 1942

?Genota H. & A.Adams, 1853

‘mitromorphid’ group of genera

Anarithma Iredale, 1916

Arielia Shasky, 1961

Vexiariella Shuto, 1983

Mem. Fac. Sci. Kyushu Univ., ser.D (Geol.) 25(1): 6

Ariella (Vexiariella) cancellata Shuto, 1983

Diptychophlia Berry, 1964

Lovellona Iredale, 1917

Maorimorpha Powell, 1939

Mitrellatoma Powell, 1942

Mitromorpha Carpenter, 1865

Mitrolumna Bucquoy, Dautzenberg & Dollfus, 1883

(=Apaturris Iredale, 1917)

(=Cymakra Gardner, 1937)

(= Helenella Casey, 1904)

(= Itia Marwick, 1931)

(= Mitrihara Hedley, 1922)

Scrinium Hedley, 1922

Zetekia Dall, 1918

‘tomopleurid’ group of genera

Drilliola Cossmann, 1903

Microdrillia Casey, 1903

(= Acropota Nordsieck, 1977, nom.nov. pro Acrobela Thiele, 1925

non Foerster, 1862

The Turridae of the European seas: 59)

Phenatoma Finlay, 1924

J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV

Pulsarella Laseron, 1954

Suavodrillia Dall, 1918

Tomopleura Casey, 1904

Maoritomella Powell, 1942

? Austroturris Laseron, 1954

? Filodrillia Hedley, 1922

? Heteroturris Powell, 1967

Indo-Pacific Moll. 1(7): 411

Heteroturris sola Powell, 1967

Subfamily? CONORBIINAE De Gregorio, 1890

Conorbis Swainson, 1840

Benthofascis Iredale, 1936

Subfamily OENOPOTINAE Bogdanov, 1987

Curtitoma Bartsch, 1941

(= Widalli Bogdanov, 1986

Zoologicheskij Zhurnal 65(1): 45

Pleurotoma trevelliana Turton, 1834)

Granotoma Bartsch, 1941

Obesotoma Bartsch, 1941

Oenopota Morch, 1852

Nodotoma Bartsch, 1941

Oenopotella Sysoev, 1988

Zoologicheskij zhurnal 67(8): 1119-1120

Oenopotella ultraabyssalis Sysoev, 1988

Propebela Iredale, 1918

Canetoma Bartsch, 1941

(=Funitoma Bartsch, 1941)

? Lorabela Powell, 1951

Subfamily MANGELIINAE Fischer, 1883

Acmaturris Woodring, 1928

Agathotoma Cossmann, 1899

Anacithara Hedley, 1922

Antiguraleus Powell, 1942

Apispiralia Laseron, 1954

Apitua Laseron, 1954

Bactrocythara Woodring, 1928

Bela Gray, 1847

Belaturricula Powell, 1951

Bellacythara McLean, 1971

The Veliger 14(1): 128

Clavatula bella Hinds, 1843

Benthomangelia Thiele, 1925

Brachycythara Woodring, 1928

Cacodaphnella Pilsbry & Lowe, 1932

Citharomangelia Kilburn, 1992

Annals Natal Mus. 33(2): 508-9

Mangilia africana Sowerby, 1903

Clathromangelia Monterosato, 1884

Cryoturris Woodring, 1928

FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA 167

Cytharella Monterosato, 1875 Thelecythara Woodring, 1928

Cyrtocythara Nordsieck, 1977

The Turridae of European seas: 34 Turrella Laseron, 1954

Pleurotoma albida Deshayes, 1834 Vitjazinella Sysoev, 1988

Rugocythara Nordsieck, 1977 Zoologicheskij zhurnal 67(8): 1122

The Turridae of European seas: 35 Vitjazinella multicostata Sysoev, 1988

Pleurotoma rugulosa Philippi, 1844

Eucithara Fischer, 1883

Euclathurella Woodring, 1928

Vitricythara Fargo, 1953

?Anticlinura Thiele, 1934

g ?Conopleura Hinds, 1844

Fehria van Aartsen, 1988

La Conchiglia 20(232-233): 232 ?Hemicythara Kuroda & Oyama in Kuroda, Habe & Oyama, 1971

Ginnania taprurensis Pallary, 1904 The sea shells of Sagami Bay: 229

2 ae: ; Pleurotoma octangulata Dunker, 1860

Gingicithara Kilburn, 1992

Annals Natal Mus. 33(2): 495-6 ?Paraclathurella Boettger, 1895

Mangelia lyrica Reeve, 1846 ;

Subfamily DAPHNELLINAE Deshayes, 1863

Glyphoturris Woodring, 1928

Abyssobela Kantor & Sysoev, 1986

Glyptaesopus Pilsbry & Olsson, 1941 Zoologicheskij Zhurnal 65(4): 492

Gites Hedley, 1918 Abyssobela atoxica Kantor & Sysoev, 1986

Euguraleus Cotton, 1947 Antimitra Iredale, 1917

Mitraguraleus Laseron, 1954

Asperdaphne Hedley, 1922

Heterocithara Hedley, 1922 Aspertilla Powell, 1944

Ithycythara Woodring, 1928 Austrodaphnella Laseron, 1954

Kurtzia Bartsch, 1944 Bathybela Kobelt, 1905

: (= Bathypota Nordsieck, 1968

Sy lbebad ae ae The Turridae of European seas: 28

Granoturris Fargo, 1953

o o Si “ a 7,

Rubellatoma Bartsch & Rehder, 1939 Pleurotoma tenellula [sic] Locard, 1897)

Kurtzina Bartsch, 1944 Buccinaria Kittl, 1887

Leiocithara Hedley, 1922 Cryptodaphne Powell, 1942

Ac todaphne Shuto, 1971

Lienardia Jousseaume, 1884 Bhi 30(1): 10 or

Acrista Hedley, 1922

Hemilienardia Boettger, 1895

Thetidos Hedley, 1899 Cenodagreutes Smith, 1967

, The Veliger 10(1): 1

ieuee Gonuer, 1924 Cenodagreutes aethus Smith, 1967

Pleurotomella biconica Schepman, 1913

Macteola Hedley, 1918 Daphnella Hinds, 1844

Mangelia Risso, 1826 Diaugasma Melvill, 1917

Hemidaphne Hedley, 1918

Mangiliella Bucquoy, Dautzenberg & Dollfus, 1883

Lyromangelia Monterosato, 1917

Marita Hedley, 1922 Eucyclotoma Boettger, 1895

Exomilus Hedley, 1918

Eubela Dall, 1889

Neoguraleus Powell, 1939

Notocytharella Hertlein & Strong, 1955 Famelica Bouchet & Waren, 1980

J. Moll. Stud., suppl.8: 88

Papillocithara Kilburn, 1993 Pleurotomella catharinae Verrill & Smith, 1884

Annals Natal Mus. 33(2): 516-7 ;

Papillocithara hebes Kilburn, 1992 Fusidaphne Laseron, 1954

Gymnobela Verrill, 1884

(= Majox Nordsieck, 1968

Platycythara Woodring, 1928 Die europaischen Meeres-Gehause Schnecken: 182

Pleurotomella bairdi Verrill & Smith, 1884)

(= Watsonaria Nordsieck, 1968 (nomen nudum)

Die europaischen Meeres-Gehause Schnecken: 182

Clathurella watsoni Dautzenberg, 1889)

Pyrgocythara Woodring, 1928 Theta Clarke, 1959

Paramontana Laseron, 1954

Pseudoetrema Oyama, 1953

Pseudoraphitoma Boettger, 1895

Saccharoturris Woodring, 1928 Isodaphne Laseron, 1954

Stellatoma Bartsch & Rehder, 1939 Kermia Oliver, 1915

Tenaturris Woodring, 1928 Kuroshiodaphne Shuto, 1965

168

Lusitanops Nordsieck, 1968

Die europaischen Meeres-Gehause schnecken: 181

Pleurotomella lusitanica Sykes, 1906

(= Pseudazorita Nordsieck, 1977 (published as nomen nudum)

The Turridae of the European seas: 31 (published as

a subgenus of Thesbia)

Pleurotoma blanchardi Dautzenberg & Fischer, 1896,

s.d. Bouchet, Waren, 1980, 1980, J. Moll. Stud.,

suppl. 8: 83)

Magnella Dittmer, 1960

Microdaphne McLean, 1971

The Veliger 14(1): 129-130

Philbertia trichodes Dall, 1910

Microgenia Laseron, 1954

Neopleurotomoides Shuto, 1971

Venus 30(1): 5-6

Clathurella rufoapicata Schepman, 1913

Nepotilla Hedley, 1918

Ootomella Bartsch, 1933

Pagodidaphne Shuto, 1983

Mem. Fac. Sci. Kyushu Univ., ser.D (Geol.) 25(1): 21

Pagodidaphne colmani Shuto, 1983

Philbertia Monterosato, 1884

(= Lineotoma Nordsieck, 1977, nom.nov. pro Cirillia

Monterosato, 1884 non Rondani, 1856

The Turridae of the European seas: 18)

Glyphostomoides Shuto, 1983

Mem. Fac. Sci. Kyushu Univ., ser.D (Geol.) 25(1): 16-17

Philbertia (Glyphostomoides) queenslandica Shuto, 1983

Phymorhynchus Dall, 1908

Pleurotomella Verrill, 1873

(= Azorilla Nordsieck, 1968

Die europaischen Meeres-Gehause Schnecken: 184

Pleurotoma megalembryon Dautzenberg & Fischer, 1896)

(= Azorita Nordsieck, 1968

Die europaischen Meeres-Gehause Schnecken: 184-185

Pleurotoma bureaui Dautzenberg & Fischer, 1897)

Anomalotomella Powell, 1966

Pontiothauma Smith, 1895

Pseudodaphnella Boettger, 1895

Raphitoma Bellardi, 1848

Cyrtoides Nordsieck, 1968

Die europaischen Meeres-Gehause schnecken: 176

Raphitoma rudis Scacchi, 1836 (= R. (C.) neapolitana Nords-

eck, 1977, nom.nov. pro R. rudis Scacchi, 1836 non Broderip)

Rimosodaphnella Cossmann, 1915

Spergo Dall, 1895

Speoides Kuroda & Habe, 1961

Stilla Finlay, 1926

Tasmadaphne Laseron, 1954

Teretia Norman, 1888

Teretiopsis Kantor & Sysoev, 1989

J.Moll.Stud. 55: 538

Teretiopsis levicarinatus Kantor & Sysoev, 1989

Thatcheria Angas, 1877

Tritonoturris Dall, 1924

J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV

Truncadaphne McLean, 1971

The Veliger 14(1): 129

‘Philbertia’ stonei Hertlein & Strong, 1939

Tuskaroria Sysoev, 1988

Zoologicheskij Zhurnal 67(7): 970-972

Tuskaroria ultraabyssalis Sysoev, 1988

Veprecula Melvill, 1917

Vepridaphne Shuto, 1983

Mem. Fac. Sci. Kyushu Univ., ser.D (Geol.) 25(1): 17

Daphnella cestrum Hedley, 1922

Xanthodaphne Powell, 1942

Zenepos Finlay, 1928

?Aliceia Dautzenberg & Fischer, 1897

? Benthodaphne Oyama, 1962

?Otitoma Jousseaume, 1898

? Thesbia Jeffreys, 1867

Subfamily? TARANINAE Casey, 1904

Taranis Jeffreys, 1870

CONIDAE INCERTAE SEDIS

Austrocarina Laseron, 1954

Austropusilla Laseron, 1954

Metaclathurella Shuto, 1983

Mem. Fac. Sci. Kyushu Univ., ser.D (Geol.) 25(1): 15

Austropusilla (Metaclathurella) crockerensis Shuto, 1983

Paraspirotropis Sysoev & Kantor, 1984

Zoologicheskij Zhurnal 63(7): 1096-1097

Pleurotomella simplicissima Dall, 1907

Teleochilus Harris, 1897

Toxicochlespira Sysoev & Kantor, 1990

Apex 5(1-2): 2-3

Toxicochlespira pagoda Sysoev & Kantor, 1990

Typhlodaphne Powell, 1951

CONOIDEA INCERTAE SEDIS

Cretaspira Kuroda & Oyama in Kuroda, Habe & Oyama, 1971

The sea shells of Sagami Bay: 219

Cretaspira cretacea Kuroda & Oyama in Kuroda, Habe &

Oyama, 1971

Graciliclava Shuto, 1983

Mem. Fac. Sci. Kyushu Univ., ser.D (Geol.) 25(1): 11

Graciliclava mackayensis Shuto, 1983

Inkinga Kilburn, 1988

Ann. Natal Mus. 29(1): 230

Pleurotoma (Clionella) platystoma Smith, 1877

Kurodadrillia Azuma, 1975

Venus 33(4): 159

Kurodadrillia habui Azuma, 1975

Lioglyphostomella Shuto, 1970

Venus 28(4): 165-166

Drillia timorensis Schepman, 1913

Meggittia Ray, 1977

Contribution to the knowledge of the molluscan fauna of

Maungmagan, Lower Burma...: 66-67

Meggittia maungmagana Ray, 1977

Thatcheriasyrinx Powell, 1969

FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA

Indo-Pacific Moll. 2(10): 405

Ancistrosyrinx orientis Melvill, 1904 (by monotypy)

Viridoturris Powell, 1964 (formerly Turrinae)

Taxa transferred to other families

Bathyclionella Kobelt, 1905 — Buccinidae (as synonym of

Belomitra; Bouchet, Waren, 1980, J. Moll.Stud., suppl.8)

Belomitra Fischer, 1882 — Buccinidae

Steironepion Pilsbry & Lowe, 1932 — Columbellidae

Surculina Dall, 1908 — Turbinellidae (Rehder, 1967, Pacific

Sci. 21(2): 182-187)

Turrijaumelia Sarasua, 1975

Poeyana 140: 12-13

Turrijaumelia jaumei Sarasua, 1975

Transferred to Columbellidae as a synonym of Steironepion

Pilsbry & Lowe, 1932 (Finlay, 1984, Nautilus 99(2-3): 73-75)

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CONTENTS

97 The status of the Persian Gulf sea snake Hydrophis lapemoides (Gray, 1849) (Serpentes,

Hydrophiidae)

A. Redsted Rasmussen

107 Taxonomic revision of some Recent agglutinated foraminifera from the Malay Archipelago,

in the Millett Collection, The Natural History Museum, London

P. Brénnimann and J.E. Whittaker

125 Foregut anatomy, feeding mechanisms, relationships and classification of the Conoidea

(= Toxoglossa) (Gastropoda)

J.D. Taylor, Y.l. Kantor and A.V. Sysoev

_ Bulletin of The Natural History Museum

ZOOLOGY SERIES

Val. 58. No. 2, November 1993

~~ THE NATURAL

| HISTORY MUSEUM

4 JUL 1994

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VOLUME 60 NUMBER 1 23 JUNE 1994

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Printed in Great Britain at The Alden Press, Oxford

Bull. nat. Hist. Mus. Lond. (Zool.) 60(1): 1-37

ic WAT JRAL

A new subfamily and genus in Achatinida HISTORY MUSEL

(Pulmonata: Sigmurethra) 15 JUL -:

ALBERT R. MEAD

Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, AZ 85721, USA

CONTENTS

SSULOPISIS Meena nctetdcsnaradetc ese ane nea Ose tes CORR MAAS ebb nieon dfdameeaee ate esmenasemsrncuadtaah cbt vet Muth metnat se dsetadte ds 1

LiniOGhUCINOTE) eccee cedar deena dence chey<teccertcer Ae Cae ap eCacOn CAeRBRREE Ee. CEE DC Cer aA aRA Mer ee cea ann Cane aero ence nee PEAT eR Er sr 1

MS RINO OS ete eta: 394 oa ite ae esta heres Sa eet RAGA SER E ciesieiinleincincin Siar esisinciiniainienaletate nina vdlslehladaseaa sadddeb actos Reeseeeeee 3

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LG DE INO BYES oto oer HOCnas elon Tee OF ae Pooch I0 Ce nee ree PRRBEREET °C Oc Sa0 050 ap Acc AACE ac REE EEE MRC CL MERE EE CEERt CRC Cem en tener tre 3

IEVACOLS ID LAMMUICS tr ga. teeee anes ho teat debt =n Aye aco ect. 5 on » wa asa peMb eet cleat ws powsiemigae tujepeniyanedeadeach boheme pe aue ews 3

Gallistoplepinide:— Mew SUDLAMMLY, s.53.15- base oie Sue caiaeins's ov ow eee aoe Maatee ve aidee Mec cash cabin gag goane qeiet ee evgagees dB oeetntee 3

[SCENT TO WL SCae LOVEE Ot PROCES ES SSHRE Se CRCMECOT: 4 SOR RCRE TCEE Sat oe can ERanE'- Bac clace casas 5: PoUicGn ek EOE ena DCC RSE cr BeGer rr ae mom EER Ceci E 4

GTS CAIESIOP CP Cte ccresgh acta Sans sania sarees aes pra td eeosic 52s on eaten c ope oe Sea ae nanos as dra aie deo neeina ste siaereles ab 5

INEN ALOIS E CICS it acranene tenes Seas el ‘apppbasippeadiaeses cidapobs'sduise osha eae Antal tane dies. Aaa ga satan scence cca ost adaeduseitasadoaee esate 5)

MEETING PNOGHLG) retina tes a soacete seine ats as cla ap sidleles ce se ects a nie 2.03 peepee cima Seed « oases ea¥laalia nid de Sab n SuayGee Hote Geddeesaemsaees 12

ISG VerOISDECICS mana tere tne ttn cs cece gteea noe ecatices fe ceteatis vas «cheitan a tah guy Septic badass a dosseans dua dawswcacs « ni teseeesaenes 13

PCNA MGIMAC 9 hac M Are ese aaene setae nc ene waisapinci ees Sree ae or cee de tes ele wo celona dale aeieiaaais dabipiaide mae mmisals Sasiiahs « daeiusla'c niet suaystccieset ss 18

TEMUISE/S COLLACHIIIE — MIGWAPCMLUS sr seis arn. te ieee aks dee tee ni'sinalsidiafeniaiweniaita arieids in ello ohn aiac da ents « se penwatelaw'acaijalsa Sesiecajeals aie 18

RS VALORSPIC ELC Stren Rteta a dela een the ome seis ete eetnutetrelaa mated ars wok npinicnia ap Saeioe dita iasleia dee tiaale wilaivast ine nincidsle te se sislnee.cejeriaictaciaee 22

ACN A Cer M| AUUSMMEE tease Re Tee dees Rcccter argent Set Ven vn es ctpemeisehine vamciiaseadedgel + eeinnissmasseasesasieeis Saas emesnelag teseanenes 32

PXOMMOWICCSEMICUES Pe. cesses setanees dee hteeises« Wee savetecb nse ccesstiodee scart ar aadadytaecsinnse semis ianedescenaeats senangusk teeaeane 35

/NGTOTUNIITS »-eaadenndbedeosoecaebantnannbpadeecesbarhe babcantinen cmt pBREREe Ct ockiae ebhc: can AOcn Ca ance ROAR cc BnCrberaur tid sdemcretdeid Oe Siiap 35

STENTS CES. icoggeacet cagoar sobre ddgoar. Uornbacr Cac HOHO irs (oc cce Cee PERRET DACEC -AE SERS n EEE aE Dict EES nse ene PAP em eee 36

Synopsis. In the Achatinidae, characteristics of the internal anatomy, particularly those in the reproductive tract,

are continuing to show greater dependability in determining phylogenetic affinities than those in the shell, radula or

jaw. In 1934, J.C. Bequaert reached the conclusion in the course of his revising the African land snail family

Achatinidae, ‘that several of the East African Achatinae are not separable from the West Africa Callistoplepae, at

any rate as far as shell characters go’. The present 5-year study of over 500 shell specimens and 50 soft anatomies,

involving 11 genus-group and 57 species-group nomina, demonstrates that the 4 anatomically distinct West African

species, to which Bequaert referred, are the most primitive in the family. Accordingly, they have been placed in a

separate subfamily. Anatomically the 5 contrastingly different East African species were found to be a distinct

genus, plesiomorphic to subgenus Achatina (sensu Bequaert, 1950). Their phylogenetic relationships show a strong

correlation with the distributional evidence in 60 recorded localities, which delineates (1) a short north central axis,

(2) a central to eastern Africa axis, and (3) a strong north-south axis extending from eastern Africa into southern

Africa. This evidence supports the emerging pattern of terrestrial gastropod distribution in Africa. The present

project forms the foundation for a revision of the family, currently in progress.

Taxonomy in the Achatinidae is based both on the shell

and on the soft anatomy, particularly that of the basal

reproductive tract. The shell is a permanent record that

INTRODUCTION

Fifty-seven species-group nomina associated with the acha-

tinid genus Callistoplepa have provided the basis for the

present 5-year project. The examination of over 500 shell

specimens and 50 soft anatomies has revealed the fact that

four West African anatomically distinct but related Lower

Guinea species, in two genera, constitute a separate, primi-

tive subfamily of the Achatinidae. Anatomical and concho-

logical studies place the five other valid species in a new

genus that is distributed in Central, Eastern and Southern

Africa.

reflects ontogenetically the influence of the environment,

whereas the soft anatomy reflects phylogenetically the influ-

ence of evolution. The latter, in the short term, is relatively

free of substantive changes, being limited to temporary

variations that reflect nutritional, developmental or repro-

ductive influence. For example, both immature and seriously

malnourished mature specimens have misleadingly attenu-

ated, thin reproductive tracts with greatly limited muscula-

ture; but the malnourished specimen can be readily

distinguished because it has a much larger reproductive tract

with the rich coloration of the mature specimen. Under

suitable conditions, both types of individuals are able to

assume the basic genital proportions that are typical of the

species. In the long term, the shell reflects and responds to

the pressures of selection in a changing or in a new environ-

ment. Wholly different molluscan stocks moving into distinct

but similar environments may evolve convergently into forms

that are so similar that conchologically they cannot reason-

ably be distinguished or identified without first knowing the

locality, e.g. Achatina (Lissachatina) craveni Smith, 1881 and

A. (Achatina) tavaresiana Morelet, 1866 (Mead, 1992).

Although in the long term, the soft anatomy undergoes

changes, it revealingly does so within a more restricted, basic,

generalized patten obviously characteristic of the larger

group of which it is a related part. At this stage of investiga-

tions, these larger groups seem to be taking shape within

genera and subgenera (sensu Bequaert, 1950).

All these factors have so entered into the present project

that nine species, once thought to be congeners on the basis

of the great similarity in their familial unique shell characters,

are now separated into two subfamilies. The shells are

remarkably similar, but the soft anatomies convincingly tell a

different story. In essence, in the Achatinidae the soft

anatomy reveals dependable, differentiating taxonomic crite-

ria at the species level and above; the shell usually reveals

supporting taxonomic criteria at the species level and often

convincing criteria at the subspecies level. A more clear

concept of subspecies is beginning to emerge in this family:

Essentially consistent, usually minor shell differences in the

members of an allopatric population that have features in

their soft anatomy indistinguishable from those of the nomi-

nate subspecies. This is precisely why the erstwhile enigmatic

Leptocala petitia is shown in the text to be a distinct species

rather than a subspecies or a synonym of the conchologically

very slightly different L. mollicella.

The shells of the nine species in this project have been

redescribed in the light of larger series of specimens and more

detailed examinations of shell characters than in previous

studies. In many full grown specimens, weathering and

environmental abrasion have obscured or removed valuable

shell characters, particularly in the upper whorls. The empha-

sis On a comparative examination of juvenile specimens in the

present study, therefore, has been especially informative. In

fact, it strongly supports the value of the collecting juvenile

specimens, along with the larger specimens, whenever they

are available. For the same reasons, determining the number

of whorls is always an imprecise measurement and thus is

recorded here only to the nearest one-quarter whorl. The

length of the last whorl (= body whorl) includes the entire

whorl and is measured from the base of the aperture to the

point on the suture immediately above where the outer lip

attaches at the periphery. All shell measurements are carried

to the nearest 0.1 mm. For many decades the basic horizontal

and vertical sculpture of the achatinid shell has been referred

to as ‘decussate’; this has been changed in the present work to

the more accurate term ‘cancellate’. From the shell dimen-

sions, two important relationships are drawn: 1) between the

greatest shell width and the shell length, and 2) between the

length of the last whorl and the shell length. These relation-

ships are expressed in percentages rather than in ratios. This

emphasizes the measurement being compared rather than the

measurement to which it is being compared; that is, in

making comparisons, it is easier to comprehend that the last

whorl is 87% of the shell length, than the fact that the length

A.R. MEAD

is 1.15 times the length of the last whorl. Aperture width does

not include the callus. The descriptive shell terminology is

largely based on Cox (1960).

Diligent searching in 49 museums and personal collections,

fortunately brought to light an unexpected fair number of

alcohol preserved specimens. The museums with the largest

and most varied collections of alcohol preserved achatinid

specimens are in Tervuren, Stockholm, Berlin, Bruxelles,

Paris, Frankfurt and London. But very valuable specimens

have been found in museums where there is relatively limited

material. It was through the convictions of Edmond Dart-

evelle of the Muséum National de l’Afrique Centrale (Ter-

vuren) that the wet collection of the museum is unparalleled

in quantity and diversity. Because the available material used

in the present project is limited, determined efforts were

made, after establishing the basic genital pattern, to conserve

the remaining specimens for future investigators.

The relationships of the basal genital system show to

greater comparative advantage in ventral view. For this

reason, the line drawings are similarly oriented in this per-

spective, with the male conduit to the left and the female

conduit to the right, unless otherwise indicated. The indi-

vidual structures of the conduits are spread apart to show

their configuration and minimize obstruction. The drawings

are idealized, where possible through multiple specimens, to

eliminate irrelevant and misleading features produced in

preservation. The origin of the penial retractor is on the male

conduit at a point that marks the division between penis and

vas deferens. As the retractor anlage reaches apically during

development, it usually inserts somewhere on the columellar

muscle system, but may insert on the body wall, diaphragm,

transverse myoseptum or other sites in the haemocoele

(Mead, 1950). The configuration of the developing viscera

may predispose the manner of insertion. The not infrequent

bifurcate (Fig. 20) and multifurcate penial retractor insertions

support the ontogenetic rather than the functional interpreta-

tion of these terms. Within a species, the site of insertion may

be consistent or variable. There is a fairly strong tendency in

this family for the basal penial retractor to proliferate muscle

and apparently connective tissue that variously produce adhe-

sions in the several parts of the male conduit. This in turn

changes the relationship of these parts, alters the extrover-

sion process, and the configuration of the resultant intromit-

tent organ. This organ is normally composed seriatim, apex

to base, of the penis, pilaster (when present), penis sheath,

penial atrium and genital atrium externally, and the basal vas

deferens, penial retractor, apical vas deferens and ejaculatory

duct internally.

Directions of left and right refer to those of the snail.

Apical, in reference to the genital system, means toward the

ovotestis, basal toward the genital atrium. The anatomical

terminology is essentially that of Mead (1950) (see Abbrevia-

tions — Anatomical below).

The discussion of each species is in the following format:

shell, soft anatomy (where available), type material, type

locality, distribution and, when applicable, remarks. The

sources and localities of the alcohol-preserved specimens are

reported in the text. A table for each species includes the

sources of all shell specimens examined, taxonomically

important specimens, illustrated specimens, a size range,

their localities, shell dimensions and shell proportions. Illus-

trations in the literature are cited in the synonymies; the

nature of the illustration, where it is other than of the shell, is

shown in parentheses. Most localities were found in the

NEW SUBFAMILY AND GENUS ACHATINIDAE

volumes of the U.S. Board of Geographic Names and their

locality figures were preferentially used. A list of acronyms of

institutions and private collections follows the text. Symbols

and abbreviations used in the Tables: Holo = Holotype, L =

Length, Lect = Lectotype, LW = Last Whorl, Para =

Paratype, PLec = Paralectotype, W = greatest Width, * =

dissected, * = see photograph.

Continued research along the lines of Mead (1950, 1978,

1992) and in the present work promises to establish a sound

taxonomic and phylogenetic base for the Achatinidae.

METHODS

Examining a great number of variously preserved whole

specimens in the present project has once again emphasized

the importance of using proper preservation procedures.

Ideally, specimens selected for preservation of the soft parts

should be put in previously boiled water that has reached

ambient temperature. They should drown normally 8-12

hours. Overdrowning will cause the basal genital structures to

evert, irreparably distorting the taxonomically valuable fea-

tures. Underdrowning permits the specimen to withdraw

excessively into the shell. Crowding the specimens or using

too small a volume of fluid promotes maceration. The

adequately drowned specimen usually contracts slightly when

placed in the initial 40% alcohol. In a few hours, depending

upon the size of the specimen, it should next be placed in a

60% solution, followed by at least one change to 70%

alcohol.

Formalin is a powerful, penetrating, irritating fixative. If it

is used at a very low percentage for a short period before the

specimen is washed thoroughly and transferred to 70%

alcohol, it can be quite effective. But in general, its use

should be avoided because formalin alters the colour of the

shell, makes the shell brittle and chalky, causes the perios-

tracum to crack and peel off upon drying, promotes adhe-

sions between the shell and soft parts, severely hardens the

muscular body wall, and precipitates great quantities of

albumin and recrystallized calcareous islands in the tissues.

As a result, extrication rarely can be accomplished without

damage to both shell and soft parts. Further, even with

prolonged soaking in a 0.5% trisodium phosphate solution,

the muscular body wall remains so hard and tough that, with

extreme difficulty it has to be snipped out, piece by small

piece, to get to the soft parts, which often are so intensely

fixed that they are brittle. Once the soft parts are removed,

they usually can be relaxed for limited manipulation only by

further soaking in trisodium phosphate and one or more

prolonged water baths. Even then, a distressing degree of

brittleness remains. The practice of ‘neutralizing’ formalin for

the preservation of vertebrate specimens by first dropping

live snail specimens in the solution is deplorable.

ABBREVIATIONS — ANATOMICAL

AVD apical vas deferens

BVD basal vas deferens

E egg

EM eversion muscle bands

FO free oviduct

GA genital atrium

OTD ovotestis duct

P penis

PA penial atrium

PIL pilaster

PR penial retractor

BS penis sheath

RCR right columellar retractor

ROR right ommatophore retractor

S spermatheca

SD spermathecal duct

SO spermoviduct

SSV secondary seminal vesicle

ar talon

Vv vagina

VA vaginal atrium

VR vaginal retentor

Achatinidae

Basal genital conduits simple, without accessory organs. A

conspicuous sheath partially, or usually, completely envelops

the penis. Spermatophores not formed. The right branch of

the columella muscle system regularly remains to the left of

the genitalia. Kidney long, two to three times the length of

the pericardium; sigmurethrous. Pulmonary vein without

major branches. Holopod. Rachidian tooth very slender and

apparently nonfunctional, rarely wide. Jaw simple; smooth or

usually striated. Shell ovate, elongate-ovate or conic-oblong,

rarely columnar; anomphalous or umbilicate; columella trun-

cate or continuous with outer lip, some forms are intermedi-

ate. Endemic in continental Africa and its adjacent small

coastal islands; four known introduced species elsewhere in

the world.

Key to Subfamilies

Vas deferens does not penetrate the penis sheath, but leaves

apically with the penial retractor through the sheath aperture.

The penial retractor inserts on the right columellar retractor;

it is extremely short, entirely or almost entirely covered by

the penis sheath; penis contains a large, conspicuous pilaster.

Rachidian tooth about as wide as the laterals ..... CALLIS-

TOPLEPINAE

Vas deferens penetrates the penis sheath. Even within a

single population, the penial retractor may variously insert on

muscle bands, body wall, diaphragm or fascia; it is usually

long to very long and entirely or almost entirely free of the

penis sheath; penis contains an ill-defined pilaster, no pilas-

ter, or a verge. Rachidian tooth usually much narrower than

the late nal S gaa ctater i-ssencee seeks toe seeecicteen ACHATININAE

CALLISTOPLEPINAE -— new subfamily

This subfamily contains the most primitive achatinids yet

known. Phylogenetically, it is at the base of the Achatinidae,

near the Subulinidae. Like that in the Subulinidae, the vas

deferens does not penetrate the penial sheath. The more

ovate, patterned shell, however, with its larger aperture and

limited number of whorls places this taxon in the Achatinidae

rather than the Subulinidae. Supportive of this are the long

kidney and the pattern of lung venation. The radula is

uncharacteristic of either family, but this is of lesser impor-

tance phylogenetically because this structure is well known to

be responsive to changing feeding habits within closely

related species. The wide, functional rachidian tooth immedi-

ately distinguishes the Callistoplepinae from most Achatini-

nae (Fig. 58-63). D’Ailly (1896:69) was the first to examine

and illustrate the radulae of both species of Callistoplepa (see

also Pilsbry, 1904:ix,xv; Thiele, 1929:560 and Ortiz & Ortiz,

1959:46). In the present study, the radulae of C. barriana

(Sowerby, 1890), C. shuttleworthi (Pfeiffer, 1856) and Lepto-

cala mollicella (Morelet, 1860) were found to follow the same

basic form and pattern. Similarly all the jaws are essentially

identical — simple, nearly smooth, fulvous, chitinoid collari-

form band that is somewhat wider in the middle and tapering

at the sides. There is no suggestion of even generic difference

in these structures. Thus, with only two alcohol specimens of

L. petitia (Jousseaume, 1884) extant, a decision was made to

leave their odontophores intact.

All four species in the Callistoplepinae are limited geo-

graphically (ca 5° N—S° S) to the tropical Lower Guinea

region of West Africa, which at this point appears to be the

cradle of the Achatinidae (Mead, 1992). The high natural

luster of their translucent, elongate-ovate shells probably

reflects selective advantage in the protractedly wet rainforest.

Prior to the present study, the soft anatomy had been

examined in one or more species in the eleven achatinine

genera except the Guinean genus Columna (none so far

available) and all except four of the thirteen achatinine

subgenera (sensu Bequaert, 1950). All that have been exam-

ined clearly are anatomically more advanced than the two

callistoplepine genera.

The calcareous, thick-shelled eggs are comparatively large

for the family and are on a par with those of Tholachatina.

D’Ailly (1896:68) felt they were somewhat small compared to

the size of the snail shell. Thiele (1929:560) echoed this point;

but d’Ailly apparently was comparing them with the rela-

tively huge eggs of some Archachatina and the subulinids. A

reticulate-microtuberculate texture covers the slender,

attenuated body. Basally, the mantle is generously covered

with variable size, fusing black to gray spots; these show

through the thin shell. Apically, the spots are smaller, more

regular and concentrated on the shoulder of the whorls. The

genital orifice appears to be unusually far posterior; Ortiz &

Ortiz (1959) made this observation in C. shuttleworthi.

Typical of the known achatinids, the anterior aorta in the

Callistoplepinae is found on the dorsal surface of the dia-

phragm where it abruptly penetrates the diaphragm to pass

vertically along the anterior edge of the sagittal myoseptum.

Anteriorly, this latter separates the right and left columellar

retractors and incompletely places the male and female basal

conduits in left and right chambers, respectively. In all four

species, the triangular kidney is long, 2-3 times the length of

the pericardium, and sigmurethrous. The ascending limb of

the urethra is closed for its entire length. Venation of the lung

is a dense, broad network on the pericardial side of the

slender principal vein, whereas on the right of the principal

vein there is a relatively narrow band of parallel, limitedly

branching veins between it and the ascending limb of the

ureter. The second largest vein, about half the caliber of the

principal vein, starts as a network in the far left posterior

corner of the lung, anastomoses, and joins the principal vein

at a right angle S—7 mm anterior to the kidney.

A.R. MEAD

Together, the four included species in this subfamily mani-

fest seriatim, from simple to complex, an impressive transi-

tion series in the basal male conduit from what surely is a

pilaster to that which appears to be a verge or penis papilla.

Callistoplepa barriana is the most primitive with an elongate,

elevated pilaster on the ventral penial wall. In C. shuttlewor-

thi the apical penis has permanently partially evaginated,

pulling dorsally the basal-most part of the vas deferens into

the pilaster and fixing it in place with tissue derived from the

adjacent penial retractor. This progression is taken one step

further in Leptocala petitia, wherein the permanently evagi-

nated apical penis becomes so greatly enlarged, thick-walled

and dorsoventrally distorted that the resultant pilaster essen-

tially fills the thin-walled, saccular basal penis. Finally in L.

mollicella, the pilaster assumes an apical position wherein it is

axially pendulous within the thick-walled basal penis. At first

glance, it appears to be a penis papilla, but the asymmetry

within betrays the fact that it is in actuality a greatly modified

pilaster. In all four species, the penial retractor is extremely

short and inserts on the right columellar retractor. In C.

shuttleworthi it inserts forward near the other branches; in the

other species it inserts far to the rear. The penis sheath is so

thin that it is difficult to trace; but it naturally enshrouds the

entire penis, allowing the penial retractor and the vas defer-

ens to pass out apically through the aperture of the sheath. A

barely discernible transparent tissue layer attached directly

on the surface of the penis is formed by the penial retractor.

This may be the forerunner of the condition found in some

Angolan achatinids, e.g. Achatina welwitschi Morelet, 1866,

in which the penis is buried in dense muscle tissue extending

from the penial retractor. Both the inner surface of the penis

sheath and the adjacent but continuous outer surface of the

penis are smooth, shiny and free from each other. This

condition facilitates seriatim extroversion.

The spermatheca is consistently attached to the spermovi-

duct well above the junction of the apical vas deferens and

the free oviduct. In the adult forms, there is no distinct

vaginal retentor between the vagina and the body wall. In

juvenile specimens of C. shuttleworthi, however, the anlage is

present, which suggests that in this subfamily, its full develop-

ment may be obviated by the highly developed, muscular

basal female conduit.

Type genus: Callistoplepa Ancey, 1888.

Key to Genera

Sculpture of body whorl coarse, with slender, tightly and

evenly placed prosocline costate ridges; shell aperture large,

usually 52% of shell length; last whorl long, usually > 73% of

shell length. Posterior foot with dorsolateral serrate ridges.

No colored band on neck. Equatorial Guinea to Nigeria-

ETERS EMCEE nao Pua cock deren Oscanostanuecoce ts Callistoplepa

Sculpture of body whorl with extremely finely engraved

microscopic rhomboids or vertical vermiculate granulae; shell

aperture modest, usually > 52% of shell length; last whorl

shorter, usually > 73% of shell length. Foot without dorsolat-

eral ridges. Dark gray band on neck between ommatophores

and mantle. Cameroon to western Zaire ........... Leptocala

NEW SUBFAMILY AND GENUS ACHATINIDAE

Callistoplepa

Ancey, 1888:69 (footnote 2 for ‘Achatina shuttleworthiana’

[sic = A. shuttleworthi Pfeiffer, 1856]); Pilsbry, 1905:viii,

ix (fig. 2), xv (radula); Germain, 1909:90; Pilsbry,

1919:54, 60, 80, fig. 25 (map); Bequaert & Clench,

1934c:114; Ortiz & Ortiz, 1959:44; Zilch, 1959:372; Mead,

1986:144.

Ganomidos

d’Ailly, 1896:66. Type species by present designation,

Achatina barriana Sowerby, 1890.

Callistopepla

Ancey, 1898:92 (type species: Achatina shuttleworthi Pfe-

iffer, 1856); Thiele, 1929:560; Germain, 1936:151 (foot-

note 3); Verdcourt, 1966:111; Meredith, 1983:30; Oliver,

1983:9; Parkinson, et al. 1987:68; Vaught, 1988:90.

Ganomidus

Boettger, 1905:170.

Ganomides

Verdcourt, 1966:111.

Callistoplepa s.s.

Mead, 1992.

After an extended trip to West Africa, a Captain Vignon

prepared a catalogue of 104 land and freshwater molluscs that

he had collected. The shells and the catalogue were subse-

quently acquired by a collector in Marseille and made avail-

able to his colleague C.F. Ancey, who was given the

opportunity to publish this catalogue. Ancey (1888) agreed to

present it ‘such as it is, but with necessary, even indispens-

able, annotations because of defective identifications, some

of which are not found to be at the level of the science’

(trans.). In one of many footnote annotations, he placed

‘Achatina shuttleworthiana’ [sic] under a new generic name

‘Callistoplepa’. If Ancey was not responsible for the misspell-

ing of the specific name ‘shuttleworthi’, then at least he did

not correct it. The greater misfortune was that he misspelled

the proposed generic name, which as revealed later (Ancey,

1898) was intended to be ‘Callistopepla’ (Gr. most beautiful

robe).

In view of Ancey’s casual manner of publishing the descrip-

tion of this genus, the spelling of the generic name ‘Callis-

toplepa’ must be considered to be the ‘correct original

spelling’ (ICZN Art. 32 (b) and is ‘to be preserved unaltered’.

According to Art. 32 (c), Ancey’s name does not qualify as an

‘incorrect original spelling’ because, ‘without recourse to. . .

external source of information,’ there is no ‘clear evidence of

an inadvertent error’ even though orthographically it would

have been desirable to have spelled it ‘Callistopepla’. His

unorthodoxy and failure to make a timely correction in

spelling suggested that he was content for ten years to leave it

in its original form. In the meantime d’Ailly (1896) unwit-

tingly proposed the generic synonym Ganomidos including

Achatina shuttleworthi along with A. barriana. Further,

Ancey’s original spelling contravenes no provisions of the

Code articles. It is only in his belated publication (1898) that

he used the spelling ‘Callistopepla’, without even implied

justification for the change in spelling. Under the circum-

stances, this constituted an ‘unjustified emendation’ of the

original spelling and therefore it is a junior objective syn-

onym (Art. 33 (b)(iii). Or, perhaps it was just another one of

his regrettable misspellings. This rationale supports Pilsbry’s

conclusions (1905:126), but not those of Germain (1936:151

footnote 3 ). Unfortunately, the confusion about the valid

spelling of the generic name has persisted in collections and

even in the more recent literature, e.g. Parkinson et al.,

1987:68, Vaught, 1988:90. It is hoped that the present expli-

cation finally will obviate any further confusion.

Ancey (1888) gave as the outstanding characteristics of this

new genus its totally different appearance, thin shell, fine

striation, and a colour pattern recalling Orthalicus gallinasul-

tana. @ Ailly (1896) was the first to describe adequately this

taxon, emphasizing the delicate, translucent, shiny, white-

flecked shell, the vertical filiform sculpture, the mammillate

apex, the inflated body whorl, the elongate, serrate-cristate

foot, the hard-shelled eggs, and the unusual radulae of both

Ganomidos shuttleworthi and the then, newly embraced G.

barriana. Pilsbry (1905) accepted broadly d’Ailly’s character-

ization of the genus and emphasized the importance of the

very thin shell, the closely ‘ribplicate’ sculpture and the broad

central tooth of the radula. In addition, he included in

Ancey’s genus Callistoplepa: Ganomidos pellucidus Putzeys,

1898, G. fraterculus Dupius & Putzeys, 1900, Achatina mar-

teli and its subspecies A. m. pallescens Dautzenberg, 1901.

Germain (1909) and Pilsbry (1919) retained this grouping.

Bequaert & Clench (1934c) added to this genus on the basis

of shell characters: Achatina nyikaensis Pilsbry, 1909 and A.

graueri Thiele, 1911. In the present work it is demonstrated

on the basis of the soft anatomy that the taxa added to

Callistoplepa since d’ Ailly (1896) are not congeneric, but are

in subfamily Achatininae.

Bequaert & Clench (1934c:114) were misleading when they

reported that C. barriana and C. shuttleworthi ‘are from

Upper Guinea’. Columbia Lippincott Gazetteer (1952)

defines Guinea as equatorial West Africa from Senegal to

Angola, being divided into Upper and Lower Guinea by the

Niger Delta. van Bruggen (1989) supports the interpretation

that the division is at the Dahomey Gap. In either interpreta-

tion, these species are limited to Lower Guinea. d’Ailly

(1896:70) states that both species live in small numbers in

shady places at the base of tree trunks and under detached

pieces of bark.

Key to Species

Second whorl with thin crescentic threads and granules; last

whorl evenly convex, expanding greatly, four times the length

of the penultimate whorl when viewed dorsally; aperture

length > shell width except in smallest specimens; peripheral

arrow-shaped pattern usually pale and diffuse, occasionally

absent or nearly so; the suture transects a smaller and often

darker pattern; white flecks sparse or abundant, irregularly

distributed; nepionic whorls 3; larger species (6 whorls =

38-52 mm long). Genital aperture complex, large, superfi-

cial; penial retractor inserts on the right columellar retractor

posterior to all other branches; penis tubular; vagina longer

thaniwides Cameroon), INigertaleas.a-n-ce- acces a cee barriana

Second whorl grossly deeply closely and evenly costate; last

whorl subcarinate, expanding proportionately, three times

the length of the penultimate whorl when viewed dorsally;

aperture length < shell width; conspicuous light castaneous

arrow-shaped pattern at periphery, with concentrations of

white flecks tending to alternate with the pattern; a smaller

similar pattern appears subsuturally, but the white flecks

there are more scattered; nepionic whorls 21/2; smaller species

(6 whorls = 26-34 mm long). Genital aperture simple, small,

lacunate; penial retractor inserts on the right columellar

6 A.R. MEAD

Fig. 1 Callistoplepa barriana, basal genital structures (MRAC no. 795.956).

Fig. 2 C. barriana, right branch of columellar muscle showing posterior attachment of the penial retractor.

Fig. 3 C. barriana, right ventrolateral view of penis to show the pilaster in profile. Contraction during preservation has telescoped the apical

penis and forced it and its fibromuscular matrix out of the penis sheath.

Fig. 4 Callistoplepa shuttleworthi, penis sheath, permanently partially evaginated penis, and pilaster (containing the basal vas deferens) are

shown in frontal plane and in dorsal view (UUZM).

Fig.5 Same, ina slightly tangential sagittal plane.

Fig. 6 Same, in ventral view with penis sheath cut longitudinally and spread laterally to show the penis within.

Fig. 7 Same, with penis cut longitudinally and spread laterally to expose the pilaster within. Penis sheath not shown.

Fig. 8 Same, with pilaster cut and spread to reveal the basal vas deferens opening dorsally into the lumen of the penis. The dense

fibromuscular webbing at the junction of the basal vas deferens and penis has been removed for clarity.

NEW SUBFAMILY AND GENUS ACHATINIDAE

retractor anterior to the retractor of the right optic tentacle;

penis permanently partially evaginated; vagina wider than

long. Cameroon, Gabon, Equatorial Guinea (‘Grand Bas-

SAIMMIOALItY ISISUSPECE)) caieliec codes aes cedenssmce tases shuttleworthi

Callistoplepa barriana (Sowerby, 1890)

Pies 23,24

Achatina barriana

Sowerby, 1890:579, pl. 56, fig. 2; von Martens, 1891:30.

Ganomidos barrianum

d’Ailly, 1896:70, pl. III, figs. S—10 (egg), text fig. (radula).

Callistoplepa barriana

Pilsbry, 1904—05:127, pl. 47, figs. 14-17 (egg), pg. ix fig. 2,

pg. xv (radula, ex d’Ailly); Germain, 1909:90; Bequaert &

Clench, 1934c:114.

Ganomidus barrianum

Boettger, 1905:170.

Callistopepla barriana

Dautzenberg, 1921:98; Oliver, 1983:9 (syntype).

SHELL. Shell ovate-conic, very thin, fragile, translucent,

shiny. Whorls 6—6'/s, rarely 6'/2, moderately convex. The

second and third nepionic whorls are nearly straight sided,

but they immediately give way to postemergent rapidly

expanding whorls, producing a mammillate or submammill-

ate, broadly conic spire and a blunt apex. Shallow sutures

form a thin, nearly even line. Last whorl large, convex, 82%

of shell length, range for 4—6'/2 whorls = 78-86% (n = 115),

swelling faintly outward directly below the suture in some

specimens. Aperture broadly ovate, nearly vertical, pale

milky within. Columella thin, slender, slightly to broadly

arcuate, concolorous, squarely to obliquely truncate, inner

rim rolled adaxially. Outer lip thin, nearly evenly arcuate;

joining the periphery at only a modestly acute angle; greatest

width is characteristically midway. Parietal callus scarcely

apparent in unweathered specimens.

From apex to base, the shell ground collar is uniformly pale

fulvous. Superimposed on this, beginning imperceptibly in

the fourth whorl, are two narrow bands of slender yellow-

brown chevrons — one at the periphery, and a less distinct one

transected by the suture. The chevrons in close juxtaposition

have their apices oriented prosocline and are about as wide as

the space between them. Much thinner, more irregular,

paler, parallel sinuate stripes may join the two bands. Speci-

mens with the most conspicuous patterns may have a second

zone of thin, pale, transverse bands between the periphery

and the base of the shell. The peripheral pattern tends to fade

with increased growth. Some specimens may have present

only the sutural band, or a unicolorous last whorl, or an

entirely unicolorous shell except possibly for a slightly darker

transverse band laid down between growth periods. Any of

the whorls may be flecked with minute circular or elongate

white spots (usually ca 0.2-0.8 mm). These are irregularly

and sparsely dispersed, but are especially conspicuous within

the costae of the last whorl. Upon close examination, they are

seen to be a consolidated white powdery substance between

the two periostracal layers. Although some are associated

with shell injuries, their formation is apparently a natural

phenomenon contributing to cryptic coloration.

The most apical portion of each nepionic whorl dips

abruptly at near-right angles adaxially to form a narrow

platform in which is embedded a strikingly uniform series of

minute shallow pits that fringe the suture. This ornamenta-

tion is limited to the nepionic whorls and is the homologue of

the diagnostic grossly costate sculpture in the second nepionic

whorl of C. shuttleworthi. The first whorl is essentially

smooth. Short faint slender crescentic threads and granules,

oriented transversely but aligned spirally in irregular series,

gradually make their appearance in the second whorl. As this

sculpture becomes more organized, the spaces between the

several spiral series seem to form shallow spiral striae. Near

the junction of the third and fourth whorls, a sharp transverse

delineation marks the end of the nepionic whorls, at which

level the threads become more symmetrical and greatly

compressed, but retain their individuality. With continued

growth, the threads remain fairly distinct or become trans-

versely variously fused into costellae, which interrupt or

obliterate in part the shallow spiral striae. Gradually the

threads become more bold and evolve into slender, closely

and very evenly placed prosocline corrugations or costae,

commonly with splitting and anastomosis. The spiral striae

remain superficial, barely transecting the costae. The

depressed cancellate sculpture below the periphery of the

upper whorls gradually becomes more corrugate until an

essentially uniformly costate sculture is finally formed on the

entire forward last whorl of the fullgrown specimen, dimin-

ishing slightly toward the columella and obliterating the

peripheral line of demarkation. The smallest shells may be

vaguely subcarinate.

SOFT ANATOMY. Alcohol preserved specimens available

31/dissected 13. Nigeria: BMNH 1/1; Cameroon: MRAC 2/2,

SMNH 10/4, SMF 14/6, UUZM 4/0. d’Ailly (1896) had access

to 34 alcohol-preserved specimens collected in Cameroon by

P. Dusén, Y. SjOstedt and J.R. Jungner. With the generous

assistance of Dr Ake Franzén, a diligent search was made in

the museums of Stockholm and Uppsala in 1987, but only 14

specimens could be found. There was no evidence of Jungn-

er’s specimens.

The body of the preserved specimen is uniformly grey

fulvous, without any apparent markings. Immediately poste-

rior to the shell, there is a depressed plateau that is fringed by

two prominent dorsolateral ridges, each composed of 12

closely aligned, truncate incisor-shaped elevations.

The most unusual feature of the internal anatomy of this

species is the penis sheath (PS) (Fig. 1). Thickest at its base

(~0.5mm) it diminshes apically to a diaphanous facia

(~0.05 mm) that, in the normal position, enshrouds the

apical penis (P), the most basal part of the vas deferens

(BVD), and the basal portion of the extraordinarily short

penial retractor (PR). As in other achatinids, the origin of the

PR marks the division between P and BVD. In contrast to

that in C. shuttleworthi, the PR inserts on the right columellar

retractor (RCR) posterior to all other branches (Fig. 2). In

the fully mature specimen (Fig. 1) the tapering attenuated

apical P appears to be cramped into a sigmoid fold in this

thinnest apical PS. A dense webbing of muscle and connec-

tive tissue fibrils, originating from the PR, obscures,

entangles and foreshortens the apical folds of the P, even to

the point in the oldest specimens where this tight, wooly mass

of fibromuscular tissue becomes histologically intimately

intermeshed with the substance of the apical penial wall. On

its outer surface, this cocoon-like network forms a smooth,

dense coating over the P that is completely free from the

equally smooth but very shiny inner surface of the PS, thus

allowing free movement between P and PS. Basally, where

the PS is thickest, this fibrous layer conversely becomes so

thin on the surface of the P and so intricately associated with

it, as to be essentially imperceptible. About midway on the P,

the PS suddenly goes from thick to thin. This creates a

transverse line of thin folds that incorrectly suggests the PS

terminates at that level (Mead, 1992, fig. 2). However, when

there is extreme contraction during preservation, the apical

edge of the PS actually does pass basally far enough to allow

the apical structures to elbow out of the PS (Fig. 3). The

contraction emphasizes the bipartite nature of the P: an

apical convoluted, transparently ensheathed portion and a

basal irregularly bulging, opaquely ensheathed portion that

contains the pilaster (PIL). Internally, the most basal P is

longitudinally plicate; above that, including the PIL, the

epithelium is vermiculate-rugate. The PIL is a simple, greatly

thickened, longitudinal, roundly elevated ridge of the ventral

penial wall that strongly projects dorsally into the lumen of

the P. Basally, this ridge terminates into a solid, inverted-

conical, pendulous verge-like process. Although its margins

are not well defined, axially the PIL has a more gross

epithelial texture than the surrounding tissue. The apical vas

deferens (AVD) is a conspicuously uniformly slender conduit

(~1.0 mm in width). It lacks the heavy muscular basal

portion found in C. shuttleworthi, thus the physical support

for the intromittent organ in C. barriana doubtless is pro-

vided by the thick, longitudinal P.

The vagina (V) is a short, nearly uniformly wide conduit,

about one-third the length of the P. Internally, it is lined with

vermiculate-rugate epithelium and is without any apparent

modifications at its junction with the spermathecal duct (SD)

and free oviduct (FO). The muscular FO is as wide or wider

than the V, 2-3 times as wide as the SD, and about as long as

the SD. For their full length, both FO and SD are tightly

bound to each other by fairly regularly appearing small slips

of muscle. The junctions of the AVD/FO and spermatheca

(S)/SD are pulled in close juxtaposition by the tissues of the

sagittal myoseptum. Just apical to this, the capitate S, about

the length of the V, is broadly attached to the basal (uterine)

portion of the spermoviduct. The SD is a thin-walled mostly

uniformly slender conduit about the caliber of the AVD. Five

gravid specimens were examined; three with full data had

been collected near the end of the rainy season in October/

November. For such a relatively small species, the eggs are

quite large (6.8 x 5.4-6.3 x 5.1 mm). Fully gravid specimens

contained 11-15 eggs, all with heavy, calcareous shells and

distributed in the full length of the spermoviduct. The ovotes-

tis acini appear in four or five discrete clusters under the

columellar surface of the right (apical) lobe of the digestive

gland. A talon with a round base and an apical, diverticulate

elongation is present.

The genital atrium (GA) in this species is unique among

the achatinids so far dissected. It is comparatively large and

so shallow that it is essentially a common genital depression,

immediately within which appear conspicuously the male and

female orifices. These latter, like twin craters, are individu-

ally surrounded by low elevated circular walls of smooth

tissue, which contiguously fuse at their inner margines (Fig.

3).

TYPE MATERIAL. Sowerby (1890:579) did not designate a

holotype. The BMNH specimen ‘89:11.19.2 purchased of

Sowerby’ is here designated the lectotype (Figs. 23, 24; Table

1). The slightly damaged and trimmed second syntype,

NMW, 1955:158.832 in the Melvill-Tomlin collection is here

designated a paralectotype (Oliver, 1983). Remeasurements

A.R. MEAD

of the lectotype confirm Sowerby’s figures except for the shell

length, which is 41.0 mm rather than ‘43 mm’. Sowerby’s

illustration is so poorly rendered that it is not precisely

identifiable with either syntype.

TYPE LOCALITY. “Calabar, Africa?’ Nigeria, 4° 57’ N, 8° 19’

E. J.C. Reid of the University of Calabar recently confirmed

this queried locality. Although he has made many excursions

into the ‘relatively undisturbed Oban Hills Forest which

yields a rich fauna’, he found only two (live) specimens along

a permanent stream at Aking (= Awsawmba) 5° 26’ N, 8° 38’

E, 78 km northeast of Calabar. One of these specimens

(BMNH) was examined anatomically and conchologically in

the present study and was found to be typical; the second

specimen is reportedly in the Tom Pain collection (NMW).

DISTRIBUTION. This species has been found essentially along

the entire expanse of coastal Cameroon from M’Bonge (=

Bonge) 4° 33’ N, 9° 05’ E in the north to Itoki 2° 24’ N, 9° 50’

E in the south. Most of the known twenty localities are

clustered in northwestern Cameroon, spilling over into south-

eastern Nigeria and extending inland as far as Yaoundé 3° 52’

N, 11° 31° E; Métet 3° 05’ N, 11° 00’ E; Ebolowa 2° 54’ N, 11°

09’ E and Sangmélima 2° 56’ N, 11° 59’ E. The nine other

localities are in the environs of Victoria 4° N, 9° E. In all

localities, seven were shared with C. shuttleworthi and five

were shared with Leptocala mollicella. Only a single general

locality record was found for Gabon (Verreaux, 1855

NHMB) and no record for Equatorial Guinea; but this

species eventually probably will be found to be limited to the

northern regions of these two countries. Data labels indicate

that specimens were collected in plantations in Kumba 4° 38’

N, 9° 25’ E (bananas), Missellele 4° 07’ N, 9° 25’ E (coca),

‘Buenga’ (oil palm), and in primary forests.

Table 1 C. barriana — Representative shells measurements.

Greatest Aperture Last %

Whorls Length Width Length Width whorlLW/L % W/L

61/4 59.0 32.4 36.1 19.5 49.0 82 55 Bonge

(UUZM)

61/4 SIPS) 30.0 32.8 16.9 46.0 80 52 Victoria

(ZMB)

6 50.7 27.8 30.9 16.8 41.9 83 55 Idenau

(SMF)*

6 49.0 29.9 30.6 17.5 40.8 83 61 Bonge

(SMNH)

5% 44.0 27.4 29.6. 16:36 37-8 86 62 Bonge

(SMNH)

6 41.0 23.3 DAIOP AES 32-3) 19 57 Calabar

(BMNH)

Lect A.

barriana

6 41.0 22.8 26.6 13.4 34.0 83 56 Kumba

(MRAC)

795.173

5/2 36.8 23.0 PB) US PAR tl 62 Idenau

(SMF)

Bibundi

(SMF)

43/4 20.5 13.8 13.2 7.6 16.8 82 67 Bibundi

(SMF)

5) DID, 15.3 16.4 8.8 20.6 82 6

Total specimens examined: 125. Sources: BMNH, CMNH, IRSN, MCZ,

MNHN, MRAC, NHMB, NHMW, NMW, SMF, SMNH, UHZI, UUZN,

ZMB, ZSM.

NEW SUBFAMILY AND GENUS ACHATINIDAE

REMARKS. This species is commonly encountered in collec-

tions and often confused with immature Achatina bandeirana

Morelet, 1866 and A. craveni E.A. Smith, 1881, both of

which have a proportionately much smaller last whorl.

Callistoplepa shuttleworthi (Pfeiffer, 1856)

Figs. 25, 26

Achatina shuttleworthi

Pfeiffer, 1856:34, 1859:603, 1868:216, 1877:275.

Callistoplepa shuttleworthiana

Ancey, 1888:69.

Ganomidos shuttleworthi

d’Ailly, 1896:69, pl. 3, figs. 11-14, text fig. (radula).

Callistopepla shuttleworthi

Ancey, 1898:92; Thiele, 1929:560, fig. 644 (radula).

Callistoplepa shuttleworthi

Pilsbry, 1904—05:127, pl. 47, figs. 18-20, pg. xv (radula, ex

d’Ailly); Germain, 1909:90, 1916:248, pl. 10, fig. 4;

Bequaert & Clench, 1934b:114; Ortiz & Ortiz, 1959:45, pl.

5, figs. 97, 98, text figs. 28-31 (genit. syst., pallial com-

plex, jaw, radula); Zilch, 1959:373, fig. 1352.

Ganomidus shuttleworthi

Boettger, 1905:170.

SHELL. Shell elongate-ovate, extremely thin, very fragile,

translucent with a subdued gloss. Whorls 51/2-5%, rarely 6,

noticeably flattened in profile. A somewhat restricting,

deeply cut second nepionic whorl produces a mammillate

obtuse apex. The following whorls form a slender conic spire

as they descend more rapidly than they expand. Sutures

between nepionic whorls are deep and regular; those between

postemergent whorls are more shallow and only slightly

irregular. Last whorl subcarinate, noticeably so in juvenile

specimens, expanding proportionately, 77% of shell length,

range for 4/6 whorls = 73-83% (n = 60). Aperture

oblique-ovate, external colour pattern sharp and distinct

from within. Columella usually straight, axial, rarely slightly

arcuate, inner rim erect with a cord-like thickened crest;

truncation oblique to very oblique, rarely at right angles.

Between the third and fourth whorls, the crest of the col-

umella rolls abaxially on itself to form a hollow tube, there-

fore an open umbilicus. Between the fourth and fifth whorls,

this tube narrows and solidifies to form a slender axial cord,

which is seen in the full grown shell. This series of changes,

from open to closed, enigmatically has been observed in

several disparate achatinid species, e.g. A. achatina (Linné,

1758) and Archachatina spp. Outer lip of shell thin, skewed

basally, joining the periphery at an acute angle; greatest

width below midway; this is emphasized by the subcarinate

nature of the shell. Parietal callus thin, vague.

Shell ground colour is pale corneous. The first 2!/2 whorls

are unicolorous. Starting near the third whorl, vague round-

ish, very pale castaneous spots appear both at the suture and

periphery. At these two levels, the spots quickly assume

| sharply angulate prosocline arrow-shaped patterns, high-

lighted with a series of parallel transverse elongate white

flecks. Similar flecks, reminiscent of those in C. barriana, are

scattered irregularly over the shell above the periphery,

rarely below. Soon the sutural band fractionates and moves

increasingly into a subsutural zone. The large arrows at the

periphery become spirally closely juxtaposed to form an

essentially continuous dominating colour band. From it,

slender, nearly parallel light castaneous stripes pass sinuously

to the subsutural band and transversely to the columella.

Only rudimentary costae appear in the last part of the

otherwise smooth first whorl. The entire second whorl is

conspicuously and uniformly ribbed from suture to suture

with elevated, deeply cut, nearly orthocline, gross costae, ca

0.2 mm wide (cf Germain, 1916 pl. 10, fig. 4). In the third

whorl the now more prosocline costae are soon reduced to

half their width. At midway in this whorl, an interruption in

the alignment of the costae marks the end of the nepionic

whorls. Gradually the costae become wider and finally regain

their original width in the fifth whorl, only to become

narrower and somewhat irregular in the last part of the sixth

whorl. Faint shallow closely spaced spiral lines, starting in the

second whorl, almost imperceptibly transect the prominent

costae. There is a delicate, greatly suppressed cancellate-

granulate sculpture on a vitreous surface below the periphery

in the upper whorls. This is invaded by the costae in the sixth

whorl until the entire whorl from suture to columella is nearly

uniformly costate. No splitting or anastomosis of the costate

has been observed.

SOFT ANATOMY. Alcohol preserved specimens available

12/dissected 5. Cameroon: SMNH 5/2, UUZM 7/3. All

specimens were collected by Y. Sjéstedt. The only two extant

mature specimens were found by A. Franzén in a medical

laboratory at UUZM. d’Ailly (1896) reported having access

to 11 alcohol preserved specimens, which apparently did not

include Sj6stedt’s Itoki specimens that were available in the

present study.

Body colour as in C. barriana; spade-shaped elevations on

the posterior foot only slightly less prominent.

Without having dissected and deciphered first the relatively

more simple reproductive tract of C. barriana, it would have

been very difficult to interpret the relationships of the genital

structure in this species. In essence, the axis of the basal male

conduit has been greatly foreshortened telescoping the

homologous structures to such and extent that the pilaster

(PIL) on the ventral wall of the penis (P) is pivoted 180°,

forcing the junction of the P and basal vas deferens (BVD)

deeply into the dorsal aspect of the infolded P, i.e., the upper

ventral wall of the P and the most basal part of the BVD are

therefore seen only in the dorsal or lateral views (Figs. 4, 5).

A dense network of muscle and connective tissue fibrils

firmly fixes the structures in this permanently partially evagi-

nated position. This places the aperture of the BVD and the

contiguous subapical part of the PIL into a basal position

within the folded penial wall to take the lead in forming the

intromittent organ at extroversion. The inner smooth shiny

surface of the extremely thin-walled penis sheath (PS) facili-

tates seriatim extroversion: PIL-P-PS and finally genital

atrium, with the BVD and the attenuated penial retractor

(PR) contained axially within the intromittent organ. Figures

6, 7, 8 show at progressively deeper levels of dissection these

relationships from the ventral view. Both PIL and the inner

penial wall are confluent with a deeply rugate epithelium. It

should be noted that since the BVD opens directly into the

lumen of the penial chamber rather than passing through the

accessory organ to open at its apex, a pilaster rather than a

verge (penis papilla) is formed.

The PR is extremely short and, as in the other species of

Callistoplepinae, it and the BVD are held tightly together by

the PS apical to the completely enclosed P (Fig. 9). In

contrast to that in C. barriana, the PR inserts on the right

columellar retractor (RCR) anterior and strongly ventral to

the retractor of the right ommatophore (RRO) (Figs. 10, 2).

Ortiz & Ortiz (1959) missed the diminutive PR in their

dissections and do not show it in their illustrations. Emerging

above the PS, the apical vas deferens (AVD) is a large

muscular thick-walled conduit that in its normal position

reaches to the peniovaginal angle and doubtless serves both

as an ejaculatory duct and a physical support for the intromit-

tent organ. Apical to this, the conduit narrows to half the

calibre and is thinner walled.

The vagina (V) is very short, but two to four times wider

than its length. Near its base, sparse, thin muscle strands

suggest a primordial vaginal retentor. Internally, the V is

muscular, thick-walled and longitudinally deeply plicate.

There is no sharp delineation between it and the broad,

somewhat thinner walled basal spermathecal duct (SD). This

latter is so large that it tends to be positioned partly between

P and V. Both upper SD and free oviduct (FO) are thin-

walled and of about the same calibre. The clavate spermath-

eca (S) is broadly attached to the spermoviduct apical to the

AVD/FO junction. The ovotestis acini are as in C. barriana.

No specimen was found to be gravid, but a single specimen

collected in October in Bonge seemed to be near it with a

very large albumen gland and an inflated spermoviduct. Ortiz

& Ortiz (1959) examined a single specimen from Fernando

P6o Island (Macias Nguema Biyogo) and found the sper-

moviduct completely crowded with four comparatively large

white eggs. Seven dried eggs (MCZ no.219224) measured in

the present study average 4.7 x 3.7 mm. A diminutive talon

is present. In contrast to C. barriana, the genital atrium is an

inconspicuous dimple without superficial embellishments.

TYPE MATERIAL. Pfeiffer (1856) described this species from

Cuming collection specimens, giving the shell size as 5'/2

whorls and the length-width measurements as 34 x 17 mm;

later (1859) he gave aperture measurements as 19 x

11.5 mm. The measurements of the three syntypes in BMNH

do not match those of Pfeiffer, but are reasonably close. The

largest syntype has a damaged and repaired last whorl and the

next largest is atypically slender; therefore the smallest

specimen (Figs. 25, 26; Table 2) is here selected as the

lectotype, the other two becoming paralectotypes. No con-

vincing evidence was found that other syntypes are extant.

Only four other specimens of the 60 examined in the present

study exceeded 30 mm in shell length.

TYPE LOCALITY. The syntypes in the Cuming collection were

reported to be from ‘ “Grand Bassam” Africae occidentalis

(Verreaux)’. All other specimens examined bearing this

locality were sold by shell dealers, viz. Da Costa, Fulton,

Geret, Paetel and Preston, who may have taken their cue for

a locality from the original description. No museum specimen

has been found with a locality record from the 1400 km

stretch of continental Africa between Grand Bassam, Ivory

Coast and the cluster of reliable locality records in northwest

Cameroon. It is suspected that this is a case of still another

erroneous Cuming record. Although under the circum-

stances, we must accept the type locality as ‘Grand Bassam’,

it is probable that Cuming’s specimens came from Cameroon,

or perhaps Gabon. The likelihood of an early secondarily

established population in Grand Bassam prior to 1856 is

extremely remote, for surely authentic collecting records

would have appeared in the meantime. Edouard Verreaux (cf

Crosse & Fischer, 1869) not only collected the syntypes of

this species, but he also collected the single known specimen

A.R. MEAD

Table 2 C. shuttleworthi — Representative shells measurements.

Greatest Aperture Last %

Whorls Length Width Length Width whorlLW/L % W/L

6 34.0 18.7 18.4 10.3 26.4 78 55 ‘G.Bassam’

(MCZ)

83441

57 ‘G.Bassam’

(BMNH)

PLec

54 ‘G.Bassam’

(BMNH)

PLec

59 ‘G.Bassam’

(BMNH)

Lect A.

shuttle-

worthi*

59 Edea

(MCZ)

61 Gabon

(MRAC)

5314

(Preston)

57 Bibundi

(SMNH)

57 Itoki

(UUZM)*

59 Bibundi

(SMF)

66 Gabon

(IRSN)

(Vignon)

6 323 18.4 18.4 10.0 24.5 76

6 31.0 16.7 16.4 DO 22E8 15

5% 30.9 18.3 17.0 10.7 24.0 78

5% 26.9 15.8 14.8 SO 2057 877

SY2 24.1 14.8 14.4 8:0) 19:05 79

Sis 20.2 AES 11.8 Gis! 164180

5 19.4 11.4 2 5:99 1540079

41/2 16.0 10.5 8.9 25) las

Total specimens examined: 60. Sources: BMNH, GNM, IRSN, MCZ, MNHN,

MRAC, NHMW, SMF, SMNH, UMMZ, USNM, UUZM, ZMB.

of C. barriana from Gabon, now in Bern (NHMB). This

raises the suggestion that Verreaux, after collecting in Gabon

and Cameroon, shipped his specimens from Grand Bassam,

which Cuming assumed was the collecting site.

DISTRIBUTION. Leonardo Fea was the first to discover this

species on Fernando Poo Island (= Macias Nguema Biyogo)

of Equatorial Guinea 3° 30’ N, 8° 40’ E (Germain, 1916:249.

Subsequently, Ortiz & Ortiz (1959:45) reported it from Basilé

and Mongola on that island. Nine reliable localities on the

mainland in Cameroon define a limited coastal belt, ca 280 x

120 km with N’dian 4° 55’ N, 8° 53’ E in the north; Métet 3°

05’ N, 11° 00’ E in the east; and Itoki 2° 24’ N, 9° 50’ E in the

south. Other Cameroon localities: Albrechts Hohe 4° 38’ N,

9° 25' E; Mukonje (= Mukonye) 4° 37’ N 9° 30’ E; Bibundi 4°

13’ N, 8° 59’ E; Edéa 3° 48’ N, 10° 08’ E; Lokoundje 3° 13’ N,

9° 55’ E. A specific locality record for Gabon was not found,

but Vignon, through Ancey (1888:69), reports them as rare in

Gabon at the edge of forest streams. They probably do not

extend south of the Ogooué River.

Callistoplepa tiara Preston, 1909 — A Misidentification

‘Preston (1909:183, pl. vii, fig. 9) described Callistoplepa tiara

from ‘Bitze [= Bitye], near the River Ja [Dja], Cameroons’

(3° 01’ S, 12° 22’ E). He indicated neither the collector nor

the number of specimens he had; however, some specimen

labels (BMNH, MRAC) specify that G.L. Bates was the

collector. Between 1908 and 1912, Preston distributed ten

known syntypes, each bearing the full locality information

NEW SUBFAMILY AND GENUS ACHATINIDAE

SSS Shor

oss

Fig.9 C. shuttleworthi, basal genital structures (UUZM).

Fig. 10 C. shuttleworthi, right branch of columellar muscle showing anterioventral attachment of the penial retractor.

Fig. 11 Leptocala mollicella, basal genital structures (MRAC no. 796.850).

Fig. 12

L. mollicella, penis sheath, penial retractor and penial wall cut longitudinally and spread to reveal the pendulous pilaster within

(MRAC no. 795.638).

Fig. 13 Same, cutaway of pilaster to show basal vas deferens joining the penial sacculus, which leads to the aperture of the pilaster.

Fig. 14 L. petitia, basal genital structures (MRAC no. 214.044).

and Preston as the source. These syntypes are currently to be

found in the following museums: BMNH (3:

no.1908.7.1.13-14, and MacAndrew Coll.), NMW (Melvill-

Tomlin Coll. no.1955.158.826; Oliver, 1983:1), IRSN

(Dautzenberg Coll. no.169), ZMB (no.62345), MNHN,

MRAC (no.5760), RMNH, UMMZ (Bryant Walker Coll.

no.142031). These vary in size from 6'/2, 63.0 x 31.3 to Sth,

44.2 x 25.7. Preston probably placed his presumed new

species in Callistoplepa because of the very thin shell, the

Cameroon type locality, and the fact that the size, general

shape and peripheral colour pattern of his specimens were

reminiscent of C. barriana. However, upon examination of

the shell sculpture in the present study, all syntypes were

found to be juvenile Achatina bandeirana Morelet, 1860.

In Cameroon, A. bandeirana and the closely related A.

iostoma Pfeiffer, 1854 and A. balteata Reeve, 1849 are

sympatric and it is not uncommon to find mixed lots of these

three species in museum collections. Preston, himself, appar-

ently had a mixed lot from which his syntypes were selected.

He sent a ‘cotype’ of Callistoplepa tiara to Dupuis (IRSN,

General Coll.); however, its locality record was simply ‘Cam-

eroon’. After Dupuis (1923) examined this specimen, he

concluded that it probably was a juvenile A. iostoma. In 1934,

Bequaert also saw this specimen and confirmed Dupuis’

conclusion. Their identifications were corroborated in the

present study because this ‘cotype’ specimen revealed the

following characters in contrast to those of the syntypes

identified as A. bandeirana: 1) upper whorls not convex, but

form a nearly straight-sided pyramid; 2) apex more acute

rather than blunt; 3) a slight but apparent peripheral carina is

present in the early whorls; and 4) sculpture is formed by

finer, more uniform, elevated beads that do not evolve into

minute prosocline arcuate welts in the sixth to seventh whorls

(cf Bequaert & Clench 1934a fig. 3). This last character is

diagnostic for A. bandeirana; but it is inadequately developed

in the very immature specimen of five to six whorls, thus such

individuals of the three species may appear to be alike.

Dupuis’ unique ‘cotype’ persuaded Bequaert to assume

that all syntypes of C. tiara were juvenile A. iostoma and he

so identified them in collections (BMNH, IRSN, ZMB,

RMNH) and in his publications (Bequaert, 1950:39; B. &

Clench 1934a:13; 1934c:114). Dautzenberg was similarly

impressed and was moved to place with his ‘cotype no.169’ an

added notation, ‘Erreur de Preston, C’est un jeune Achatina

iostoma Pfeiffer’. This was unfortunate because Dautzen-

berg’s specimen, with full C. tiara field data, is shown now to

be an immature A. bandeirana. IRSN thus has a true syntype

in the Dautzenberg Collection and questionable ‘cotype’ in

the General Collection, which latter is here confirmed to be

A. iostoma and not a bona fide syntype. A somewhat similar

situation exists at BMNH, which has three valid syntypes. A

fourth specimen in the Connolly Collection (BMNH

no.1937:12.30.3684) was sent by Preston and labelled ‘Callis-

toplepa tiara Pr.’ (apparently in his writing) but without any

locality data, except ‘Bitz’ in the accession book. Connolly

had his doubts about the identification and relabelled it

‘Achatina ? balteata Rve juv.’ Bequaert also saw it in 1933

and referred to it as A. iostoma. This now proves to be still

another juvenile A. bandeirana and is here considered a

doubtful eleventh syntype of C. tiara.

It should be noted that A. bandeirana is a wide spread,

highly variable Lower Guinea species complex involving A.

b. arenaria Crowley & Pain, 1961; A. b. mayumbensis C. &

P., 1961; A. paivaeana Morelet, 1866, (1868); and A. dohrni-

A.R. MEAD

ana Pfeiffer, 1870. It is found from Cameroon to northern

Angola (7° N—10° S) and fans north and east into Gabon,

Central African Republic, Congo Republic and Zaire. A

study of this complex is in progress.

Preston did not designate a type, but he retained in his own

collection the specimen that was illustrated in his description

of this species. This syntype is here selected as the lectotype

(measurements: 6; 49.4 x 26.7; aperture 30.9 x 13.8; last

whorl 40.0 mm). It is now in Tervuren (MRAC no.5760) and

can be precisely identified by the unique configuration of the

map-like pattern on the last whorl. This pattern is caused by

the irregular lifting up of the thin outer periostracal layer

from the durable inner periostracal layer, allowing an air

space between. This produces blotchy grey-white patches,

which probably provide cryptic coloration. The juvenile and

mature specimens of both A. bandeirana and A. iostoma

commonly have these patches, which have been referred to as

‘hydrophanous streaks’ (Bequaert & Clench, 1934a:15). They

apparently are homologous to the conspicuous white flecks

on the shells of Callistoplepa barriana and C. shuttleworthi

and may have contributed to Preston’s decision to put his

species in this genus.

Leptocala

Petitia

Jousseaume, 1884:171 (non Chitty, 1857); d’Ailly,

1896:71; Bequaert, 1950:138 (type species: Petitia petitia

Jousseaune, 1884).

Leptocala

Ancey, 1888:70, 1898:92 (type species: Achatina mollicella

Morelet, 1860); Thiele, 1929:560; Bequaert & Clench,

1934¢:116; Ortiz & Ortiz, 1959:24.

Achatina (Leptocala)

Pilsbry, 1904:72; Spence, 1928:213; Bequaert, 1950:138;

Zilch, 1959:366; Vaught, 1988:89.

Achatina (Leptocola)

Kobelt, 1910:66 (non Gerstaecker, 1883).

Leptocala (Leptocala)

Bequaert & Clench 1934b:272.

Pilsbry (1904:73, 75) reduced genus Leptocala to subgeneric

rank in Achatina and placed within it his new Section

Leptocallista. Thiele (1929:560) returned Ancey’s Leptocala

to generic rank and retained within it Sections Leptocala and

Leptocallista. Bequaert & Clench (1934b:274) elevated these

sections to genus and subgenus, respectively. In 1950,

Bequaert placed both names as subgenera of Achatina. Zilch

(1959:366) followed suite. The present studies of the soft

anatomies demonstrate that these two genus-group taxa are

in separate subfamilies because the East African Leptocallista

is anatomically allied to Lissachatina and therefore is an

achatinine.

Bequaert & Clench (1934b,c) announced that the Cam-

eroonian Pseudoglessula efulensis Preston, 1908 might belong

to Leptocala and stated that the type could not be located in

the British Museum. The holotype (no.5309) and the

paratype (no.97435) of this species were found during the

present study in Tervuren (MRAC) and clearly proved to

belong to the Subulinidae. Ancey (1888:71) incorrectly

placed Achatina polychroa Morelet, 1866 in Leptocala;

Bequaert (1950:48) believed it belongs in subgenus Pintoa of

Achatina. A final decision depends upon a study of its soft

anatomy.

NEW SUBFAMILY AND GENUS ACHATINIDAE

The ancestral stock of the two closely related, remaining

species in this genus, L. mollicella and L. petitia, probably

became separated in fairly recent times by a vicariance event

— possibly the development of the Ogooué River.

Because of the unique microsculpture and the somewhat

smaller shell aperture, Leptocala up until now has escaped

suspicion of being closely related to Callistoplepa. The genus

is limited to the southwestern portion of Lower Guinea from

northwestern Cameroon to far western Zaire.

Key to Species

Shell 6-6'2 whorls; spire conic; exceedingly fine distinct

vertical and spiral lines form shallow minute engraved rhom-

boids. Pilaster verge-like, cylindrical, vertically suspended

from the apex of a dome-shaped penis; basal vas deferens

obscured by penial retractor. North of Ogooué River in

Gabon, Equatorial Guinea, Cameroon and probably south-

SINE etter alae occa ceemsacovadelasoaanest kosdae mollicella

Shell 6/7 whorls; spire slender conic; exceedingly fine

closely appressed vertical vermiculate-granulate sculpture

obliterates the spiral lines, especially on the upper whorls.

Pilaster potato shaped, somewhat compressed, attached for

nearly its full length along a diagonal right ventrolateral axis

of a hull shaped penis; basal vas deferens conspicuous in

ventral view. South of Ogooué River in Gabon, Congo

Republic, western Zaire and probably Cabinda, Angola-

ion necoe « vugled ges AGERE DR SRBC Ge eee An na ses <AAMBA ee Ree aes petitia

Leptocala mollicella (Morelet, 1860)

Figs. 27, 28

Achatina mollicella

Morelet, 1860:189; Pfeiffer, 1868:216; 1877:275; Vignon

(in Ancey, 1888:70); Pilsbry, 1904:29.

Achatina pulchella

von Martens, 1876:258, pl. 3, figs. 1, 2 (syntype) (non Spix

& Wegner, 1827; non Pfeiffer, 1857); Ancey, 1888:70;

Pilsbry, 1904:73, pl. 34, fig. 14 (ex von Martens).

Leptocala mollicella

Ancey, 1888:70, 1898:92; Thiele, 1929:560; Bequaert &

Clench, 1934b:273.

Achatina smithi

Sowerby, 1890:579, pl. 56, fig. 3 (holotype, monotypy;

non Craven, 1880).

Achatina sowerbyi

E.A. Smith, 1890:392 (new name for A. smithi).

Petitia pulchella

d’Ailly, 1896:71; Boettger, 1905:170.

Achatina (Leptocala) mollicella

Pilsbry, 1904:73; Spence, 1928:213, pl. 2, fig. 5; Bequaert,

1950:138; Zilch, 1959:366, fig. 1342.

Achatina (Leptocala) pulchella

Germain, 1916:154, 241, pl. 6. figs. 11, 12.

Leptocala (Leptocala) mollicella

Bequaert & Clench, 1934b:273.

Leptocala mollicella zenkeri

Bequaert & Clench, 1934c:118, pl. 1. figs. 5-7, pl. 2, fig.

13 (holotype, 3 paratypes).

Leptocala mollicello zenkeri

Bequaert & Clench, 1934¢:119 (/apsus calami).

Achatina (Leptocala) mollicella petitia

Bequaert, 1950, pl. 58, fig. 4.

Achatina (Leptocala) mollicella zenkeri

Bequaert, 1950:138.

Leptocala pulchella

Ortiz & Ortiz, 1959:25, pl. 5, fig. 99.

SHELL. Shell obovate, glossy, translucent, thin but sturdy;

periostracum tenaceous. Whorls 6—-61/2, moderately convex.

Spire conic; apex broadly obtuse; sutures moderately deep,

fine, straight or slightly irregular. Last whorl expanding at a

somewhat greater rate than the upper whorls, 72% of shell

length, range for 4°4-61/2 whorls = 69-75% (n=34). Aperture

elongate inverted ear-shape, pale milky within. Columella

short, variably straight to slightly arcuate and twisted, trans-

versely to obliquely truncate, basal crest slightly elevated in

juvenile specimens. Outer lip thin, joining the periphery at a

broadly acute angle. Parietal callus minutely granular, shiny,

concolorous.

Shell ground colour is pale fulvous, rarely somewhat

darker. Most specimens have a distinct but subdued pattern

of pale yellow-brown, slender, strongly parallel, nearly

straight or somewhat sinuous stripes, usually 0.2-0.3 mm

wide, alternating with ground colour bands of about the same

width. The banding may be slightly coarser and more con-

spicuous in the fifth whorl and above. Often apparently

unicolorous or weathered specimens under proper lighting

and magnification will be seen to have this characteristic

pattern at least in limited areas. This is witnessed in Bequaert

& Clench’s (1934c) figure 7 of their Leptocala mollicella

zenkeri, which they report is ‘without any darker markings’.

The first 1—-1'/ whorls are smooth and very shiny. Short

vertical or arcuate lines begin to appear in the second whorl,

often concentrated at the suture below. These soon elongate

into delicate, narrow, closely packed vertical lines that span

the full width of the whorl. At 2'2 whorls, there is a

conspicuous diagonal demarkation between the nepionic and

the postemergent whorls. At this demarkation, spiral lines

that immediately previous to this were sparce, short and

ghost-like, quickly form 35-40 nearly evenly spaced exceed-

ingly shallow, but sharply engraved lines. These lines cross

the vertical lines and cut the surface into minute engraved

thomboids (see Bequaert & Clench, 1934c fig. 13). In the

following whorls, these spiral lines become more numerous

and somewhat wavy, suggesting the surface had been evenly

and shallowly combed. Beginning at 2'/2 whorls, subtile

prosocline, more sparce growth wrinkles compete with the

vertical lines; these may impact the suture directly or arcu-

ately. There is little reduction in the intensity of this engraved

pattern below the periphery, although there is a slight reduc-

tion in caliber. In the last half of the last whorl, the rhomboid

pattern may essentially disappear, leaving the growth

wrinkles to dominate. Throughout, the shell characteristically

remains remarkably smooth and shiny. A subcarina is present

in the early whorls, but this disappears in the fifth whorl.

SOFT ANATOMY. Alcohol-preserved specimens available

12/dissected 4. Cameroon: MRAC 3/3, UHZI 1/0; ZMB 7/0;

ZMUC 1/1.

Body colour of preserved specimens, including the head

and anterior edge of the mantle, is pale-cream fuscous.

Dorsally, there is a diffuse grey wash that shades darker

anteriorly. A 1-2 mm wide dark gray band with diffuse

borders, appears immediately behind the ommatophores and

extends posteriorly along the middorsal neck to the edge of

the mantle. Surface of body reticulate-microtuberculate. Foot

without structural elaborations.

The most prominent feature of the genital system (Fig. 11)

is the massive, muscular basal female conduit that seems to

be an ill defined fusion of vagina (V), spermathecal duct (SD)

and free oviduct (FO). Sessile to this, obliterating the penio-

vaginal angle, is the contrastingly short, broad penis sheath

(PS). Crowded together and projecting from the apical collar

of the PS are the long apical vas deferens (AVD) and the very

short penial retractor (PR). In fact, this latter is so short that

only its insertion on the broad right columellar retractor

(RCR) may be seen. Like that of L. petitia and Callistoplepa

barriana, it inserts far posterior on the RCR (Fig. 2). The

tripartite AVD starts as a slender tube, but soon enlarges to

form the thick-walled ejaculatory duct. This in turn narrows

and then once again enlarges into a thin-walled conduit

(possibly a secondary seminal vesicle) before fusing with the

FO to form the spermoviduct. When the PS is cut longitudi-

nally, it is found to be extremely thin and attached basally

only about half way down on the stubby, dome-shaped penis

(P). Both the inner surface of the PS and the outer surface of

the P are smooth and shiny, facilitating extroversion. In this

aspect, the PR is seen to hood over the apical P and, about

one-quarter the way down, blend with the substance of the

thick-walled P. When the PR is split longitudinally and its

muscle bands spread apart (Fig. 12), the basal vas deferens

(BVD) is seen within, discreet and without muscle or connec-

tive tissue attachment. Vertically cutting the penial wall

reveals a conspicuous, pendulous, vertically oriented pilaster

(PIL) whose thick transverse-diagonally textured brownish,

glandular epithelium is continuous with the inner wall of the

penis. Ventrolaterally on the PIL is a 1 mm vertical eccentric

apertural slit. This leads internally to a funnel shaped penial

sacculus, which joins the extension of the BVD in a dense

mass of connective tissue. It is clear at this point that the PIL

has been formed by a permanent partial eversion of the P. In

the process, the basal-most BVD, with its diagonal anchoring

muscle strands, forms the axial conduit of the PIL (Fig. 13).

During extroversion, to form the intromittent organ, the PIL

would take the lead, followed by the basal P, and finally the

PS, which would contain the BVD, PR, and a portion of the

RCR. Both PR and RCR would be involved in the introver-

sion process.

The alignment of the V, SD and FO insures that the

intromittent organ will be channeled directly to the SD. This

has been accomplished not only by a large knob of tissue

eccentrically blocking the narrow lumen of the FO, but also

by a massive buildup of muscular tissue surrounding the large

lumen of the basal SD. These modifications, in turn, tend to

wedge the SD between P and FO (Fig. 11). Internally, there

is a Sharp division between the thick-walled, heavily muscular

V, with its many narrow, tightly compressed, vertical plicae,

and the thick-walled basal half of the SD (functionally an

extension of the V), with its several bold, deep, coarse

vertical plicae. Cutting across these latter plicae are trans-

verse vermiculate rugae that produce a grossly serrate texture

on the crests of the plicae. This rough texture seems to

complement that of the P and PIL. All these structures

obviate the necessity of a distinct muscular vaginal retentor

found in many species of Achatininae. The apical SD is

thin-walled, as is the clavate spermatheca (S). The apical

saccular FO probably serves to hold the large egg immedi-

ately before expulsion; basally, however, it is thick-walled,

with a narrow lumen. An ‘elbow’ tends to form at the

junction of the two parts. The lower portion and the collar-

like thick-walled V doubtless serve as an ovijector.

A.R. MEAD

Three of the four dissected specimens had the uterus and

oviduct crowded with 4, 6 or 8 relatively large, off-white

hard-shelled eggs, measuring 4.3 x 3.7-5S.1 x 4.4 mm. Their

long dimension is ca. 15% of the adult shell length, which is in

strong contrast, for example, to that relationship in Achatina

achatina at ca. 5%. All gravid specimens were collected

September—November, just before the long dry season, and

because of the demands of producing eggs, they manifested

considerable emaciation, especially in the digestive system.

Such observations raise the unanswered question of longevity

in this small species. In a single specimen, six ovotestis acini

were found under the columellar surface of the apical lobe of

the digestive gland. The talon is extraordinarily long and

slender and without a basal enlargement.

TYPE MATERIAL. The type of Morlet’s Achatina mollicella,

collected by Vignon, has never been illustrated and its very

existence has been uncertain. It is not in the Morelet material

in the Paris, Geneva or Tervuren collections. Fulton (1920)

reported that he had purchased the Morelet land and fresh-

water shells in 1892, including ‘all the types’, but that in

transit between Dijon and London many of the fragile

specimens, including some types, were broken. The BMNH

accession book under date 2 April 1893 confirms this infor-

mation. It lists the accessioned types (pp. 230-254, 2049

entries) including a single entry indicating that there were

only two specimens: ‘93.2.4.119-120 Achatina mollecella,

Gabon’. However, neither of these two syntypes have the

length-width measurements of ‘18 x 12 mm’ given by More-

let (1860) for an individual 6 whorl specimen. Under the

circumstances, it is most likely that the 6 whorl syntype is the

one Morelet had in hand. Further, there clearly is an error in

his reported measurements because in the 35 specimens

examined in the present study, the shell width averages 52%

of shell length, not 67% as would be the case if Morelet’s

measurements were correct. For these reasons, the larger

syntype BMNH no.93.2.4.119 is here selected as the lecto-

type (Figs. 27, 28; Table 3) and BMNH no.93.2.4.120

selected as paralectotype. Morelet’s incorrect measurements

have contributed greatly to the confused synonymies of his

valid species and Jousseaume’s valid species Petitia petitia.

The lectotype and 10 paralectotypes of von Martens’ junior

subjective synonym Achatina pulchella are in Berlin (ZMB;

Kilias, 1992), type locality Bonjongo, Cameroon. Additional

single specimens here designated as paralectotypes have been

found and labeled in Stockholm (SMNH no.4282) and Ter-

vuren (MRAC no.5315). The holotype (monotypy) of Sower-

by’s junior subjective synonym and junior primary homonym

Achatina smithi no.89.11.18.1 is in London (BMNH), type

locality ‘Calabar, Africa?’ The holotype of Bequaert &

Clench’s Leptocala mollicella zenkeri (1934c fig. 6) plus two

paratypes (their fig. 7 and one unfigured) are in Berlin

(ZMB,; Kilias, 1992). A third paratype is at Harvard (MCZ)

under No. 98687, which was identified by the distinctive mark

at the junction of the ultimate and penultimate whorls (cf.

their fig. 5). All these were from Yaoundé. A fourth

paratype, unfigured but listed on their page 119 is from Bitye

(BMNH, no.1908.6.3.2; Table 3).

TYPE LOCALITY. Morelet (1860) lists it as, ‘Habitat, rara, in

sylvis Guinea.’ His two syntypes in BMNH were more

specifically labeled ‘Gabon,’ which is included in the early

broad generic geographic term ‘Guinea.’

DISTRIBUTION. Sowerby (1890) described Achatina barriana

NEW SUBFAMILY AND GENUS ACHATINIDAE

Table 3 L. mollicella— Representative shells measurements.

Greatest Aperture Last %

Whorls Length Width Length Width whorlLW/L % W/L

61/4 39.8 18.7 17s OST 27 10 47 Bitya

(BMNH)

Para L. m.

zenkeri

1908.6.3.2

6h S75) Wee: 17.3 8:9) 926:5. 711 47 Ebalowa

(UMMZ)

61/4 35.8 7S 18.4 8.7 26.5 74 49 Yaoundé

(UMMZ)

6h 33.3 WEB: SO LOSI 237470. 52 Olounou

(MRAC)

796.850

6 30.5 16.1 15.6 8.2.) 2253073 53 Kribi

(MRAC)

795.638"

14.8 15.0 TS eA ail 50 Calabar

(BMNH)

Holo A.

smithi

6 26.7 13.4 12.8 6.2 18:9 71 50 Gabon

(BMNH)

Lect. A.

mollicella

93.2.4.119

6 26.6 14.0 13.6 6.9) 192k) 72 53 Bonjongo

(MRAC)

5315 PLec

pulchella

5, 23.6 Sei 12.4 6.6) 17.3: 73 55 Gabon

(BMNH)

PLec A.

mollicella

93.2.4.120

49/4 16.6 10.4 9.1 SO ler 73 63 Nyong

(ZMUC)'

Total specimens examined: 35. Sources: BMNH, IRSN, MNHN, MRAC,

NHMW, NMW, SMNH, UMMZ, ZMUC.

and A. smithi (= L. mollicella) at the same time and indicated

for both that the locality was ‘Calabar, Africa?’ (4° 57’ N, 8°

19’ E). Both specimens were from the Cuming collection, in

which a number of other locality records from time to time

have been questioned or proven incorrect. J.C. Reid of the

University of Calabar has collected near Calabar what is now

confirmed as Callistoplepa barriana, but so far no L. molli-

cella. It is altogether possible that this latter species eventu-

ally will be found in Nigeria because Mbonge, Cameroon, a

known endemic locality for this species, is only ca. 70 km to

the southeast in a similar environment. Ortiz & Ortiz

(1959:26) have reported the western-most records for this

species from four localities on Fernando Poéo Island (=

Macias Nguema Biyogo) of Equatorial Guinea 3° 30’ N, 8°

40’ E. Seventeen locality records on the continent cluster in

the northwestern corner of Cameroon, with the extremes

being Mbonge 4° 33’ N, 9° 05’ E in the North, Molobo 4° 01’

N, 14° 19’ E in the East, and Efulen 2° 42’ N, 10° 30’ E in the

South. Vignon through Ancey (1888:70) records this species

as being very rare in the forests of Gabon. It probably is not

found south of the Ogooué River.

In the specimens examined, there was a high direct correla-

tion between greater shell size and distance from the sea-

coast, e.g. the largest specimen seen in this study is from

Bitya on the river Dja, ca. 260 km from the coast 3° 01’ N, 12°

22’ E (BMNH no.1908.6.3.2; Table 3).

Leptocala petitia (Jousseaume, 1884)

Figs. 29, 30

Petitia petitia

Jousseaume, 1884:172, pl. 4, fig. 4a, holotype, monotypy

(non Chitty, 1857); Bequaert, 1950:138.

Achatina (Leptocala) mollicella petitia

Pilsbry, 1904:73, pl. 34, fig. 15 (ex Jousseaume, 1884);

Bequaert, 1950:138, pl. 58, fig. 4.

Leptocala mollicella petitia

Bequaert & Clench, 1934b:273.

SHELL. Shell ovate-elongate, thin but not fragile; last whorl

shiny, upper whorls less so. Whorls 6'/2-7, moderately con-

vex. Spire slender conic; apex narrowly obtuse; sutures fine,

almost without irregularities. Last whorl expanding propor-

tionately to upper whorls, 70% of shell length (n=7); fourth

and fifth whorls subcarinate. Aperture elongate-oval, milky

within. Columella short, straight, transversely to obliquely

truncate. Outer lip thin, arcuate, joining the periphery at an

acute angle. Parietal callus minutely granular, shiny, concol-

orous.

Shell ground colour is pale corneous. At the junction of the

fifth and sixth whorls, diffuse yellow-brown stripes

(0.3-0.5 mm wide) alternate with wide ground colour bands

(0.5—0.7 mm); these are approximately the same width on the

early whorls, but become slightly or much narrower, more

distinct and closer together on the last whorl, or nearly

disappear; they may be variously straight, diagonal or

rippled.

The last quarter of the first whorl has nearly imperceptible

surface irregularities that originate close to the suture, where

they evolve into a series of closely packed crescentic lines.

They quickly multiply axially into five or six horizontal series

of short crescentic lines. These gradually fuse vertically to

form very narrow, crowded, thread-like, prosocline welts. A

fairly conspicuous diagonal line, near mid third whorl, marks

the end of the nepionic whorls. Near there, the welts become

superficially engraved with a vertically oriented, exceedingly

fine vermiculate-granulate sculpture, which is reminiscent of

the much coarser sculpture of Achatina (Tripachatina)

vignoniana Morelet, 1874. Gradually, the welts diminish and

the more sparce growth lines emerge, leaving the rash-like

microscopic sculpture to dominate. This is best seen in

subdued light at a low angle. The sculpture may diminish and

become more sparce between the third and fourth whorls, as

it does in the holotype, or it may continue at essentially the

same caliber until the fourth or fifth whorl. At a certain point

in the diminution, and if the light intensity is properly

adjusted, ghost-like, engraved spiral lines, here and there,

spaced as in L. mollicella, can be distinguished, especially

below the periphery, where the sculpture is somewhat

reduced in calibre. The sculpture may extensively obscure

these spiral lines and all but traces of a rhomboid pattern, or

it may become so sparce on the lower whorls as to allow the

sharp spiral lines to dominate. It is almost as if the

vermiculate-granulate sculpture were superimposed upon the

typical sculpture of L. mollicella in a variably decreasing

intensity from apex to base. As a result, the surface of the

16 A.R. MEAD

s\\ SSS

Fig. 15 L. petitia, penis sheath and penial wall cut diagonally in right ventrolateral view and spread to expose the pilaster. The cutaway

shows the basal vas deferens joining the aperture of the pilaster (MRAC no. 214.044 & 212.583).

Fig. 17 Bequaertina pintoi, basal genital structures (NM).

Fig. 18 B. pintoi, penis sheath cut and spread laterally. Conduits transected. Most of the obscuring basal eversion muscle bands have been

removed.

Fig. 19 B. pintoi, hermaphroditic duct system.

Fig. 20 Bequaertina graueri, basal genital structures (MRAC no. 610.342 & 610.303).

Fig. 21 8B. graueri, penis sheath cut and spread.

Fig. 22 8B. graueri, frontal plane through basal male and female conduits. Bar scale = ~ 1 mm. A.R.M. del.

NEW SUBFAMILY AND GENUS ACHATINIDAE

shell has less luster than in this latter species. This peculiar

microscopic sculpture on the upper whorls is determinative.

SOFT ANATOMY. Alcohol preserved specimens available

2/dissected 2. Congo Republic: MRAC 1/1; Zaire: MRAC

1/1. These apparently are the only such specimens extant.

Both had small body masses and were withdrawn far into

their thin shells because they were collected during the dry

season and were inadequately drowned before preservation.

However, most of the soft parts were successfully extracted

with only minimal damage to one shell.

Body colour and texture as in L. mollicella.

Upon exposing the reproductive tract (Fig. 14), the most

noticeable anatomical feature is that both the unusual hull-

shaped penis (P) and the large basal vas deferens (BVD)

show through the thin, nearly transparent, but substantial,

penis sheath (PS). Typical of the Callistoplepinae, the PS also

enshrouds the very short penial retractor (PR). As in L.

mollicella and Callistoplepa barriana, this latter inserts far

posterior on the large right columellar retractor (RCR). Also

conspicuous is the apparently inordinately long bipartite

apical vas deferens (AVD), with a nearly evenly broad

muscular basal portion and a thin-walled, somewhat undulant

apical portion. When the PS was cut vertically and the edges

pulled laterally, it was found to extend essentially to the base

of the P. In the first dissected specimen, from Lukula, Zaire,

the exposed, large BVD appeared out of proportion and

excessively deeply wedged into the ventral surface of the P.

Similarly, the navicular P, with its diagonal left ventrolateral

orientation, seemed enigmatically distorted. But the second

specimen, from Kayes, Congo Republic, ca. 180 km to the

north, had almost identical proportions and alignment, thus

essentially removing the suspicion that there had been exces-

sive distortion. The relatively thin penial wall of the first

specimen was cut along a midventral, vertical line. Immedi-

ately below the surface was a large, obstructing mass of penial

wall tissue whose angulate orientation could not safely be

explored. Consequently, in the second specimen, a diagonal

cut was made along the long axis of the oblong P. This

revealed in right ventrolateral aspect a comparatively huge,

somewhat compressed potato-shaped pilaster (PIL) attached

diagonally along nearly its full length ventrolaterally on the

inner basal penial wall, parallel to the adjacent crowded BVD

(Fig. 15). In essence, the wall of the basal half of the P was

hardly more than a thin, tight-fitting cover for the PIL. The

surface of the PIL and the inner wall of the P, similar to that

of L. mollicella, was covered with transverse, diagonal,

anastomosing rugae. Irregularities in the rugae revealed a

small basal aperture. Cutting basally into the 2.6 mm PIL

exposed the short basal-most BVD narrowing rapidly

through dense connective tissue to connect with this aperture.

Apically, the PIL is a solid mass of penial wall tissue.

Collectively, the relationships in these structures are reminis-

cent of those in C. shuttleworthi, particularly with respect to

the exposed BVD pushing ventrally far down into the par-

_ tially evaginated P (Figs. 4, 5).

In this species the basal female conduit, externally and

_ internally, is much less gross than in L. mollicella. The vagina

(V) is a distinguishable, more slender porion of the conduit.

Similarly, the basal spermathecal duct (SD) is less muscular

and tends less to interject itself between PS and the free

oviduct (FO). However, the FO, muscular at the base and

thin-walled apically, is less robust yet comparatively more

prominent in this species. Although shown spread apart in

Fig. 14 for clarity, the FO and basal SD are actually held

tightly together by many small, short muscle bands, probably

providing support for the SD at termination of copulation. In

that natural position, the clavate spermatheca (S) is attached

by thin muscle bands and connective tissue to the uterine

portion of the spermoviduct, well above the junction of AVD

and FO. The eggs are not known but are probably on a par

with those of L. mollicella.

TYPE MATERIAL. The holotype (monotypy) (Figs. 29, 30;

Table 4) in the Jousseaume collection in Paris (MNHN) was

collected by L. Petit.

TYPE LOCALITY. At the mouth of the River N’toc, which

disappears in the Mayumba Lagoon, Gabon 3° 25’ S, 10° 39’

DISTRIBUTION. Gabon: type locality. Congo Republic: Sibiti

3° 41’ S, 13°21’ E (SMNH), Kola 4° 03’ S 11° 44’ E(MRAC),

Kayes 4° 26’ S, 11°23’ E(MRAC). Zaire: Lukula 5° 21’ S, 13°

02’ E(MRAC). All known localities are south of the Ogooué

River of Gabon. This species will probably be found in

Cabinda, Angola.

REMARKS. In addition to the holotype in Paris (MNHN),

there are only six known specimens of this species, five in

Tervuren (MRAC), collected by Dartevelle, and a single

specimen in Stockholm (SMNH). The explanation for its

apparent rarity probably rests in the fact that there has been

much less professional collecting in south coastal Gabon and

the Congo Republic than in Cameroon, where L. mollicella is

not a rarity. In this limited number of specimens extant, there

is a north to south gradient of more intense vermiculate-

granulate sculpture and reduced rhomboid pattern. If the

substantive differences in the soft anatomies had not been

known, this taxon might well have been assumed to be no

25° 30° 35° 40°

, SOMALIA J»

(

O ucanoa Z

KENYA

RWANDA

ZAIRE

Sane 24 s°

a7 TANZANIA

25° 30° 35° 40°

Fig. 16 Distribution of Bequaertina. O = Bequaertina fraterculus,

@ = B. graueri, 0 = B. pellucida, @ = B. pintoi, A = B. marteli.

Where possible, all localities were checked with the volumes of

the U.S. Board on Geographic Names.

Table 4 L. petitia — Representative shells measurements.

Greatest Aperture Last %

Whorls Length Width Length Width whorlLW/L % W/L

21.5 66 45 N’toc

(MNHN)

Holo *

61/2 30.9 14.2 1457 72 e2ArA Soo 46 Sibiti

(SMNH)

61/4 30.8 14.8 1339 S72 A SGO9: 48 Lukula

(MRAC)

212.583*

6'/2 29.0 13.7 14:0) 6s 20217169 47 Kola

(MRAC)

196.340

6 28.6 14.6 14.59 /Alee20ES, 72 51 Kola

(MRAC)

196.341

6 23.9 12.3 11.6 6.0 16.4 69 51 Kayes

(MRAC)

214.004*

5 20.8 13.3 bles Gs) Sly 7 64 Kola

(MRAC)

791.389

oT Si}) 14.8 ahs} Saeco)

Total specimens examined: 7. Sources: MNHN, MRAC, SMNH.

more than a subspecies of L. mollicella, as Bequaert &

Clench concluded (1934b:273). This case is reminiscent of the

conchologically very similar but anatomically contrasting

Achatina reticulata Pfeiffer, 1845 and A. albopicta E.A.

Smith, 1878 (Mead, 1950:232).

Jousseaume’s illustration of the holotype is misleading

because the artist has shown the apex acuminate; actually,

the first and second whorls are noticeably larger, producing a

narrowly obtuse apex.

Apparently neither Leonardo Fea (Germain, 1916) nor

Captain Vignon (Ancey, 1888) went far enough south in

Gabon to encounter true L. petitia.

ACHATININAE

Bequaertina new genus

Thin, fragile, anomphalous, medium to large, ovate to ovate-

elongate shells, 40-80 mm in length. Spire tends to be

mammillate, apex obtuse. Aperture large, columella long and

slender, squarely or obliquely truncated. Whorls 6-6/2, rarely

7; second and third nepionic whorls sculptured; last whorl

ventricose, ca. 80% of shell length. Sculpture may be vari-

ously cancellate-granulate, lirate, malleate or nearly smooth.

Surface of shell lusterless; its abrasion reveals a brilliant inner

periostracal layer. An occasional specimen may show in the

periostracum of the lower whorls limited areas of an

extremely fine decussate micromesh, commonly seen in a

wide variety of achatinids.

The generic characters in the soft anatomy are based on

features that are shared by the two available species — B.

pintoi (Bourguignat, 1889) and B. graueri (Thiele, 1911).

Because of the similarity in the basic anatomical pattern in

these two species, and because, on the basis of shell charac-

ters, each of the species represents a different dichotomous

group, it is felt that the following anatomical characters will

prove to be valid for the genus.

The most prominent features of the genital system are the

long free oviduct, the apical vas deferens and the large,

A.R. MEAD

elongate sacculate spermatheca — all held in close juxtaposi-

tion by a distinct fascia. In contrast, the penis and penis

sheath are inconspicuous. The penis sheath enshrouds a short

basal portion of the long vas deferens. Without exception in

26 dissected specimens, the penial retractor inserts on or near

the diaphragm where the latter joins the mantle and the body

wall of the neck region. At the origin of the penial retractor,

muscle fibrils pass snugly over the apical penis and then fan

out into a network that covers the basal vas deferens and the

inner wall of the penis sheath, except for a limited smooth,

shiny zone on the approximately upper half of the left side.

Below this, the fibrils infuse intimately with the tissues of the

basal penis and penis sheath to create an ill defined section of

the male conduit that contains the penial atrium. This atrium

connects the lumen of the penis with the genital atrium. At

this level, abundant hypertrophied eversion muscle bands

obscure the genital atrium and its junction with the male and

female conduits. There is no pilaster or verge.

The spermathecal duct is much shorter than the spermath-

eca and often sessile on the vagina. This significantly places

the spermatheca in a basal position with its usually attenuated

apex stretching to its connection by fascia to the junction of

the free oviduct and the apical vas deferens, well below the

spermoviduct. The hermaphroditic duct has in its midsection

an enlarged glandular structure of unknown function. The

ductules to the five hermaphroditic acini are inordinately

gross and highly convoluted.

The eggs are covered with a hard calcareous shell and are

proportionately larger than those of Achatina, i.e. on a par

with those of Tholachatina (sensu Bequaert, 1950). There is

no evidence of ovoviviparity.

The anterior aorta is on the floor of the diaphragm and

passes ventrally through the diaphragm to the sagittal myo-

septum in the haemocoele. The second largest vein in the

lung drains the region near the extreme left mantle and joins

the primary vein near the apex of the kidney. The large last

whorl of the shell allows for a highly vascularized left side of

the lung. The secondary ureter is completely closed. The

rachidian tooth of the radula is either slender and question-

ably functional or broad and about half the size of the

adjacent laterals. The jaw is narrow and broadly arcuate,

with many slender vertical ribs irregularly distributed.

Six specimens of B. pellucida (Putzeys, 1898) and one of B.

marteli (Dautzenberg, 1901), as well as several of Achatina

craveni, have been found with a single, almost perfectly

circular hole, 0.6-4.0 mm in diameter, cut usually in the

dorsal part of the last whorl. These are thought to be caused

by bird pecks (Meredith, 1983a:25).

The five species in this genus were place in the genus

Callistoplepa on the basis of similar shell characters: thin

shell, large aperture and a tendency to form a mammillate

spire (Pilsbry 1919, Bequaert & Clench 1934c). But as shown

in the Key to Subfamilies, a study of the internal anatomies

revealed major differences. Bequaertina reflects strongest

phylogenetic affinities to subgenus Achatina (sensu Bequaert,

1950), particularly with respect to the configuration of the

basal male conduit and to the fact that the spermatheca is

attached to the adjacent free oviduct and apical vas deferens

rather than to the spermoviduct. In Bequaertina, the apical

penis is free from the apical penis sheath and therefore can

evert independently, with the sheath following seriatim at

extroversion, whereas in subgenus Achatina the penis is

completely enmeshed in a dense network of muscle fibrils and

connective tissue that requires the penis and the sheath to

NEW SUBFAMILY AND GENUS ACHATINIDAE ~

Figs 23, 24 Callistoplepa barriana (Sowerby, 1890); lectotype Achatina barriana BMNH no. 1889.11.19.2. 25, 26 C. shuttleworthi (Pfeiffer,

1856); lectotype Achatina shuttleworthi BMNH. Bar scale + ~ 10 mm.

Figs 27, 28 Leptocala mollicella (Morelet, 1860); lectotype Achatina mollicella BMNH no. 93.2.4.119. 29, 30 L. petitia (Jousseaume, 1884);

holotype Petitia petitia MNHN. 31, 32 Bequertina pellucida (Putzeys, 1898); lectotype Ganomidos pellucidus MRAC no. 5135. 33, 34 B.

pellucida; paralectotype G. pellucidus (unicolorous) MRAC no. 5136. 35, 36 B. pellucida; lectotype Serpaea foai Germain, 1905 MNHN.

37,38 8B. marteli (Dautzenberg, 1901); lectotype Achatina marteli IRSN. 39, 40 B. marteli; lectotype A. marteli pallescens (Dautzenberg,

1901) IRSN. 41, 42 B. pintoi (Bourguignat, 1889); holotype, Serpaea pintoi MNHN. Bar scale = ~ 10 mm.

Figs 43, 44 B. pintoi; lectotype Achatina fragilis Smith, 1899 BMNH no. 97.12.31.9. 45, 46. _B. pintoi; holotype, Callistoplepa thielei,

Bequaert & Clench, 1934c ZMB no. 53177. 47, 48 B. pintoi; BMNH no. 1885.5.25.47. 49, 50 B. pintoi; BMNH MacAndrew (1563).

51 B. pintoi; BMNH no. 1907.7.25.3. 52,53 B. fraterculus (Dupuis & Putzeys, 1900); lectotype Ganomidos fraterculus MRAC no. 5140.

54,55 B. graueri (Thiele, 1911); lectotype Achatina graueri ZMB no. 101935. 56,57 8B. graueri; lectotype Callistoplepa babaulti,

Germain, 1936 MNHN. Bar scale = ~ 10 mm.

NEW SUBFAMILY AND GENUS ACHATINIDAE

evert together as a unit, forming a quite different intromittent

organ. The very shell characters that seemed to link these five

species with Callistoplepa now are seen conchologically to

distinguish the more primitive genus Bequaertina from Acha-

tina S.S.

On the basis of somewhat overlapping shell characters,

available distributional records, and the limited anatomical

evidence, the species of Bequaertina break into two groups:

(1) the malleate, mammillate B. fraterculus (Dupuis &

Putzeys, 1900) and B. graueri, and (2) the cancellate B.

pellucida, B. marteli and B. pintoi. B. pellucida of southeast

Zaire, close to what is believed to be its ancestral home, is

plesiomorphic within the group and stands between an ances-

tral achatinid stock of the Zaire Basin and Achatina s.s.,

which today is largely restricted to that basin. A branch of the

ancestral stock moved north and northeast to give rise to the

apomorphic B. fraterculus and B. graueri. A more conserva-

tive second branch moved east to give rise to B. pellucida and

B. marteli. This second branch continued further east and

then into a strong north-south axis to give rise to the closely

related B. pintoi. The known distribution of the genus (Fig.

16) embraces a vast area of the Rift Valley —Lake Region and

the Lualaba branch of the Zaire River in central, eastern and

southeastern Africa. Greater field collecting will probably

extend the limited distributions of B. fraterculus and B.

marteli.

Pilsbry & Cockerell (1933), on the erroneous assumption

that Achatina graueri ‘represented an intrusion of a South

African type into the Central African region’, initially

decided to place it in the genus Cochlitoma. They softened

their stand on the advice of M. Connolly and designated it

‘Achatina (Cochlitoma) graueri’. However, this species can-

not possibly be considered congeneric with Cochlitoma zebra

(Bruguiére, 1789), which Pilsbry (1904:xiii, 78) selected as

the type species of the genus Cochlitoma, because Mead

(1992) shows Achatina zebra anatomically belongs to subge-

nus Tholachatina of Archachatina. Since the present group of

five species is not congeneric with the species in either

Callistoplepa or Achatina s.s., and since this group also is not

congeneric with Achatina hortensiae Morelet, 1866, which

Pilsbry (1904:21) selected as the ‘type’ of Serpaea, there is no

other available genus-group name. Ganomidos cannot be

considered because it is a junior subjective synonym of

Callistoplepa. For these reasons, the generic name Bequaer-

tina is proposed. It is named in honour of the late Dr Joseph

C. Bequaert, Agassiz Professor of Zoology at Harvard Uni-

versity, who will remain the classical authority not only in the

Achatinidae, but also in several families of insects and

arachnids that he mastered in his long lifetime. Because B.

graueri is the largest and most conspicuous of the five species,

and because it has departed farthest from what is believed to

be the ancestral stock, it is here selected as type species of the

genus.

Early in the present study, it became obvious that this

group of five species anatomically was not congeneric with

Callistoplepa. This information was shared with colleagues

who considerately referred to this new genus in general terms

(van Bruggen, 1978:912, 921, 1988:10; van Bruggen &

Meredith, 1984:161). Also, the present author made refer-

ence to this new genus in an earlier manuscript as ‘Callis-

toplepa s.1.’ (Mead, 1992).

5 Shell large (61/4 whorls =

A.R. MEAD

Key to Species

1 __ Last whorl distinctly granulate above the periphery, or at least

in a limited subsutural zone; growth wrinkles conspicuous to

GOMUMANE Ts ae. ebisn cence ieqeelsneaeaai+)sacieae nen. -0 9p eee eee eRe Renee p2

— Last whorl faintly granulate, malleate, lirate or smooth except

for modest irregular growth wrinkles .................ec0eceeeeeees 4

2 Apex broadly obtuse; 6 whorls = > S50 mm; yellowish, ochra-

ceous or olivaceous; gross granulate sculpture; first nepionic

whorl 2-3 mm in diameter; second whorl expanding broadly;

sculpture of second whorl coarse and either distinctly granular

or depressed and poorly defined; transverse measurement at

junction of third and fourth whorls is 21/4 mm; outer lip

increasingly arcuate basally; growth wrinkles bold or moder-

ately HEAVY... cshescdsssicscecendoses-eeecteas cee ne cess eee eRe ee ROME 3

— Apex subacute to narrowly obtuse; 6 whorls = ~ 40 mm, 7

whorls = 53-60 mm; translucent dull fulvous to dull olivaceous-

brown; moderately coarse to fine granulate sculpture; first

nepionic whorl 2 mm in diameter; second whorl tends to be

slightly constricted, expanding limitedly; sculpture of second

whorl finely engraved, delicate; transverse measurement at

junction of third and fourth whorls is 2—2'/2 mm; outer lip evenly

arcuate; growth wrinkles thin, of modest calibre. Southeast

Zaire, northeast Zambia and west central Tanzania . pellucida

3. Last whorl large, rarely strikingly so; ground colour intense

olivaceous-yellow to subdued olivaceous; prominent closely

aligned somewhat irregular costate transverse ridges embrace

the gross elongate granules with bold vertical emphasis, domi-

nating the spiral lines; strongly contrasting zigzag castaneous

flammules usually present, pale unicolorous forms uncommon;

first nepionic whorl 2'/-3 mm in diameter; sculpture of second

whorl coarse, granular, elevated, tightly packed; transverse

measurement at junction of third and fourth whorls is 3-4 mm;

third whorl deeply and grossly granulate. Middle west and east

shores of Lake Tanganyika, Zaire and Tanzania ........ marteli

-— Last whorl large, often very large to ventricose; ground colour

dark olivaceous to pale olivaceous yellow; coarse granulate

sculpture above periphery, reduced or absent below periphery

(varies within a single whorl); transverse ridges moderate,

slender, fairly uniform, in balance with the spiral lines, con-

spicuous below periphery but obscured by granulate sculpture

above periphery; usually unicolorous, but narrow fairly straight

light castaneous stripes may be present; first nepionic whorl

2-2'/2 mm in diameter; sculpture of second whorl coarse, but

superficial, vaguely and irregularly impressed, patchy, poorly

defined, often worn smooth; transverse measurement at junc-

tion of third and fourth whorls is 2'/2-3 mm; third whorl

delicately to moderately granulate. East Africa, almost reaching

the Limpopo River in the south (4-20° S, 27-39° E) ..... pintoi

4 Apex of shell obtuse and noticeably mammillate; shell conspicu-

ously to obscurely malleate; opaque or dark and translucent,

uniformly or somewhat variably brown or yellow-brown, band-

ing limited and irregular; coarse growth wrinkles or extremely

fine lirae dominate the sculpture, 6 whorls = > 43 mm ...... >

— Apex of shell subacute to narrowly obtuse, somewhat elevated

but not mammillate; shell not malleate; translucent dull fulvous

to dull olivaceous-brown, usually with moderately broad casta-

neous flames and stripes irregularly distributed, but may be pale

unicolorous; very fine granulate-cancellate sculpture dominates;

6 whorls = < 43 mm. Southeast Zaire, northeast Zambia and

west central Wanzamiaeeceesea- sees see see eeeeeeee eee eee pellucida

60-80 mm), thin but substantial,

essentially opaque; usually conspicuously malleate; not cari-

nate; growth wrinkles prominent, rather regular; lirae of fifth

whorl distinctly transacted by spiral striae; unicolorous or

NEW SUBFAMILY AND GENUS ACHATINIDAE

Table 5 Locality records — Bequaertina. Numbers in the first

column correspond to the locality numbers in Figure 16. Sources

of specimen information are shown in the last column.

1. Lake Tanganyika, 1800-2000m 4° 30’ S, 29°00'E NHMW*

2. Kiambi 7° 20'S, 28°01’ E Dautz. &

Germ.,

1914

3. Sampwe (non ‘Sangue’) 9° 20°S, 27°26 E - SDautznd&

Germ.,

1914

4. Ibahi, Ugogo (=Ougogo) Riv. 5° 04’ S, 34°04" EE Ancey,

1902

5. Mbwe (=Mbwego) > 21'S, 38 598 8 Ancey;

1902

6. Mamboya (=Mamboa) 6° 16'S, 37°06’ E BMNH

7. Morogoro 6° 50’ S, 37° 45'E BMNH

8. Ngerengere, Oukani, Kingoni 7° 03'S, 38°31'E _Bourg.,

1889

9. Ufipa (=Sumbawanga) 8° 00’ S, 31°30’ E Ancey,

1902

10. Rukwa Lk. 8° 00’ S, 32°25’E BMNH*

11. Mbaya, 1700m 8° 45' S, 33°27'E BMNH,

LNK

12. Utengule 8° 54’ S, 33°20’ E BMNH,

MCZ,

ZMB,

SMF

13. Misuku Hills, Mughoma, 9° 40’ S, 33°33’E RMNH‘

1500m

14. Deep Bay (=Chilumba, 10° 27' S, 34° 16’ E BMNH

=Hengwa)

15. Nyika Plateau, 6000-7000 ft. 10° 48’ S, 33° 48’ E BMNH,

IRSN,

MCZ et

al.

16. Nkota-Kota 12° 55' S, 34° 18' EE BMNH

17. Nchisi (=Ntchisi) Mt. 13° 20’ S, 34° 05' E HM"

18. Chinyama 13° 43’ S, 33° 43’ E HM

19. Zomba, Shirwa Lk., Mpita 15723"'S;,35° 23’ E BMNH,

IRSN,

RMNH

20. Chiradzulu Mt., Lisau 15° 41’ S, 35°09’ E HM"

21. Nyambadwe Hill 1S248/'Sg5- lo) Ee ING?

22. Soche Mt. [S251 'S; 35° 01-8 ING"

23. Cheri Bridge, Upper Lauangwal3° 35’ S, 31° 30’ E MCZ

24. Broken Hill (=Kabwe) 14° 27' S, 28°27'E NMW

25. Kafue Riv., Mumbwa 15° 56’ S, 28°55’ E Beq. &

Cl., 1934¢

26. Pemba 16° 40' S, 27°25’ E SAM

27. Mazoe Valley 162 32° 'S5 33225" NMW

28. Salisbury 17° 50’ S, 31°03’ E NM

29. Vumba, Zonwi Bridge, 2500 ft. 19° 07’ S, 33°05’ E NM‘

30. Bulawayo 20° 09" S,.28°.35E.RMS,

SAM

31. Chirinda, Selinda Mt., 4000 ft. 20° 26’ S, 32°42’ E BMNH,

MCZ

32. Macequece, Vilade Manica 18°56’ S, 32°53’ E BMNH,

NMW,

SAM

33. Nsendwe, Maniema 2951-S;252 S60 ES BMNH,

MRAC

34. Uvira 3° 24’ S, 29°08'E ZMB

35. Mpala (=Pala) nS 29> 31 BS IRSN

36. Mweru (=Moero) Lk. 9° 00’ S, 28° 45'E BMNH,

IRSN

37. Dilolo 10° 42’ S, 22°20’ E SMF

38. Rumonge 3°11’ S, 29°08’ E MRAC

39. Kapuri (=Piani Kapuri) 3° 34’ S, 26°53’ E BMNH,

IRSN,

MRAC et

al.

40. Luaye 4° 42'S, 27° 23’ E MRAC

variably transversely striated with yellow-brown to dark brown;

nepionic whorls densely granulate. Lake Kivu district of Zaire,

Rwanda; Wie an apy mss f.8 6c gamete lostes n satanianco pia aetna graueri

— Shell small (6'/s whorls = ~ 50 mm), extremely thin, fragile,

translucent; malleations very shallow, often sparse; subcarinate

at periphery, producing a bend in the arc of fine prosocline

lirae; lirae of fifth whorl not transacted; ground colour dull dark

brown-olive with irregular castaneous brush marks that are

closely highlighted adaperturally with buff; nepionic whorls

faintly granulate. Lualaba River, Zaire ............... fraterculus

Bequaertina pellucida (Putzeys, 1898)

Figs. 31-36

Ganomidos pellucidus

Putzeys, 1898:84, text fig. 20, 21.

Callistoplepa pellucida

Pilsbry, 1905:128, pl. 43, fig. 3, 4; Germain, 1909:90;

Pilsbry, 1919:81; Bequaert & Clench, 1934c:114; Haas,

1936:13.

Serpaea foai

Germain, 1905:255; 1908:631.

Achatina foai

Verdcourt, 1966:111; 1983:219.

Callistopepla pellucida

Oliver, 1983:9.

SHELL. Shell ovate-achatiniform to elongate-ovate,

extremely thin, very fragile, translucent. Whorls 6-7, rarely

7/2; a conspicuous demarcation at or near the end of the third

whorl sets off the nepionic whorls. Spire conic, with a

narrowly obtuse apex that is slightly elevated; occasionally

the second nepionic whorl is somewhat constricted, produc-

ing a submammillate profile. Whorls slightly convex, expand-

ing and descending proportionately. Sutures moderately deep

Table 5 continued

41. Kabambare 4° 42'S, 27° 43'E ZMB

42. Lukuga Riv. mouth 5250! 9y 297 12) Be INICZ

43. Gandajika 6: 45°S; 2375/7" E MRAC

44. Pweto 87 2608928799 4E) 0 FIRSN

45. Kamina 8° 44’ S, 25°00'E MRAC

46. Abercorn (=Mbala) 8250! S312) EM GZ,

47. Kungwe, Sitete (=Nkungwe) 6° 07'S, 29°48'E Verdcourt,

1966

48. Beni 0° 30’ N, 29°28’ E _IRSN,

MRAC

49. Kitembo 2793483272375 7 MINN

50. Lobengera Mission 2031 S9297250 Es AMRAG

51. Ibanda 0° 08’ S, 30°30’ E MRAC

52. Loashi Valley 1°14’ S, 28° 45'E MRAC

53. Burungu, Ruasa 1S20"S,29". 02) ANSE

54. Nyabukere 1° 29’ S, 28° 33’ E MRAC

55. Kirotche, 1250 m 1°37’ S,29°02"E MRAC

56. Lwiro Riv. 2° 00' S, 28°52’ E AMNH,

FMNH

57. Idjwi (=Kwidschwi, Kwidjwi) 2° 09'S, 29°04'E ZMB,

MRAC,

UMMZ et

al.

58. Katana 2713S, 28° 50°E “MRAC

59. Tshibinda 2° 20’ S, 28° 45' E ANSP,

MRAC

60. Bukavu 2304S; 28252415 SMINEIN:

NMB

t = specimens dissected in the present study.

and irregular. Last whorl large, 80% of shell length; range for

5-7/2 whorls, 76-84% (n = 52). Aperture oval, faint milky

wash within. Columella concolorous, slender, moderately

long straight or slightly arcuate and rectangularly to very

obliquely truncate. Outer lip extremely thin, evenly arcuate,

receding at base in profile. Parietal callus scarcely detectable.

The first two to three whorls are light horn colour. Pale,

obscure, castaneous streaks begin to appear in the third or

fourth whorl; these characteristically are broader at the

suture below, becoming increasingly darker, larger and more

irregular on the last whorl. These streaks may be variously

vertical, diagonal, angulate, flammulate, interrupted, or

reduced to spots and blotches. Ground colour is dull buff to

dull olivaceous-fulvous. Of 56 specimens checked precisely

for colour, 63% have a definite pattern, 7% are nearly

unicolorous, and 30% are unicolorous. There was no correla-

tion between colour pattern and locality.

The first whorl is essentially without sculpture. Minute,

faintly engraved crescentic granulations usually appear early

in the second whorl; these are formed by nearly equidistant

spiral lines and irregular, scalloped transverse lines. The

latter become straighter and compressed in the third whorl,

producing narrow elongate granulations and irregularly

appearing prosocline growth wrinkles that are crenulate at

the suture. The sculpture becomes more disperse in the

fourth and fifth whorls, producing a dominant, fairly uni-

form, subquadrate, often welt-like, cancellate-granulate

sculpture, which usually fades quickly at the periphery. In the

sixth to seventh whorls, this sculpture becomes more and

more subdued and diffuse until the increasingly prominent,

yet modest, growth wrinkles dominate both above and below

the periphery. In the largest specimens of seven whorls the

cancellate-granulate sculpture may feebly or strongly return

both above and below the periphery. The dull, extremely

thin, tenaceous outer periostracal layer wears off in very

limited areas, highlighting the sculpture with the exposed

glossy inner periostracal layer.

SOFT ANATOMY. No known alcohol preserved specimens.

TYPE MATERIAL. As nearly as can be determined, Putzeys

had 14 syntypes of his Ganomidos pellucidus, for which he

gave a range of shell dimensions (1898). The specimens were

collected by P. Dupuis. Putzeys retained a select series of 7

syntypes in his own collection (MRAC no.5132-5138). He

did not designate types, but selected the largest specimen

(no.5132) for an abaperatural view and a small specimen with

slender flames (no.5133) for an aperatural view in his line

drawing illustrations. Regrettably, the larger specimen had

been rather badly damaged and mended in nature, and the

smaller specimen was excessively small. The second largest

syntype in his series (no.5136) is a unicolorous specimen that

is representative of only about a quarter of the known

specimens (Figs. 33, 34). Hence, the flamed, third largest

syntype in his series (no.5135) is here selected the lectotype

(Figs. 31, 32; Table 6), with the other syntypes becoming

paralectotypes (BMNH 1no.1904.5.18.68, IRSN 5, MRAC 6,

NHMW 1).

On the basis of two specimens collected by Edouard Foa

during his 1897-98 expedition to the Lake District of Africa,

Germain described (1905) and figured (1908) the junior

subjective synonym Serpaea foai from ‘Tanganika est’, later

corrected to ‘les bords du Lac Tanganyika’. Bequaert (1950)

placed Serpaea in the synonymy of Achatina, but apparently

A.R. MEAD

Table 6 B. pellucida — Representative shells measurements.

Greatest Aperture Last %

Whorls Length Width Length Width whorlLW/L % W/L

7 60.4 31.4 3277 1955" 45,9" 76 52 Mweru

(BMNH)

56 Kamina

(MRAC)

581.196

7 S4e5 2910) S05") L720)) 42-5578 53 Mweru

(BMNH)

1907.11.

115

7 5353" 3012 S102 1728) AZ Sao) 57 Mpala

(IRSN)

61/2 48.0 28.0 29:7 15:8, 38-0) 79: 58 Piani

Kapuri

(MRAC)

5132 PLec

G.p.

61h 46.9 29.6 294 116:5" 372) 80 63 Tanganyika

(MNHN)

Lect S.

foai *

61/4 45.8 28.0 28.0 16.7 37.0 82 61 Piani

Kapuri

(IRSN)

PLec G.p.

61/2 45.0 28.2 26:8 155% 355099 63 Piani

Kapuri

(MRAC)

5136 PLec

G.p. *

6's 43.2 24.6 Pisyoy IERIE MS See)! 7s" 57 Piani

Kapuri

(MRAC)

5135 Lect

Gip..

61/4 40.4 22.6 23:0) 13:6 3317s 56 Piani

Kapuri

(MRAC)

5133 PLec

G.p.

6 39.0 23.3 24.4 14.1 32.0 82 60 Tanganyika

(MNHN)

PLec S:

foai

5% 3525) 20/0 20.9 12.0 28.4 80 56) Blec Gzp:

(BMNH)

1904.5.

18.68

Th 58.6 32.8 36.0 16.6 46.0 79

Total specimens examined: 62. Sources: BMNH, IRSN, MCZ, MNHN,

MRAC, NMW, USNM, ZMB.

overlooked Germain’s species. Only Verdcourt (1966) has

acknowledged the existence of this species, and then only as

an East African species unknown to him. A study of the two

syntypes in Paris (MNHN) confirmed the fact that they are

indeed Putzeys’ species. His larger, sharply photographed

‘seul adulte’ specimen (Figs. 35, 36) is here selected as the

lectotype of Germain’s Serpaea foai (Table 6).

Deshayes (1824-37, 1864) described and illustrated a small

fossil snail Agathina pellucida (=Achatina pellucida) from the

Paris basin. Lamarck (1838:313) also refers to this species.

This very acuminate, slender specimen is possibly a subulinid.

It does not enter into homonymy with Putzeys’ G. pellucidus

because the latter was never included in the genus Achatina.

NEW SUBFAMILY AND GENUS ACHATINIDAE

TYPE LOCALITY. Forest of Piani Kapuri, Maniema

(=Manyema), Zaire 3° 34’ S, 26° 53’ E.

DISTRIBUTION. Next to B. pintoi, this is the most wide

spread species in the genus (Fig. 16). The known specific

localities delineate essentially the southeastern quarter of

Zaire, with Nsendwe, near Kindu-Port-Empain, Maniema

region in the northwest; Uvira, Kivu region in the northeast;

Mpala, Tanganyika region in the east; Lake Mwero

(=Moero), Kantanga region in the southeast; and Dilolo,

Lualaba region in the southwest. The only records outside

Zaire are (1) in Abercorn (= Mbala) at the southern tip of

Lake Tanganyika, Zambia, and (2) on the Tanzanian east

shores of this lake, based on Germain’s synonym Serpaea foai

(1905, 1908). Meredith (1983b) and N. Gray (correspon-

dence) failed to find it during extensive collecting in Malawi.

The largest and finest specimens extant were collected in the

Lake Mwero region (BMNH) and Kamina (MRAC).

REMARKS. This plesiomorphic wide spread species is most

closely related to B. marteli. Specimens have been found in

mixed lots along with B. marteli and Achatina craveni. The

juvenile specimens of all three species are easily confused.

Further, the full grown specimens are quite variable in shape,

colour, sculpture and pattern, with the not uncommon atypi-

cal forms of each species contributing to the difficulty of

identification. The young specimen that Grauer collected in

the virgin forest SO km east of Kasongo, Zaire, and identified

as Achatina fulminatrix von Martens, 1895 by Thiele

(1911:205) was examined in Berlin (ZMB) and found to be B.

pellucida. Extensive series of this species are in Bruxelles

(IRSN) and Tervuren (MRAC).

Bequaertina marteli (Dautzenberg, 1901)

| Figs. 37-40

Achatina marteli

Dautzenberg, 1901:3.

- Ganomidos marteli

Dautzenberg, 1901, pl. 1, fig. 1.

Achatina marteli pallescens

Dautzenberg, 1901:3.

Ganomidos marteli pallescens

Dautzenberg, 1901, pl. 1, fig. 2.

Callistoplepa marteli

Pilsbry, 1905:129, pl. 47, fig. 21 (ex Dautzenberg); Ger-

main, 1909:90; Pilsbry, 1919:81; Bequaert & Clench,

1934c:114.

Callistoplepa marteli var. pallescens

Pilsbry, 1905:129, pl. 47, fig. 22 (ex Dautzenberg);

Bequaert & Clench, 1934c:114.

“Achatina sp. near tavaresiana’

Verdcourt, 1966:106, fig. 12; 1988:219.

Callistopepla marteli

Germain, 1936:151; Oliver, 1983:9.

SHELL. Shell ovate-achatiniform, opaque, thin but not frag-

| ile. Whorls 6-6/4, rarely 61/2. Spire moderately broad, conic;

apex obtuse; only one out of 69 specimens examined had a

_mammillate apex. Upper whorls only slightly convex,

descending proportionately but expanding somewhat more

rapidly. Sutures fine and regular in nepionic whorls, shallow

to moderately deep and irregular in the following whorls.

_ Last whorl large and more convex, 80% of shell length; range

for 41/262 whorls, 77-84% (n = 69). Aperture inverted

auriform to ovate-elongate; pale blue-white within; surface

pattern and flames show through. Columella straight or

weakly arcuate, somewhat slender, concolorous but with a

thin calcareous film; usually moderately obliquely truncated.

Outer lip thin, extending basally only a slight way below the

truncation; its arc is characteristically greatest below midway

in the mature specimens. Parietal callus thin but apparent

even in the smaller specimens.

The nepionic whorls (first 2!/2) are unicolorous pale buff-

white. This changes imperceptibly to a uniform dull ground

colour that varies in specimens from a rather intense oliva-

ceous yellow to subdued olivaceous. In most specimens,

faint, very diffuse light castaneous blotches appear in the

fourth whorl. At first these are vertical, evenly spaced and

broader at their base; but they soon become fragmented

apically, darker, and strikingly distorted into diagonal even

spiral, irregular streaks, bands and flames that are approxi-

mately as wide as the ground colour space between them. In

the present study of 69 specimens, 72% are flammate, 13%

are vaguely flammate but only on the last whorl, and 15% are

without flames, i.e. ‘pallescent’. In some of the latter, e.g. the

lectotype of Achatina marteli pallescens, lines of arrested

growth are highlighted with thin bands of dark brown.

A delicate beaded or slightly semilunar sculpture starts in

the second quarter of the first whorl and quickly assumes in

the early second whorl the diagnostic sculpture of strikingly

coarse, elevated, round or crescentic, discreet but tightly

packed beads that are neatly aligned in 5-7 spiral rows. This

pattern persists almost uniformly throughout the second

whorl. In the mid-third whorl, the transverse rows become

greatly compressed, producing growth wrinkles and convert-

ing the beads into transverse welts 2-3 times as long as wide.

This doubtless marks the first postemergent growth. Adaper-

tural to this, the growth wrinkles become prosocline, the

sculpture gradually becomes less compressed, the spiral striae

become more numerous and deeper, and the individual welts

become larger, more variable in size, more rectangular, and

often cleft. The remarkably evenly and closely spaced coarse

growth wrinkles embrace and intensify the prosocline rows of

welts, producing the characteristic prominent ribbed sculp-

ture of this species. Apically, the ribs may bifurcate and form

crenulations. Below the periphery, the welts rather abruptly

reduce to one-quarter their calibre, or are absent, leaving

prominently the growth wrinkles. An extremely fine decus-

sate micromesh of the periostracum appears on the last whorl

of some specimens. It is more noticeable on the shiny inner

layer of the periostracum where the latter is exposed through

wear or injury. It is apparent that the micromesh is formed at

the time that the inner periostracal layer is laid down and that

it is largely obscured by the preformed, smoother outer

periostracal layer. It is likely that the micromesh assists

structurally in bonding the two periostracal layers.

SOFT ANATOMY. No known alcohol preserved specimens.

TYPE MATERIAL. In his description of this species and its

synonymous unicolorous ‘variety pallescens’, Dautzenberg

(1901) announced that he was dedicating them to Colonel

Martel and that specimens had been collected by R.P.

Guillemé ‘en nombreux exemplaires’ in the region of Lake

Tanganyika. He did not specifically designate types and

paratypes, although he selected a fine flamed specimen and

an equally fine unicolorous specimen that were photo-

graphed, both in apertural view only, as representative of the

two proposed taxa. These are in the type collection in

Bruxelles (IRSN) and are here selected as lectotypes of

Dautzenberg’s Achatina marteli and A.m. pallescens, respec-

tively (Figs. 37-40; Table 7). As he pointed out in a footnote

in the original descriptions, the pronounced flame pattern of

his figure 1 unfortunately did not reproduce well. Pilsbry’s

copies (1905) therefore reflected this deficiency. In this

species, neither the lack of colour pattern nor the greater

degree of ventricosity is taxonomically valid for establishing a

trinomen.

In the IRSN collection there are several mixed lots totalling

48 mostly juvenile, damaged or weathered specimens. All

these specimens were very carefully examined in the present

study and were found to be a mixture of the flamed and

unicolorous forms of this species and, in addition, juveniles of

Bequaertina pellucida and Achatina craveni. These cannot

reasonably be considered to have been a part of Dautzen-

berg’s type series. Dautzenberg, however, did distribute his

specimens widely. Those bearing the type locality and R.P.

Guillemé as the collector are here selected as paralectotypes.

The known distribution of these flamed/unicolorous speci-

mens are BMNH 1/0, NMW 1/1, IRSN 8/5, MCZ 2/1, MRAC

7/1, NMB 2/1, MNHN 6/1, NHMW 1/0.

Table 7 8B. marteli— Representative shells measurements.

Greatest Aperture Last %

Whorls Length Width Length Width whorlLW/L % W/L

64/2 68.3 34.0 42.3 20.8 55.4 81 50 Mpala

(IRSN)

IML

61/4 67.2 38.6 BOI | 2222) D4-4. 8h 57 Mpala

ao)

6 63.7 3372 S7a* 19SIGE Sl S81 5

61/4 62.9 82.3 36.7 18.2 49.8 79 5

<x:

no]

eas

6 60.8 34.8 38.0 19.4 51.3 84 57 Mpala

5% 58.4 Bias 37.3. 18.5 49.0 84 55 Mpala

5129 PLec

654.032.233.817.043.78160

54.0 B22) sB8 IZA BP Sil 60 Mpala

(MNHN)

JAL@E

5¥% 48.4 26.9 S27 1S. OM SOrsF 82 55 Mpala

(BMNH)

Pec

1937.12.

30.1934

52 48.4 27.9 32.0 16.7 40.5 84 58 Mpala

(MRAC)

5125 PLec

6 48.4 M33 SO: ISS. 3394 3? 56 Mpala

(MRAC)

5131 PLec

Total specimens examined: 69. Sources: BMNH, IRSN, MCZ, MNHN,

MRAC, NHMW, NMB, NMW.

A.R. MEAD

TYPE LOCALITY. Mpala (= Pala) 6° 45’ S, 29° 31’ E, region of

Lake Tanganyika, Zaire (cf van Burggen, 1988:9).

DISTRIBUTION. All specimens examined are from the type

locality. Verdcourt kindly sent the author a photograph of a

specimen, which earlier had been identified as Achatina

tavaresiana (Pain & Verdcourt, 1962; Verdcourt, 1966). This

specimen is clearly B. marteli and establishes this species in

Tanzania on the eastern shore of Lake Tanganyika in the

Mahari Peninsula at Nkungwe (= Kungwe) 6° 07’ S, 29° 48’

E. It was collected in ‘litter in thick scrub at head of stream,

altitude 4500 ft’. With the many locality records known for B.

pellucida, including Mpala, it is strange that the distribution

of the present species is, to date, so contrastingly limited.

REMARKS. This is the least fragile and the most boldly

sculptured species in the genus. Phylogenetically, it appears

to stand between B. pellucida and B. pintoi. Mixed lots of B.

marteli, B. pellucida, and Achatina craveni suggest that these

species are sympatric. The juveniles in particular are confus-

able. By far the largest series of this species is to be found in

Bruxelles (IRSN).

Bequaertina pintoi (Bourguignat, 1889)

Figs. 41-51

Serpaea pintoi

Bourguignat, 1889:86, pl. 4. fig. 4.

Achatina fragilis

Smith, 1899:591, pl. 35, figs. 3,4 (non Achatina fragilis

Deshayes, 1864); Ancey, 1902:278, text fig. 6; Pilsbry,

1904:63, pl. 9, figs. 25, 26 (ex Smith); Dautzenberg &

Germain, 1914:26.

“Achatina.... sp. nov?’

Ancey, 1902:277, text fig. 4.

Achatina pintoi

Pilsbry, 1904:63, pl. 41, fig. 8 (ex Bourguignat); Bequaert,

1950:11; Verdcourt, 1966:111, 1983:219.

Achatina nyikaensis

Pilsbry, 1909:113, 1919:79; Connolly, 1925:168, 1939:321;

Germain, 1935:9; van Bruggen, 1965:81, 1988:10; van

Bruggen & Meredith, 1984:161.

Callistoplepa nyikaensis

Bequaert & Clench, 1934c:115, 116.

Callistoplepa thielei

Bequaert & Clench, 1934c:115, pl. 2, figs. 8-10, 12.

Callistopepla nyikaensis

Verdcourt, 1966:111, 1983:219; Meredith, 1983a:29, fig.

10, 1983b:247.

SHELL. Shell thin, fragile, highly variable in shape, usually

elongate-ovate or ovate-subsuccineiform, but may be ovate,

globose-ovate or slender conic-ovate. Whorls 6-6'/2, rarely

slightly larger. Apex obtuse; vaguely mammillate in some

specimens. Spire usually inscribes a short broad based tri-

angle that appears to be nearly equilateral, 27-36% of shell

length. Less commonly, the spire is more produced, with the

sides of the connate triangle appearing longer than the width

of the base, 37-43% of shell length; such specimens may or

may not have a more slender last whorl. Sutures fine, distinct,

deeply impressed, quite regular, but may be faintly crenulate

in the last part of the sixth whorl. Whorls slightly, moder-

ately, or distinctly convex, usually expanding rapidly to form

a large last whorl, 84% of shell length; range for 31/2674

whorls, 78-87% (n = 87). Aperture ovate-acuminate; charac-

NEW SUBFAMILY AND GENUS ACHATINIDAE

teristically widest slightly below middle; translucent pale

milky opalescent within; fine, closely aligned internal riblets

mirror the external sculpture. Columella concolorous or

white; long, slender, feebly arcuate or nearly straight; trun-

cated obliquely or abruptly. Outer lip extremely thin, fragile,

somewhat receding, arcuately skewed and evenly rounded

toward the base. Parietal callus not apparent in young or

fresh specimens; thinly calcareous white in others.

Nearly 90% of the specimens examined in the present

study (77/87) were essentially unicolorous in a colour gradient

from pale olivaceous yellow to olivaceous brown to deep

olivaceous green. In individual specimens, the colour tends to

be quite uniform, except for darker bands where there was

cessation of growth. Because of their very thin two periostra-

cal layers, the apical whorls, which are pale straw colour,

soon become calcareous white with wear and exposure. Ten

of the specimens examined had on their lower whorls, very

faint, narrow light castaneous irregular, sometimes inter-

rupted, transverse stripes that were one-half to one-third the

width of the ground colour between them. In some speci-

mens, only a fraction of a whorl was involved.

Extremely fine spiral engraved lines appear at the end of

the first whorl. Transverse lines appear in the second whorl,

giving rise to depressed beads that become more conspicuous

and more abundant until near the middle of the third whorl.

At that point, which marks the end of the nepionic whorls,

the beads become compressed into very narrow transverse

ridges. In the fourth whorl, this compression is relieved and a

fairly even granulose-cancellate sculpture emerges. In the

fifth and sixth whorls, the granulae swell to become welts that

occasionally anastomose along the transverse growth

wrinkles, with the shallow spiral lines remaining strongly in

evidence (cf Bequaert & Clench 1934c, fig. 12). Usually, this

sculpture diminishes rapidly in caliber at the periphery, with

essentially only the growth wrinkles continuing into the

otherwise smooth surface. In other specimens, the sculpture

may continue strongly below the periphery, but at a reduced

calibre. In still others, subdued patches of this sculpture

| appear irregularly below the periphery, and perhaps contrast-

ingly so with resumed growth after diapause. Particularly in

the parietal area, the dull, tenaceous microscopically granular

outer periostracal layer may wear off, exposing the smooth,

shiny inner periostracal layer. Only rarely has a periostracal

decussate micromesh been observed in this species, and then

only spottedly below the periphery in the inner periostracal

_ layer of the sixth whorl, e. g. in the holotype of Callistoplepa

thielei Bequaert & Clench, 1934c.

SOFT ANATOMY. Alcohol preserved specimens available

29/dissected 17. Tanzania: BMNH 1/1, NHMW (no.47996)

1/1. Malawi: CMNH 13/2, HM 5/5 (all now at RMNH), NG

| 4/3 (2 now at BMNH), RMNH 3/3. Zimbabwe: NM 2/2.

| Additional Malawi specimens are in collections HM and NG.

No others are known. Most specimens examined were well

extended from their shells.

In the individual specimen, the body colour varies from

unicolorous pale buff to dark grey. A black thin-lined,

coarsely reticulate pattern is characteristically present later-

| ally on the foot. Sole of foot is uniformly pale dusky, without

variation in texture. Usually a delineated or diffuse narrow

dark grey or black stripe, originating behind each ommato-

phore, passes posteriorly toward the mantle on either side of

the paler neck region. The mantle varies from unicolorous

dark or light grey to a grossly maculate pattern. The Chirinda

2if

Forest, Mount Selinda, Zimbabwe field notes of A.C. van

Bruggen in litt.) record the body colour varying from black to

pale grey marbled with black, and the longitudinal stripes

varying from white to greyish white.

The short, slender penis (P) and the slightly shorter penis

sheath (PS) seem diminutive compared to the relatively gross

structures of the basal female conduit. Of the 17 specimens

dissected, 11 had the apical-most P and the basal-most basal

vas deferens (BVD) projecting slightly above the rim of the

PS. In the other 6 specimens, the P was completely covered

by the PS, 2 of which were gravid, 2 were over-drowned with

consequent eversion of the genital atrium (GA), one was

immature and one was severely distorted because of

improper fixation. There was no positive correlation in this

latter group of 6 with such other possible influencing factors

as latitude, shell size, length of penial retractor, month of

collection or size of albumen gland. The specimen (NM)

depicted in Fig. 17 collected by A.C. & W.H. van Bruggen in

Zimbabwe (Vumba Circular Drive, Zonwi River Bridge) has

thus been selected as representative of the first group and

typical of this species.

In all specimens, the penial retractor (PR) inserts either on

the anterior diaphragm, on the body wall of the neck, or at

the forward junction of the diaphragm, mantle and body wall.

In well extended specimens, it can appear inordinately long.

Cutting and spreading the thin-walled PS reveals the slender

P and BVD within (Fig. 18). Fine muscle strands pass basally

from the PR to cover thinly the apical P. About half way

down the P, these strands attach to the smooth, shiny inner

surface of the PS and then proliferate into voluminous strands

that completely cover the BVD and infuse with the tissues of

the basal P and PS. At this level the tissue layers are not

distinct, but below the PS the male conduit continues as a

short penial atrium (PA) that joins the vaginal atrium (VA).

These fuse to form the genital atrium (GA). Strap-like

overlapping, glistening eversion muscle bands (EM) connect

the basal male and female conduits to the inner right body

wall. During precopulatory behaviour, contraction of these

muscles cause the GA, then the PA and VA to evert, to

protrude as a stimulatory organ, and subsequently to initiate

the extroversion of the intromittent organ. In some older

specimens, these bands can be so voluminous and so high on

the PS that they seriously obscure relationships. The thin-

walled BVD passes through the PS to emerge as a much

larger, thick-walled apical vas deferens (AVD), which basally

functions as an ejaculatory duct and as a support for the

thin-walled everted P. About midway apically, the AVD

gradually becomes thin-walled until at the AVD/FO junction,

it forms a glandular funnel-shaped chamber that internally is

crowded with extremely thin epithelial partitions. This may

function as a secondary seminal vesicle (SSV). There is no

pilaster or verge in this species.

In the basal female conduit, the vagina (V), free oviduct

(FO) and the spermathecal duct (SD) form, without distinct

delineations, an impressive muscular Y-shaped structure

(Fig. 18). Upon dissection, the broad lumen and the promi-

nent longitudinal plicae of the V are seen to be uninterrupt-

edly confluent with those of the SD. In contrast, the lumen of

the basal, muscular thick-walled FO is slender and it is

confluent with that of the V through a small recessed pore.

This arrangement ensures the passage of the intromittent

organ into the SD during copulation. The thick-walled FO

functions basally as an ovijector; apically its thin walls

accommodate the descending eggs. The clavate-elongate

spermatheca (S) (Fig. 17) frequently is apically attenuated

because thin fibers firmly attach it 4-5 mm _ below the

AVD/FO junction. That junction, in turn, is held tightly

together on the right internal body wall by fibres from the

transverse myoseptum. Any contraction or extension thus

provides maximum pull in this part of the body. As a result,

under certain conditions, the S apex and SD may become

inordinately elongate or nearly disappear into a huge saccu-

late S. Severe contraction of the viscera may even produce

the artifact of a tandem bilobed S. The only important

character of the S is its consistent position below the

AVD/FO junction. The FO, though shorter than the AVD, is

conspicuously long, nearly uniformly wide, and approxi-

mately twice as wide as the AVD. Although there are thin

facia binding together V, SD, S, FO and AVD, there is no

formation of a vaginal rententor per se.

Two specimens were gravid. One from Chinyama, Malawi

(HM) was collected in February 1983 (mid-rainy season) and

had 56 light yellow, moderately large eggs averaging

6.3 X 5.2mm. The other, from Lake Rukwa, Tanzania

(BMNH), was collected in 1938 (month not indicated) and

had 51 similar eggs measuring 7 X 5.5-6.5 mm. The eggs,

without discernible embryos, were closely embraced by uter-

ine tissue folds as depicted by Mead (1950, fig. 48) for

Achatina fulica Bowdich, 1822.

All specimens dissected showed a remarkably gross devel-

opment of a trimerous hermaphroditic duct system (Fig. 19).

Basally, there is a 10 mm characteristically deeply convoluted

portion just distal to the talon. Next, there is a 3 X 2 mm

discreet, compacted saccular portion that appears to be

glandular. And finally, there is a 6mm slender, weakly

convoluted portion that quickly forms a series of five larger,

tightly convoluted ductules leading to the five gonadal acini

buried in the right lobe of the digestive gland. The talon is

elongate, capitate and diminutive (1.5 x 0.5 mm). The sper-

moviduct is characteristic of the family.

The following anatomical characters distinguish this species

from B. graueri: basal genital fascia diaphanous; AVD about

twice the width of BVD and half the width of FO; P slender,

much longer than wide, normally projecting slightly above

PS; BVD slender, much longer than wide.

TYPE MATERIAL. For over a century, Bourguignat’s (1889)

Serpaea pintoi has been an engima to conchologists. The

principal contributing factor has been the artist’s rendering an

excessively bold, wide, broadly truncated columella. A sec-

ond factor is that the exceedingly thin, fragile shell of the only

know specimen unfortunately had become broken sometime

since it was drawn. The several pieces, including a figure ‘4’

label, had been placed in a separate vial. This specimen (Figs.

41, 42; Table 8) is in Paris (MNHN) and is here considered to

be the holotype by monotypy (Code Art. 73(a)(ii); 74(b). Its

conchological features and the type locality in eastern Tanza-

nia support the conviction that this is conspecific with Smith’s

junior subjective synonym Achatina fragilis (1899). Pilsbry

(1904:1, 21, 63) placed Serpaea in the synonymy of Achatina,

indicated (1909:113) that Smith’s name was a primary hom-

onym of Deshayes’ fossil species (1864), proposed the

replacement name Achatina nyikaensis, reproduced Smith’s

figures [fig. 26 is way too intensely coloured], and perspica-

ciously placed Bourguignat’s species and Smith’s species

seriatim under Achatina in his Manual.

Smith’s syntype lot of seven specimens from Nyika Plateau,

Malawi, unfortunately contains a single specimen of Achatina

A.R. MEAD

Table 8B. pintoi — Representative shells measurements.

Greatest Aperture Last %

Whorls Length Width Length Width whorlLW/L % W/L

61/2 74.6 42.8 AT6.. 26.1 62:5) 184 57 Nyika

Plateau

(BMNH)

Lect A.

fragilis*

6/4 74.4 42.0 AGS, 7 24:6) 1614982 56 Macequese

(SAM)

6 66.8 40.6 46.9 24.0 58.0 87 61 Nyam-

badwe

(NG)'

64 62.2 32.7 38:3). 1955 50.6781 52 Utengule

(ZMB)

Holo C.

thielei *

6 56.2 37.8 S80) 213) A785 67 Bulawayo

(RMS)

6 56.0 33.0 35.0) | 196) 4615) 83 59 Utengule

(BMNH)*

Mac

Andrew

(1563)

6 53.3 Die? 33:0) 5 16:7 “43:5, 82 51 Chirinda

(BMNH)*

1907.7.

25:3

6 50.1 32.6 33.0 19.8 42.4 85 65 Mamboya

(BMNH)*

1885.5.

25.47

6 49.7 33.6 31.8" 195" 48:25:87 68 Ngrengre

(MNHN)

Holo S.

pintoi*

55 Tanzania

(NHMW)'*

68 Pemba

(SAM)

66 Usagara

(BMNH)

61/4 49.3 D2 2823) S20 38:5 478

RY 49.3 SiE// 34:9" 19:3" 42-7, 87,

5% 43.7 29.0 2910 17S" 37-2. 83)

Total specimens examined: 92. Sources: BMNH, FMNH, HM, IRSN, LNK,

MCZ, MNHN, MRAC, NG, NHMW, NM, NMW, RMNH, SAM, SMF,

UMMZ, ZMB.

craveni Smith, 1881. Since Smith, as its author, was thor-

oughly familiar with this latter species, since he did not

designate the number of syntypes, and since he separately

discussed (p. 35) and figured (figs. 1-4) specimens of both

this species (BMNH no.97.12.31.1-7) and A. fragilis (BMNH

no.97.12.31.8-14) in his 1899 paper, it is here safely assumed

that the division between the two adjacent accessioned speci-

men lots was an inadvertent curatorial error, with no nomen-

clatural implications for the misplaced specimen. The

unicolorous, largest and _ finest specimen (BMNH

no.97.12.31.9) of the six syntypes (Smith’s fig. 3, Pilsbry’s fig.

25), here shown in Figs. 43, 44 bears a handwritten note,

‘Lectotype “3’’ A.C. van Bruggen, May 1974’. This selection

is here endorsed. Other known and examined paralectotypes

are single specimens in Berlin (ZMB no.101934) and Vienna

(NHMW/R).

Ancey (1902) described and illustrated a specimen which

he labelled ‘Achatina . . . sp. nov.’ from Ugogo (5° 4’ S, 34°

4' E). Ancey’s collection was distributed widely by Geret and

this specimen so far has not been located. However, the very

{

NEW SUBFAMILY AND GENUS ACHATINIDAE

large aperture, the slender arcuate columella, the transverse

stripes, the translucence of the shell showing the pattern

through the aperture, and the locality indicate that his

specimen is Bourguignat’s species.

Bequaert & Clench (1934c) found nine purchased (Her-

mann Rolle) specimens from Utengule, Tanzania in the

Berlin Museum (ZMB) that were marked as new and given a

preempted manuscript name, but were undescribed.

Although they felt the new taxon ‘might perhaps prove to be

a local race’ of Callistoplepa nyikaenisis, its more slender,

more tapered shape convinced them it should be established

as the new species Callistoplepa thielei. The largest specimen

previously had been broken, so they selected the second

largest specimen as the holotype (Figs. 45, 46). In addition to

the holotype, they figured two paratypes. This series of four

types (ZMB no.53177) are in Berlin (Kilias, 1992). Examina-

tion of these types in Berlin in 1989, showed evidence that the

paratypes had been broken apparently on their return to

Berlin because their fragile shells had been stuffed exces-

sively with cotton, leaving only the holotype undamaged. The

five other original specimens, 3 full grown and 2 juveniles,

are labeled paratypes (MCZ no.98686) in the Harvard collec-

tion. Three other broken specimens with the same data are in

Frankfurt (SMF); there is no evidence that these were seen

by Bequaert & Clench.

TYPE LOCALITY. Bourguignat (1889) states, ... provient

des environs de l’Ougerengere, vallée du Kyngani, dans

VOukani’. Inside the lip of the holotype, in Bourguignat’s

handwriting, is ‘Serpaea pintoi Ougerengere (Oukami). A

small label bears the inscription ‘M. Requin 1846 30’.

According to B. Verdcourt (in litt.), Ukami is a large

geographic district that surrounds and includes the Uluguru

Mountains in Tanzania. The Kingoni river and the Ngeren-

gere settlement and stream, however, place the type locality

close to 7° 03’ S, 38° 31’ E, 125 km west of Dar-es-Salaam (cf

Verdcourt, 1966:111).

DISTRIBUTION. This is by far the most widespread species in

the genus. The 32 recorded localities define a 1800 = 1200

km territory 4-20° S, 27-39° E that includes eastern Zaire,

eastern and western central Tanzania, nearly all of Malawi,

south central Zambia, southern and eastern Zimbabwe, and

far west central Mozambique (Fig. 16). The northern outpost

of this species was established in Zaire by R. Grauer in 1910,

when he found and preserved in alcohol a specimen in ‘the

primary forest behind bordering hills of the northwest shores

of Lake Tanganyika, 1800-2200 m’ (trans.) 4° 30’ S, 29° 00’ E.

It is noteworthy that along the west, there is an almost

straight N-S line of demarkation 27—28° E from eastern

central Zaire to south central Zambia and southwestern

Zimbabwe. Connolly, in his writings (1925, 1939) and on

some of his specimen labels, juxtaposes the geographic names

Macequece (a district in eastern central Mozambique) and

Lourencgo Marques (the major port now known as Maputo in

southern Mozambique). Specimens may have been shipped

from this port, but there is no convincing evidence that this

species has ever been collected in Mozambique south of the

Macequece district. The remarks of Germain (1935:4) seem

to clarify when he explains that Portuguese East Africa is

generally divided into two large regions separated by the

Zambeze River, ‘le Mozambique au Nord, le Lourenzo

Marques au Sud’. When adequate population studies can be

made in this widespread species, valid subspecies may

emerge.

REMARKS. This apomorphic species is most closely related to

B. marteli. Both Achatina craveni E.A. Smith, 1881 and B.

pintoi are highly variable and are often confused in collec-

tions, particularly where the field data are the same or the

individuals are small. Despite rather considerable overlap in

the extremes of the conchological characters of these two

species, the shell of A. craveni can be differentiated on the

basis of the following: nepionic whorls smooth; last whorl

equals only 70-75% of shell length (vs 80-90%); one to two

more whorls for the same length; apex more acute; columella

much shorter, broader, straighter and more squarely trun-

cated; finer and deeper granulate-cancellate sculpture; shell

usually much less fragile.

The wide distribution and the independently great variabil-

ity in shell characters, even within a single population, are

responsible for the long and confused synonymy of this

species. The soft anatomies of antipodal specimens, and

many between, support the conclusion that only one species

is involved.

Ecological notations with specimen data (HM, NG) indi-

cate that active specimens were found in Malawi lowland

evergreen forests, along the banks of earth roads, crawling on

leaf litter, and on the underside of banana leaves in the rain.

Specimens from Mbeya, Tanzania were found in luxuriant

herbaceous vegetation (LNK). The director of the Imperial

Institute of Entomology earlier reported that in the Nkota-

Kota district of Malawi there was a ‘very serious outbreak in

November 1937’, implying that this species has the potential

under certain conditions of becoming an agricultural pest. In

Zambia this snail is know as ‘chuzuya’.

The fine series of specimens in BMNH convincingly dem-

onstrates the wide range of variability in the shells of this

species (Figs. 41-51).

Bequaertina fraterculus (Dupuis & Putzeys, 1900)

Figs. 52, 53

Ganomidos fraterculus

Dupuis & Putzeys, 1900:xiii, text fig. 18.

Callistoplepa fraterculus

Pilsbry, 1905:129, pl. 47, fig. 23 (ex Dupuis & Putzeys);

Germain, 1909:90; Pilsbry, 1919:80; Bequaert & Clench,

1934¢:114.

SHELL. Shell ovate-turrite, extremely thin, translucent.

Whorls 6—61/4. The second and third whorls are comparatively

large, long and nearly straight-sided, producing characteristi-

cally a collared blunt mammillate apex. The fourth and

subsequent whorls are convex and expanded proportionately.

Sutures moderately deep. Last whorl large, but not inordi-

nately so, 78% of shell length; range for 6—6'/s whorls,

77-80% (n = 10). Aperture oval, pale milky within. Col-

umella brown, long, slender, nearly straight, obliquely trun-

cated. Outer lip thin, evenly arcuate; receding at base in

profile. Parietal callus diaphanous.

Highly irregular castaneous streak and spot brush marks,

some of which are closely highlighted with buff adaperturally,

are characteristically found on the last whorl; however, these

may be reduced to a few obscure dull buff spots more or less

limited to the peripheral carina. The earliest signs of this

diagnostic colour pattern are seen in the third whorl. The

ground colour intergrades from pale horn colour of the

unicolorous nepionic whorls to an obscuring dark olive brown

of the last whorl.

Faint, closely oppressed minute crescentic granulations

may appear in the second half of the otherwise smooth first

whorl. These granulations dominate the second whorl, giving

it an evenly, delicately engraved appearance. In contrast,

rather bold, diagonally oriented arcuate plicae, arising from

the suture below, disrupt this sculpture in a narrow basal

zone. This plication, which may start even in the last part of

the first whorl, rather abruptly disappears in the third whorl,

leaving a delicate, closely aligned series of nearly transverse

lirae, uninterrupted except for small, sparce, ghost-like

patches of granulations in some specimens. This marks the

end of the nepionic whorls. In the fourth and fifth whorls, the

lirae become increasingly strongly prosocline, more regular,

apically arcuate immediately below the suture, and eventu-

ally so prominent that they obscure the growth lines. In the

sixth whorl, the lirae become finer, less regular, more nearly

orthocline, and often interspersed with short parallel lirellae.

In the fifth whorl, spiral lirae, which seem to join rather than

interrupt the prosocline lirae, appear somewhat irregularly,

producing a slight checkerboard effect, reminiscent of the

sculpture of Achatina tracheia Connolly, 1929. These spiral

lirae, which are quite close together near the suture, irregu-

larly diminish in number and intensity toward the columella.

Very shallow malleations, starting subtly in the fourth whorl,

intensify the checkerboard effect. A subdued but apparent

carina appears at the periphery, where the lirae are seen to

bend slightly. The carina is more conspicuous in the younger

specimens. Abrasion of the shell along the periphery and

along some of the more prominent lirae visually intensifies

the carina and the sculpture by exposing the brilliantly shiny

inner periostracal layer. The outer layer of the periostracum

imparts a characteristically dull, corneous luster to the shell.

SOFT ANATOMY. No known alcohol preserved specimens.

TYPE MATERIAL. Nine of the ten known specimens of this

rare species were collected by Dupuis and are considered

syntypes. Six of these were identified and labelled as

‘cotypes’: three in Tervuren (MRAC no.5140-5142) and

three unnumbered specimens in Bruxelles (IRSN, General

Collection). The seventh specimen (MRAC no.5139) was

labelled as the one figured by Dupuis & Putzeys (1900); their

figure 18 (reproduced by Pilsbry, 1905) is so generalized that

it cannot specifically identify with any of the syntypes.

Unfortunately, the apex of this specimen had been broken

and cemented together, with a resultant alteration of the shell

configuration and length. The damage possibly occurred

during the precarious period when the artist had the speci-

men. Two additional syntype specimens (IRSN) were given

to Dauzenberg by Dupuis, one of which was collected in

Nsendwe. Under these circumstances, and since the authors

did not designate a holotype, the largest and finest of the four

MRAC syntypes (no.5140) is here selected as the lectotype of

Dupuis & Putzeys’ Ganomidos fraterculus (Figs. 52, 53;

Table 9). The tenth known specimen, acquired by Preston,

was passed on to V.W. MacAndrew, and is now in the

BMNH. Its apex was damaged and repaired-naturally. This,

too, most probably was collected by Dupuis and should be

considered a paralectotype.

TYPE LOCALITY. The Island of Mvula on the Lualaba

River, Zaire. J.C. Bequaert was unable to find this locality

on any map (Pilsbry, 1919:11, 19), nor is it listed in the

A.R. MEAD

Table 9 B. fraterculus — Representative shells measurements.

Greatest Aperture Last %

Whorls Length Width Length Width whorlLW/L % W/L

6 51.0 28.7 30.0 16.4 40.7 80 56 Mvula

(BMNH)

PLec

6 50.5 29.0 30:0°) 16:9539'34a78 57 Mvula

(IRSN)

PLec

61/4 50.4 28.5 30S 20) S89 77; 56 Mvula

(MRAC)

5140 Lect*

6 47.5 27k4 2728) <16:0\ 6375079 58 Mvula

(MRAC)

5139

6 46.3 26.8 28: Tale) 50a 19 58 Mvula

(MRAC)

5141 PLec

6 44.4 26.6 27 DMA Sie 3500079 60 Nsendwe

(IRSN)

PLec

6 44.3 26.4 2Osiem 14-Dien 54500 70, 59 Mvula

(IRSN)

PLec

5% 40.2 26.4 26:2" * 14:2 “326% 81 66 Mvula

(MRAC)

5142 PLec

5% 37.5 23.0 22.6 13.2 29:7 79 61 Mvula

(IRSN)

PLec

5% 36.5 QF, 22:0) Si2 2s 28.207 59 Mvula

(IRSN)

PLec

Total specimens examined: 10. Sources: BMNH, IRSN, MRAC.

current USBGN series. The only lead is the fact that a

single specimen of this species in the Dautzenberg collec-

tion (IRSN) bears the data: ‘Nsendwe, Congo. P. Dupuis

Coll., leg & ded’. At the same time that Dupuis & Putzeys

described Ganomidos fraterculus (1900), they described

the new species Perideriopsis mvulaensis, giving its locality

as ‘ile de Mvula (en face de Nsendwe)’. Bequaert in Pilsbry

lists Nsendwe as 3° 05’ S, 26° E. This location nearly

coincides with the important crossroad Kindu-Port-Empain

2° 57’ S, 25° 56’ E, hence Mvula must be very close to 3° S,

ZOE.

DISTRIBUTION. The two known localities for this species are

the Island of Mvula and the nearby onshore village of

Nsendwe in Zaire. Given the nature of the riverine environ-

ment, this species may well be found on other of the many

small islands and possibly in shore sites along this northward

flowing remote section of the Lualaba.

REMARKS. Phylogenetically, this species appears to stand

between B. graueri and the plesiomorphic B. pellucida.

Based on a single specimen from Nsendwe (BMNH, Preston

‘L/K 13/11/01’), B. pellucida may be sympatric with B.

fraterculus.

Bequaertina graueri (Thiele, 1911)

Figs. 54-57

Achatina graueri

Thiele, 1911:205, pl. 5, fig. 43; Pilsbry, 1919:78.

NEW SUBFAMILY AND GENUS ACHATINIDAE

Achatina (Cochlitoma) graueri

Pilsbry & Cockerell, 1933:366, pl. 1, fig., 1, la.

Callistoplepa graueri

Bequaert & Clench, 1934c:115.

Callistoplepa graueri

Schouteden, 1935a:110; 1935b:287; Oliver, 1983:9.

Callistoplepa babaulti

Germain, 1936:151, text fig. 46.

SHELL. Shell lacrimoid-subsuccineiform; the thick, durable

periostracum appears to provide more support than the thin

calcareous shell. Whorls 6-6/2, rarely 7. Spire tapered,

mammillate, elevated-conic, clearly shorter than aperture

length. The first 1'/2 whorls form a bluntly obtuse dome. The

second and third whorls expand only slightly, but descend

rapidly; this produces the strongly mammillate apex. The

fourth and following whorls descend proportionately, are

increasingly more convex, but expand rapidly to produce an

inordinately large last whorl. Sutures moderately deep,

increasingly so between fifth and sixth whorls. Last whorl

80% of shell length; range for 5—7 whorls, 77-83% (n = SO).

Aperture oval to elongate; very thin blue-white shelly layer

within. Columella markedly slender, almost entirely concol-

orous, slightly arcuate or nearly straight; typically very nar-

rowly and obliquely truncate. Outer lip thin, usually

somewhat obscured by the more rapidly advancing thick

periostracum, which tends to curl into the aperture in dried

specimens. The arc of the outer lip is often greatest below

midway and basally extended well below the columellar

truncation in the more mature specimens. Parietal callus

appears to be virtually absent in the smaller specimens and

barely visible in the larger and older specimens.

The first three whorls are unicolorous beige-buff to buff-

horn colour. Faint, diffuse wide transverse castaneous bands

may emerge in the second whorl and become darker, more

conspicuous and fragmented in the third and fourth whorls.

These gradually give way to narrow transverse bands of

various shades of brown, irregularly appearing coincident or

alternating with growth bands. The darkest, broadest bands

usually indicate interrupted growth. In some specimens the

transverse bands may be essentially absent. Ground colour

varies within and between specimens in a spectrum of olive-

buff, light yellow-brown, olivaceous brown and medium dark

brown. The fifth whorl often has a contrastingly paler ground

colour; hence, a 5-whorl specimen, with its short last whorl,

may appear to be a different species.

Pronounced spiral striae, usually 5-7, starting in mid-first

whorl are offset by beaded or semilunar granules that are

irregular in size and not aligned in vertical rows. The spiral

rows of granules increase to 12-17 or more and the now very

small granules become more uniform, more prominent, and

gradually more transversely aligned in the second and third

whorls. At the end of the third whorl, there is a prominent

delineation that marks the end of the nepionic whorls. At this

point, the granulate sculpture is rather abruptly taken over by

thin, slightly prosocline growth wrinkles that increase in size

and number until they dominate in the fourth whorl. In the

fifth whorl, there is a reemergence of the granulate sculpture

_ in the form of notched, subquadrate, tile-like plates that

often resemble the block letters K,H,W,V,Y & M, as is

strikingly seen in Achatina reticulata Pfeiffer, 1845. This

reappearance of the granulate sculpture, accentuated by the

deeper spiral striae, dominates the fifth whorl, although the

growth wrinkles continue to increase in calibre and become

subcrenulate apically. In the usually somewhat darker

coloured last whorl, the granulate sculpture is once again

greatly reduced, but may be highlighted here and there by an

isolated sporadic deep section of a spiral stria. The earliest

malleations appear subtly in the third or fourth whorl and

intensify in the following whorls. They consist of usually short

spiral or diagonal ridges that join or distort the growth

wrinkles to form a coarse, irregular raised network of welts.

Often entering into this is a very faint elevation at the

periphery, below which the otherwise nearly uniform sculp-

ture is reduced. An occasional specimen is entirely without

malleations. Their irregular appearance is probably explained

by thin shell, tough perostracum, and environmental impacts.

Continued shell deposition from within ‘fixes’ the dents in

place. Rarely is the outer layer of the thick periostracum

broken enough to reveal the shiny inner layer.

SOFT ANATOMY. Alcohol preserved specimens available

82/dissected 9. Zaire: MRAC (Mulungo, no.204.632-633)

2/2; (Kahusi-Tshibati no.610.302—305, 342-343) 79/6; ZMB

(lectotype) 1/1. Unfortunately, all dissected specimens except

the lectotype, were exposed excessively to formalin during

their preservation; thus, even with prolonged special treat-

ment, their tissues remained hard and the specimens were

exceedingly difficult to dissect. Pilsbry and Cockerell (1933)

described the living animal as very pale ochreous with head

and broad based tentacles faintly bluish.

Clearly, the most conspicuous and characteristic aspect of

the basal genital conduits is the vaginal retentor (VR) muscle

system, grossly dominating in ventral view (Fig. 20). Slender,

parallel, glistening, partly fused muscle bands pass ventrally

from the vagina (V) to the right body wall along a dorsolat-

eral line from immediately posterior to the genital aperture to

the junction of the mantle and the right body wall. From the

left side of the vagina, this system gives rise to a series of

muscle bands that starts at the peniovaginal angle and binds

tightly together the equally prominent apical vas deferens

(AVD) and the free oviduct (FO). A more bold series of

distinct, but laterally fused muscle bands pass from the left

lateral aspect of the AVD to the same right dorsolateral line

of attachment, further obscuring the basal genital structures.

(These bands are shown cut short in this figure.) Apically this

whole muscle system is reduced largely to a thin, transparent

membrane attached to the junction of the AVD and the FO.

At this same junction, a similar membrane passes to the

spermatheca (S) and attaches it to the FO. The S is propor-

tionately extremely large in this species. In four of the nine

specimens examined, the slender apex of the S is folded back

on itself, as shown here; this is apparently a common artifact

of preservation. In the rest, the apex is extended, and

although it may appear to go apically beyond the junction of

the AVD and FO, it is not attached to the spermoviduct (SO)

as it is in many achatinids. The spermathecal duct (SD) is so

short and broad that when the S is gorged, it seems to be

sessile on the V.

Short, often diagonal or anastomosing muscle bands, sepa-

rate or combined, are found at the base of the penis sheath

(PS) and the V. These eversion muscle bands (EM) initiate

the precopulatory extroversion of the genital atrium (GA).

The PS normally completely envelops the short, diminutive

penis (P); in only one of nine specimens, the apical P

projected slightly. The penial retractor (PR) has its origin on

the apical P and inserts on the mid-forward diaphragm at or

near the junction with the body wall. A single specimen,

shown here, had a bifurcate insertion; a multiple insertion is

not rare in the achatinids. At the origin of the PR, the muscle

fibrils enshroud smoothly and completely the apical P and the

robust basal vas deferens (BVD), greatly obscuring the

relationships in the basal male conduit (Fig. 21). Contributing

to this, the PS is free from the P only in the approximate

upper half of the left side. On the right side, fibrils from the

PR extend basally to form a dense webbing that seems to

invade the substance of the outer wall of the P and the inner

wall of the PS. Below the PS, this infusion of tissues, along

with the EM, obscurely defines the wall of the penial atrium.

This atrium connects the lumina of the P and GA (Fig. 22).

The lumen of the P is thickly carpeted with vermiculate

rugae, which become slender and elongate near the GA,

resembling the plicae of the basal V. No verge or pilaster is

present. Eccentrically in the apex of the P, a small aperture

leads to the narrow lumen of the extremely thick-walled BVD

and AVD. These two structures provide the physical support

for the extroverted, highly expansile P. Since they have

supportive and ejaculatory functions, they may explain the

extreme development of the VR. Approximately 8-10 mm

basal to the junction of the AVD and FO, the lumen of the

AVD enlarges considerably and forms an elongate, thin-

walled chamber, which conceivably functions as a secondary

seminal vesicle (SSV). Apically this chamber becomes saccu-

lar with thin elongate rugae.

A single specimen (610.343) was gravid. Six large, fully

formed eggs, 9.5 X 6.5 mm, were in the apical (oviductal),

cream coloured portion of the spermoviduct; no eggs were in

the contrastingly light brown uterine basal portion. This

specimen and the five others in the same lot had robust,

mature coloured, fully formed reproductive tracts. The field

data thus indicate that in the Kivu, breeding takes place in

October.

Kidney is large, typical of the subfamily, broad anteriorly

and truncated posteriorly. Five ovotestis acini are embedded

in the columellar surface of the right (apical) lobe of the

digestive gland. The inconspicuous anterior aorta is on the

left posterior surface of the lung, where it penetrates the

diaphragm. The hermaphroditic duct, similar to that of B.

pintoi (Fig. 19), is trimerous with an abruptly enlarged

saccular central portion, 5.3 x 2.4 mm.

The following anatomical characters distinguish this species

from B. pintoi: basal genital fascia gross, forming a VR with a

massive system of muscle bands; AVD, BVD and FO are all

about the same width; P is strikingly short and stubby,

normally retained entirely or nearly entirely within the PS;

BVD wide, about as long as wide.

TYPE MATERIAL. Thiele (1911) described this species from a

single mature specimen (ZMB no.101937) in the Schubotz

collection and two small juvenile specimens from the Grauer

collection. The mature specimen is nearly full grown but only

moderately large (Figs. 54, 55; Table 10). Thiele’s fine line

illustration shows it slightly larger than natural size in aper-

tural view only. This specimen, whose soft parts are in

alcohol, I labelled as the lectotype of Thiele’s Achatina

graueri when I examined it in East Berlin (ZMB) in August

1989. At that time, the single available very dark coloured

juvenile specimen (Table 10) therefore was labelled paralec-

totype. Since then a second small paralectotype has been

found there and so labelled by Kilias (1992).

Germain (1936), without any apparent knowledge of

Thiele’s species, described and figured the junior subjective

A.R. MEAD

synonym Callistoplepa babaulti from two specimens collected

by Babault in Kitembo, Kivu, comparing it only with C.

marteli. The two Paris syntypes (MNHN) are large, typical

specimens of Bequaertina graueri (Table 10). The larger,

finer specimen is here selected as the lectotype of Germain’s

C. babaulti (Figs. 56, 57).

TYPE LOCALITY. Idjwi Island (= Kwidschwi, Kwidjwi,

Idjewi), Lake Kivu, Zaire 2° 09’ S, 29° 04’ E.

DISTRIBUTION. This species occupies a 450 km long, narrow,

north-south corridor in the upper Rift Valley of Zaire

between Beni and Uvira (Fig. 16). So far, it has been found

only as far west as Kitembo and projects slightly east of Zaire

into Lobengera Mission, Rwanda and into Ibanda, Uganda.

Eventually, it will also be found in Burundi.

REMARKS. This is the largest and most distinctive species in

the genus. It is most closely related to Bequaertina fratercu-

lus. Because of its size and colour, and because in some

localities it is sympatric with Achatina stuhlmanni von Mar-

tens, 1892, it has been confused with that species. However,

since A. stuhlmanni has a shorter, broader spire, a more

obtuse apex, and a distinctive spirally fine-combed wavy

sculpture (Bequaert & Clench, 1934a:3), it can readily be

Table 10 B. graurei - Representative shells measurements.

Greatest Aperture Last %

Whorls Length Width Length Width whorlLW/L % W/L

dl 93.2 45.7 52:33, 266) 718" 77 49 Katana

(MRAC)

610.304

61/4 89.7 47.0 55:5) $28!457256) ‘Sil 52 Tshibinda

(MRAC)

5115

if 89.3 43.6 50, 6 255: 0078 49 Katana

(MRAC)

610.302

61/4 78.4 42.5 46.8 24.3 62.5 80 54 Kitembo

(MNHN)

Lect C.

babaulti*

6 77.0 38.6 45.9 23.5 63.0 82 50 Uvira

(MRAC)

607.170

61/4 71.8 41.4 41.9 23.4 56.1 78 58 Kitembo

(MNHN)

PLec C.

babaulti

6 62.0 36.5 37.8 20.6 49.1 79 59 Idjwi

(ZMB)

101935

Lect A.

graueri'*

5% 49.8 29.1 31.8 16.4 40.0 80 58 Idjwi

(MCZ)

6 49.2 22.8 31.0 145 39.5 80 46 Beni

(MRAC)

5119

514 42.0 25u 27.0 14.0 33.0 78 61 Idjwi

(ZMB)

101936

PLec A.

grauert

Total specimens examined: 54. Sources: IRSN, MCZ, MNHN, MRAC,

NMW, RMNH, SMNH, UMMZ, ZMB.

NEW SUBFAMILY AND GENUS ACHATINIDAE

distinguished. Pilbry & Cockerel (1933) reported seeing two

living specimens ‘crawling about 5 feet up on the trunk of a

tree in the forest above Tshibinda at about 2100 m’ in Zaire.

The largest collection of this species, including many alcohol

specimens, is to be found in Tervuren (MRAC). B. graueri is

the type species of the genus.

Radulae and jaws

D’Ailly (1896:69) was the first to describe and illustrate the

radulae of Callistoplepa shuttleworthi and C. barriana. Pilsbry

(1904:ix, xv) referred to d’Ailly’s work but reproduced only

the illustration of the latter species. He also reported (p. 72)

that G. Schacko (1881) (nec ‘Schako’) found ‘A. pulchella has

. a very small central tooth’. The present work shows

Schacko’s specimen was therefore misidentified. Thiele

(1929:560) examined and illustrated in part the radula of C.

shuttleworthi. He also examined but did not illustrate the

radula of Leptocala mollicella and pointed out that the middle

tooth was a little smaller than the neighbouring teeth. Possi-

bly on the basis of this observation, he prophetically juxta-

posed ‘Callistoplepa and Leptocala. More recently, Ortiz &

Ortiz (1959:47) also illustrate the radula of C. shuttleworthi,

but the focal plane of the microscope was apparently too low

and the configurations of the teeth are misleading. The

radulae in the present project were prepared according to the

recommendations of Solem (1972) and the emphasis has been

placed on the rachidian teeth and the adjacent laterals.

Because the soft anatomies of the four callistoplepine

species are so similar, it was not surprising to find the radulae

of C. barriana, C. shuttleworthi and L. mollicella (Figs.

58-63) to be remarkably similar. This fact supported the

earlier decision to conserve intact the odontophores of the

two extant soft anatomy specimens of L. petitia. It is assumed

with confidence that the radula of this latter species is

essentially like the others. In those examined, all have bold

rachidian basal plates and a broad functional rachidian tooth

that is one-half to two-thirds the size of the adjacent lateral

teeth. A second type of tooth is found in the first series of

laterals, which similarly consists of broad, solitary meso-

cones, but with conspicuous laterally asymmetrical basal

plates. A third type of tooth arises in the second series of

laterals, wherein the mesocones angle increasingly more

mesad and small ectocones gradually arise. These merge

almost imperceptibly into a fourth type, the tricuspid margin-

als with minute irregular endocones, broad shorter serrate

mesocones, and increasingly reduced basal plates that no

longer contact the teeth posterior to them. The greatest

irregularity within and between specimens occurs in the

gradient between the bicuspid laterals and the tricuspid

marginals. Hence the following formulae (tooth numbers

from centre to right) are only approximate: C. barriana

C-31-55-84, C. shuttleworthi C-17-28-49, and L. mollicella

C-19-28-66.

The available radula specimens in only two of the five

Bequaertina species have produced an incomplete and some-

what confusing picture in this genus. The basal genital

| systems of B. pintoi and B. graueri are fundamentally similar

| — both reflecting affinities with the Zaire Basin subgenus

Achatina (sensu Bequaert, 1950). It thus was anticipated that

the radulae also would be similar. The radula of B. pintoi

(Figs. 64, 65), not surprisingly, was found to be of the same

type as that of Achatina craveni E.A. Smith, 1881 (Figs. 68,

69). Both have greatly diminished, essentially nonfunctional

rachidian teeth that are almost concealed by the adjacent

laterals. And both have broad based, nearly tricuspid laterals

with angular ectocones, broad mesocones and endoconal

flanges. In addition, the mesocone column of each lateral

tooth directly contacts and supports the broad basal plate of

the tooth immediately posterior to it. It should be noted that

A. craveni, on the basis of its soft anatomy, belongs in

Bequaert’s subgenus Achatina rather than where he has

placed it in his subgenus Lissachatina. The surprise came in

the radula of B. graueri (Figs. 66, 67), with its large functional

rachidian tooth, attenuated massive basal plates, more

restricted contact support between horizontal rows of teeth,

and an imperceptible gradient into the marginal teeth. Within

the genus, B. pintoi and B. graueri are at the conchological,

geographic and ecological antipodes. The known plasticity in

molluscan radulae suggests that undetermined different feed-

ing demands in dissimilar habitats have produced the con-

trasts in the radulae of these otherwise two closely related

species. B. graueri and B. fraterculus appear to be very

closely related conchologically. There is a question now

whether the radulae will support this assumption. In reality,

the relationships in Bequaertina will not be understood until

both the soft anatomies and radulae of B. fraterculus, B.

pellucida and B. marteli are known. Radula formulae: B.

pintoi C-42-25, B. graueri C-59, A. craveni C-34-24.

The castaneous callistoplepine jaw forms an unusually

broad middle section that quickly tapers on each side to about

half its width and curves inward at the ends into a collariform

structure. Its surface is featureless except for microscopic

horizontal growth increments best seen under transmitted

light. Measurements: C. barriana2 x 1.3 mm, C. shuttlewor-

thi 1.5 x 0.6 mm, L. mollicella 1.7 x 0.4 mm. The illustra-

tion of Ortiz & Ortiz (1959:46) for C. shuttleworthi appears

excessively broad.

The jaw of B. pintoi forms a light castaneous nearly uniformly

slender rainbow arc, 5.5 X 1.1 mm, with ca 36 irregularly placed

vertical ridges. In B. graueri the jaw forms a lower arc,

4.4 x 1.3 mm, with very obscure vertical lineations.

The more slender jaw of A. craveni forms a fulvous,

somewhat depressed arc, 4.5 x 0.7 mm, with ca 35 fairly

uniformly distributed vertical riblets.

ACKNOWLEDGEMENTS. Special thanks are given to the curators and

their assistants of 28 museums, and to three private collectors, all

individually referred to in the Acronyms — Institutional & Personal

Collections. I am especially grateful to Drs A.C. van Bruggen

(Rijksmuseum van Natuurlijke Historie, Leiden), B. Verdcourt

(Royal Botanic Gardens, Kew), and W.F. Sirgel (Stellenbosch

University) for reviewing the manuscript and offering valuable

suggestions; and to Dr P.K. Tubbs, Executive Secretary of the

International Commission on Zoological Nomenclature, for consulta-

tions on the Code. I am also indebted to many others who have

provided important assistance, among them: Donald B. Sayner,

Charlotte Ernstein and Virginia Childs of the Scientific Illustration

staff at the University of Arizona for photographs in Figures 27-42,

45-46, 52-57; the Photography Service staff of the British Museum

(Natural History) for photographs in Figures 23-26, 43-44, 47-51;

David L. Bentley of the Electron Microscope Facility, Division of

Biotechnology, and Professor Michael A. McClure, both of the

University of Arizona, for assistance in the SEM photographs;

Emilee M. Mead of the University of Arizona Teaching Center who

drew the map and prepared the photographic layouts; Yolanda

Baldonado Whigham of the U.A. Department of Ecology and

Evolutionary Biology for word processing; and my wife Eleanor who

has helped in countless ways.

34 A.R. MEAD

Figs 58-63 Dorsal and dorso-right lateral views of radulae: 58,59 Callistoplepa barriana (SMF, O. Boettger). 60, 61 C. shuttleworthi

(SMNH no. 10). 62, 63 Leptocala mollicella (MRAC no. 795.638).

NEW SUBFAMILY AND GENUS ACHATINIDAE

Figs 64,65 Bequaertina pintoi (BMNH no. 1953.8.15.562—564, mixed lot). 66,67 B. graueri (MRAC no. 204.633). 68, 69 Achatina

craveni (BMNH no. 1953.8.15.562—564, mixed lot). Bar scale = 1 pm.

ACRONYMS - INSTITUTIONAL &

PERSONAL COLLECTIONS

AMNH_ New York: American Museum of Natural History (W.K.

Emerson)

ANSP Philadelphia: Academy of Natural Science (A.E. Bogan)

BMNH _ London: British Museum (Natural History) (P. Mordan,

F. Naggs)

BV Kew, Royal Botanic Gardens: Bernard Verdcourt

CMNH Pittsburg: Carnegie Museum of Natural History (J.E.

Rawlins)

FMNH_ Chicago: Field Museum of Natural History (A. Solem)

GNM G6teborg: Naturhistoriska Museet (I. Levinsson, H.W.

Waldén)

HM Newquay, Cornwall, England: Hazel Meredith

IRSN Bruxelles: Institut Royal des Sciences Naturelles (J. van

Goethem)

LNK Karlsruhe: Landessammlungen ftir Naturkunde (H-W.

Mittmann)

MCZ Harvard: Museum of Comparative Zoology (K.J. Boss)

MNHN Paris: Muséum National d’Histoire Naturelle (S. & A.

Tillier)

MRAC_ Tervuren: Musée Royal de l'Afrique Centrale (P.L.G.

Benoit, F.A. Puylaert)

NG Blantyre, Malawi: W. Noel Gray

NHMB Bern: Naturhistorisches Museum (J.J. Oberling)

NHMW Wien: Naturhistorisches Museum Wien (E. Wawra, O.E.

Paget)

NM Pietermaritzburg, South Africa: Natal Museum (R.N. Kil-

burn)

NMB Basel: Naturhistorisches Museum (C. Stocker-

Unternahrer)

NMW _ Cardiff: National Museum of Wales (A. Trew, P.G.

Oliver)

RMNH Leiden: Rijksmuseum van Natuurlijke Historie (A.C. van

Bruggen, E.Gittenberger)

RMS Edinburgh: The Royal Museum of Scotland (D. Heppell)

Cape Town: South African Museum (J. Pether)

SMF Frankfurt: Forschungsinstititut Senckenberg Natur-

Museum (R. Janssen)

SMNH _— Stockholm: Naturhistoriska Riksmuseet (A. Warén, C.

Holmquist)

UHZI Hamburg: Universitat Zoologisches Institut und Museum

(R. von Cosel)

UMMZ Ann Arbor: University of Michigan Museum of Natural

History (J.B. Burch)

USNM_ Washington, D.C.: U.S. National Museum of Natural

History (R. Hershler)

UUZM _ Uppsala: Universitets Zoologiska Museum (A. Franzén,

L. Wallin)

ZMB Berlin: Museum ftir Naturkunde der Humboldt-

Universitat (R. Kilias)

ZMUC_ K@gbenhavn: Zoologisk Museum — Universitet (J. Knud-

sen, T. Schigtte)

ZSM Munchen: Zoologische Staatssammlung (R. Fechter)

A.R. MEAD

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Bull. nat. Hist. Mus. Lond. (Zool.) 60(1): 39-104

On Recent species of Spiraserpula

Regenhardt, 1961, a serpulid polychaete

genus hitherto known only from Cretaceous

and Tertiary fossils

T. GOTTFRIED PILLAI

Marine Biological Services Division, Department of Zoology, The Natural History Museum, Cromwell

Road, London SW7 5BD

HARRY A. TEN HOVE

Universiteit van Amsterdam, Instituut voor Taxonomische Zodlogie, Postbus 94766, 1090 GT

Amsterdam, Nederland

CONTENTS

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Synopsis. A group of Recent serpulid species related to the genus Serpu/a Linnaeus, 1758, but differing from it in

two important characters, is described in this paper. The first is a hitherto undescribed character, namely, the

possession of internal tube structures which consist of longitudinal ridges and other structures, the form and

arrangement of which, in combination with characters of the worms themselves, served to separate the various

species. The second is that the thoracic membranes of the two sides in the worm do not unite ventrally at the end of

the thorax to form a flap or apron as in Serpula. These characters are also common to 18 species, including three

previously described ones. On the basis of the tube structures, these Recent species can be referred to the genus

Spiraserpula Regenhardt, 1961, which was previously known only from fossils (Pillai, 1993). Scissiparity was

observed in at least three of its species. A key to the known Recent species of Spiraserpula and a discussion on the

Issued 23 June 1994

systematics of the genus are included.

INTRODUCTION

In the course of a study of the serpulids currently referred to

the genus Serpula Linnaeus, 1758, it was observed that the

worms of larger species could frequently be extracted undam-

aged from the anterior ends of their tubes with a pair of fine

forceps, while they were invariably damaged in the process in

certain small species as, for instance, in the well-known

Mediterranean species Serpula massiliensis Zibrowius, 1968.

In almost every collection of the latter, the worms which had

been previously extracted from their tubes were incomplete

posteriorly, and the ends of their longitudinal musculature

provided evidence of their having been forcefully broken off

from the rest of the abdomen. The cause of the difficulty in

extracting complete worms was revealed by opening their

tubes carefully from their anterior ends all the way to their

posterior ends. It was found that the posterior end of the

abdomen was retracted very tightly into the posterior coiled

part of the tube and, quite surprisingly, against a longitudinal

row of sharp serrations projecting from the inside of the tube.

Examination of more material showed that these were consis-

tent for S. massiliensis.

Study of similar material from various other geographical

localities revealed the existence of species with other forms of

internal tube structures. Evidently, these serve for anchorage

of the worm when withdrawn into the tube, and thereby,

have an additional protective function. The form and

arrangement of the internal tube structures, in combination

with characters of the worms themselves, served to separate

the various species. They are absent in Serpula Linnaeus,

1758, and have not been described in any of the other known

genera of Serpulidae. They differ from the transverse tabulae

of certain serpulids, an account of which is given by Lom-

merzheim (1979). Another important character common to

the group is that, unlike in Serpula, the thoracic membranes

of the two sides are not united posterior to the thorax to form

a ventral flap or apron.

In the search for a name for this group, the genera

Pseudoserpula Straughan, 1967 and Protoserpula Uchida,

1978, were considered, among others. The former was found

to be invalid, and an account of the study which led to this

conclusion is provided under Spiraserpula minuta,

(Straughan, 1967), in this paper. It was not possible to

examine the type specimen of Protoserpula to establish

whether it has ITS or not. It is not in the National Science

Museum, Tokyo, and other efforts to locate it were unsuc-

cessful.

H. Zibrowius of Station Marine d’Endoume, Marseille,

who went through the manuscript of this paper, and the

second author discussed the group with M. Jager of Rohrbach

Zement, Dotternhausen, Germany,who re-examined the fos-

sil serpulids studied by him (Jager, 1983), and other material,

and found that some of them too possessed internal tube

structures, although they had not been reported earlier. The

collaboration which followed (pers. comm.) led to a study of

likely Cretaceous and Tertiary serpulid genera and species

(Pillai, 1993), which revealed that the group belongs to the

genus Spiraserpula Regenhardt, 1961, previously known only

from fossil species. Spiraserpula Regenhardt, 1961, has prior-

ity over Protoserpula Uchida, 1978, even if the latter were to

possess internal tube structures, henceforth referred to in the

text as ITS (vide Jager 1993)). Zibrowius (1972) described a

Recent spirorbid species belonging to the genus Neomicrorbis

Rovereto, 1904, which was previously known only from

Cretaceous and Tertiary fossils.

In three of the Recent species of the genus Spiraserpula

definite proof of asexual reproduction was found, in the form

of branching tubes, corroborated by the presence of a parent

with a schizont in one tube of Spiraserpula snellii sp. nov.

Asexual reproduction had previously been reported for the

genera Filograna Berkeley, 1835, Filogranula Langerhans,

1884, Josephella Caullery & Mesnil, 1896, Salmacina Cla-

parede, 1870 and Rhodopsis Bush, 1905 (ten Hove, 1979;

Ben-Eliahu & ten Hove,1989). Pillai (1993) reports the

occurrence of tube branching in the fossil species Spiraserpula

versipellis Regenhardt, 1961. It would not be surprising if it

turns out that scissiparity takes place in most, if not all,

species of the genus Spiraserpula, in view of their aggregated

occurrence.

Nineteen species of Spiraserpula, including the three

known ones referred to above and an unnamed one, are

described. They come from the Mediterranean, Madeira,

Canary and Cape Verde Islands, Gulf of Mexico, the Carib-

bean and Panama, the northern Red Sea, Mozambique, the

eastern islands of Indonesia, Eastern Australia, Japan and

New Caledonia.

T.G. PILLAI AND H.A. TEN HOVE

METHODS AND MATERIALS

The tubes and their internal structures, as well as whole

worms and parts were examined and drawn under a stereo

microscope fitted with a drawing attachment. Measurements

were taken with a pair of fine dividers against a scale having

an accuracy of 0.5 mm, of total length of the tube when

possible, external diameter of the tube, total length of the

worm, width of the thorax just posterior to the pair of collar

fascicles, length and diameter of operculum, length of the

opercular peduncle, and length of the longest radiole and its

pinnule-free tip when present. Radioles and thoracic seg-

ments were counted on both sides, while the abdominal

uncinal tori of one side were counted to determine the

number of abdominal segments. The chaetae were mounted

in polyvinyl lactophenol or aquamount and figured under the

oil immersion lens of a high power microscope fitted with a

drawing attachment. Measurements of chaetae were made

with an eyepiece micrometer standardised with a stage

micrometer. Scanning electron micrographs of chaetae of

some of the species are also provided (Plates 1-5).

The sources of material have been detailed under the

respective descriptions as well as the acknowledgements. Full

details of E. Atlantic stations surveyed by the ‘Tydeman’

Canary and Cape Verde Islands Expeditions of 1980, 1982

and 1986 (CANCAP-IV, VI and VII), (e.g. CANCAP

4.D14, 6.134) can be found in van der Land (1987); of E.

Indonesian stations sampled during the Indonesian-Dutch

Snellius II Expedition (e.g. Snellius II 4.051) in van der Land

& Sukarno (1986). The following abbreviations have been

used in the text: AM: Australian Museum, Sydney; AMNH:

American Museum of Natural History, New York; BM(NH):

British Museum (Natural History), London, presently, The

Natural History Museum, London; FSBC I: Florida Depart-

ment of Natural Resources, Invertebrate collection, St.

Petersburg, Florida; HUJ: The Hebrew University, Jerusa-

lem; MCZ: Museum of Comparative Zoology, Harvard;

NNM: Nationaal Natuurhistorisch Museum, Leiden (for-

merly Rijksmuseum van Natuurlijke Historie); NSMT:

National Science Museum, Tokyo; QM: Queensland

Museum, Brisbane; RMNH: Collection numbers of NNM;

SME: Station Marine d’Endoume, Marseille (most material

will be deposited later in the Musée Nationale d’Histoire

Naturelle,Paris); USNM: United States National Museum of

Natural History, Washington DC; V.Pol: Polychaete collec-

tion numbers of ZMA; ZLU: Zoological Laboratory,

Utrecht; ZMA: Zodlogisch Museum, Instituut voor Taxo-

nomische Zodlogie, Amsterdam; ZMH; Zoologisches Insti-

tut und Zoologisches Museum, Hamburg; ZMK: Zoologisk

Museum, Kgbenhavn.

TERMINOLOGY

The terminology used in this paper is explained in Figs 1 and

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961

Po MN ie, is

pe +

way

1967). B, S. ypsilon sp. nov.

| DIAGRAMMATIC REPRESENTATIONS OF

TUBES

| The various arrangements of ITS in the species described are

_ diagrammatically represented in Fig. 3.

‘Plate 1 Scanning electron micrographs of fractured ends of tubes showing internal structures. A, C & D, Spiraserpula lineatuba (Straughan,

DIAGNOSIS OF SPIRASERPULA Regenhardt,

1961

The original generic diagnosis of Spiraserpula was based only

on the tube of its fossil type species, S. Spiraserpula Regen-

hardt, 1961. Pillai (1993) provides an emended definition for

fossil species based on characters of the tube. However, the

recent species described here are distinguishable not only by

characters of their tubes but also of the worms themselves,

T.G. PILLAI AND H.A. TEN HOVE

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Plate 2 Scanning electron micrographs of chaetae. A-D, Spiraserpula massiliensis (Zibrowius, 1968): A, bayonet chaetae. B, thoracic uncini.

C, anterior abdominal uncini. D, flat trumpet-shaped abdominal chaetae. E & F, S. singularis sp. nov.: E, bayonet chaetae. F, abdominal

uncini.

and they have been taken into consideration in the following

diagnosis:

Tube with internal structures, usually towards its earlier

formed, coiled, posterior portions. They consist of internal

longitudinal ridges which vary in form and complexity in the

different species; they may be dorsal, along the convex inner

wall of the tube, and/or ventral, along the opposite side. They

may be laminar, serrated or unserrated, or have other forms,

and accessory lateral ridges or other structures may also be

present. An umbilicus and peristomes may be present. There

is usually an external granular overlay which bonds together

coils of individual tubes, or those of other tubes to form

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961

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| Plate3 Scanning electron micrographs of chaetae. A-D, Spirserpula nudicrista sp. nov.: A, bayonet chaeta. B, thoracic uncini. C,

abdominal uncini. D, flat trumpet-shaped abdominal chaetae. E-G, S. lineatuba (Straughan, 1967): E, bayonet chaetae. F, abdominal

uncini. G, flat trumpet-shaped abdominal chaetae.

44 T.G. PILLAI AND H.A. TEN HOVE

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Plate 4 Scanning electron micrographs of chaetae. A-D, Spiraserpula zibrowii sp. nov.: A, bayonet chaetae. B, thoracic uncini. C,

abdominal uncini. D, flat trumpet-shaped abdominal chaetae. E & F, S. caribensis, sp. nov.: bayonet chaetae.

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961

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Plate5 Scanning electron micrographs of chaetae. A—C, Spiraserpula caribensis sp. nov.: A, abdominal uncini. B, flat trumpet-shaped

abdominal chaetae. C-E, collar chaetae from Grenada material. F—-H, S. snellii sp. nov.: F, bayonet chaetae. G, abdominal uncini. H, flat

trumpet-shaped abdominal chaetae.

T.G. PILLAI AND H.A. TEN HOVE

Fig. 1 Terminology. A, Tube: d.cx.w., dorsal convex wall; d.r., dorsal ridge; v.cv.w., ventral concave wall; v.r., ventral ridge. B, Radioles

of both sides and operculum: o.p., opercular peduncle; p., pinnules; pf.t., pinnule-free tip; r., radius; rl., radiole; r.I.s., radioles of left

side; r.o., rudimentary operculum; r.r.s.,radioles of right side; t.r.I. triangular radial lobe; z.o., zygomorph operculum. C—G, Opercula:

cn., constriction between operculum and peduncle; i.r.g., inter-radial groove; ut.c., unthickened cuticle. bs.o., bell-shaped operculum;

r.r.1., rounded radial lobe; t.t.c., thickened transparent cuticle.

mutually bonded aggregations of a few to numerous individu-

als.

An operculum similar to that in Serpula, which is a

modification of the second most dorsal radiole, is often

present on one side with, correspondingly, a rudimentary

operculum on the other. There may only be a rudimentary

operculum on each side in certain species, while they may be

present in juveniles and completely lost in older specimens in

others. The shape of the fully developed operculum is charac-

teristic for a particular species; it may be funnel-shaped,

bell-shaped, zygomorphic or spherical. Its distal end may be

concave or convex and usually bears radii which end as

triangular or rounded lobes at the rim; but radii may also be

lacking in some species. Its cuticle may be unthickened or

thickened and transparent.

The number of branchial radioles is usually small, rarely up

to 14 pairs. Palps absent. A pair of prostomial ocellar clusters

is usually present. The number of thoracic chaetal tufts may

exceed the seven pairs commonly occurring in many genera

of Serpulidae, including Serpula, and those of the two sides

are more frequently asymmetrical than symmetrical. Up to 14

have been counted on each side. Histological work is needed

to ascertain the real extent of the segments, and their relation

to numbers of chaetal tufts and uncinal rows, etc. The term

‘chaetiger’ is, therefore, used here in the literal meaning of

‘hair bearer’ and not as a synonym of segment. The thoracic

membranes end anterior to the last thoracic chaetigers, also

more frequently asymmetrically than symmetrically. Unlike

in most species of Serpula sensu stricto, therefore, a post-

thoracic ventral flap (apron) is absent.

Collar fascicles bear chaetae of two kinds: bayonet

chaetae and limbate chaetae, the blades of both of which

are usually finely serrated. In the former, there are a few to

several comparatively large teeth, located at the distal end

of the shaft, separated from the bayonet-like blade by an

unserrated area (unserrated notch). The range in the

number of such teeth and the length of the unserrated

notch varies in the different species. Limbate chaetae bear

simple, more or less curved, blades. Thoracic and anterior

abdominal uncini may bear teeth in a single row (saw-

shaped), or are partly (saw- to rasp-shaped) or completely

rasp-shaped. Abdominal chaetae bear distally flat trumpet-

shaped ends, and are replaced by capillaries in the poste-

rior segments. The distal ends of the abdominal chaetae of

Serpula have been described as ‘trumpet-shaped’ in ser-

pulid literature. We have discussed the inappropriateness

of the comparison, as demonstrated by the scanning elec-

tron micrographs and drawings of these chaetae presented

in this paper, and our attention has also been drawn to this

by Zibrowius (pers. comm.). In order not to create confu-

sion, it was decided to retain ‘flat trumpet-shaped’, for the

present.

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961

Fig. 2 Terminology. A & B, Body of worm: a.t., abdominal tori; cap., capillary chaetae; col., collar; col.s., collar chaetae; d.l.g., dorsal

longitudinal groove; t.s., thoracic chaetae; t.t., thoracic tori; v.I.g., ventral longitudinal groove. C-G, Bayonet-shaped collar chaetae (after

ten Hove & Jacobs, 1984; all same magnification): ac.t., accessory teeth; b.b., basal boss; f.t., few teeth; I.s.b.,long serrated blade;

m.l.s.b., moderately long serrated blade; s.s.b., short serrated blade; s.t., several teeth; u.n., unserrated notch; v.s.dl.b., very short

dagger-like blade; v.s.t.b., very short tapered blade. H, Uncini, showing orientation in relation to the body of the worm: a., anterior; p.,

KEY TO THE KNOWN RECENT SPECIES OF

posterior.

_ SPIRASERPULA REGENHARDT, 1961

(See Figure 3 for terminology of ITS)

Tube with either dorsal or ventral internal longitudinal ridge

OM Wye rec itga sina dat ae enc R eRe ach ocieaie snGieisinaeeinpsraicesins

Tube with dorsal and ventral internal longitudinal ridges ....

Tube with dorsal longitudinal ridge only (Fig. 3, B—E)

Tube with ventral longitudinal ridge only, exceptionally with

fewasolatedttcethn (Eig Sail) tp pecn see cesses eeeeeetee esac eee ee 6

Dorsal ridge unserrated, shaped like an inverted V (Fig. 3, B)

Siseieoisieisla ci mta’apmisiains quatyctsea sheet egbildereeta S. singularis sp. nov. p.62

Worsalinidsersetrated (ica 3. CE) eense- cee eee nese eee eeeeee 4

Serrations of dorsal ridge deltoid (Fig. 3, C)

Beaten, etatavodae th ot eeome even tactic adakereiads S. deltoides sp. nov. p.80

Serrations of dorsal ridge not deltoid (Fig. 3, D & E) ........ 5)

48 T.G. PILLAI AND H.A. TEN HOVE

Fig. 3 Diagrammatic representations of ITS in the various species. A, Generalized drawing with all the main ITS. The orientation of the

tube and terminology used are the same for all the diagrams. Ant., anterior direction. d, dorsal side. d.cx.w., dorsal convex wall.d.I.r.,

dorso-lateral ridge. d.r., dorsal ridge. v, ventral side.v.cv.w., ventral concave wall. v.r., ventral ridge. B, S. singularis sp.nov. C, S.

deltoides sp. nov.; d.s., deltoid serrations. D, S. massiliensis (Zibrowius, 1968). E, S. capeverdensis sp. nov.; k.c.d., knob-like calcareous

deposits. F, S. nudicrista sp. nov. & S. snellii sp.nov.; v.v.r., variant form of ventral ridge. G, S. ypsilon sp. nov. & S. paraypsilon sp. nov.

H, S. sumbensis sp. nov. 1, S. iugoconvexa sp. nov. J, S. vasseuri sp. nov. K, S. plaiae sp. nov. L, S. caribensis sp. nov. & S. lineatuba

(Straughan, 1967). M, S. discifera sp.nov.; t.d., transparent discs. N, S. karpatensis sp. nov. & S. minuta (Straughan, 1967). O, S. zibrowit

sp. nov.

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961

10.

12.

13:

14.

Maximum number of radioles 6 pairs, abdominal segments

about 50. Operculum may or may not be present. A shallow

water species, down to about 60 m, rarely in deeper water

Ree aio seteiate «oeieh epee S. massiliensis (Zibrowius, 1968) p.49

Maximum number of radioles 8 pairs, abdominal segments

about 145. Operculum absent. A deep water species occurring

at depths of about 75—200m ..... S. capeverdensis sp. nov. p.54

Tube creamish white, trapezoidal in cross-section. Operculum

absent. Maximum number of radioles 9 pairs, pinnule-free tips

very long. Prostomial ocelli prominently seen through collar.

Unserrated notch of bayonet chaetae very short

ech a cieciines ote senrancocessnctitele S. nudicrista sp. nov. p.76

Tube mustard coloured, circular in cross-section. Operculum

present. Maximum number of radioles 5 pairs, pinnule-free tips

short. Prostomial ocelli not seen through collar. Unserrated

notch of bayonet chaetae moderately long

ec acpia ia ccgneen'nissie sen vieidtie se cw wins S. snellii sp. nov. p.84

Dorsal ridge serrated, ventral ridge Y-shaped (Fig. 3,G) ... 8

Dorsal ridge unserrated, ventral ridge having other forms ... 9

Thoracic uncini without lateral tubercles. Maximum number of

radioles 7 pairs, abdominal segments more than 100, about

PSR ee eta ales vale els incite oe os soleinniiclg ah Sesion S. ypsilon sp. nov. p.56

Thoracic uncini with lateral tubercles. Maximum number of

radioles 11 pairs, abdominal segments less than 100, about 90

S. paraypsilon sp. nov. p.60

Wentral mdsennsenmated (Rig: 3.) s....cine- te eaidereveagesseens 10

Ventral ridge serrated (Fig. 3, J-0)

Tube white to faintly pinkish, circular in cross- section, external

longitudinal ridges absent; dorsal and ventral internal longitudi-

nal ridges pink, wedge-shaped in cross-section (Fig. 3, H).

Operculum with up to about 21 triangular radial lobes, cuticle

unthickened. Maximum number of radioles 5 pairs, abdominal

segments below 100 (about 70); bayonet chaetae with several

teeth on the basal boss ............... S. sumbensis sp. nov. p.82

Tube bright rose, quadrilateral to trapezoidal in cross-section, a

pair of external longitudinal ridges, and a faint median one in

places; dorsal and ventral internal longitudinal ridges white,

dorsal ridge T-shaped in cross-section, ventral ridge very small

(Fig. 3, I), may or may not be present. Operculum with up to

about 12 rounded radial lobes, cuticle unthickened. Maximum

number of radioles 14 pairs, abdominal segments over 100

(about 120); bayonet chaetae with two teeth on the basal boss

PRP ctiaciec Yoccadentttodch denacetctes S. iugoconvexa sp. nov. p.82

Bayonet chaetae with long blades and several teeth on the basal

LOTS: © RE GETeRRARB ERR ONEBMRO ee eaee ae Uneere tere S. vasseuri sp. nov. p.78

Bayonet chaetae with short to moderately long blades and few

@=Miteethion the basal’bOss ........0......secnseoeseecosecuensees 12

Accessory lateral ridges present (Fig. 3, K & L) .............. 13

Accessory lateral ridges absent (Fig. 3, M—O) ................. 15

Accessory ridges dorso-lateral. Dorsal ridge wedge- to

Y-shaped in cross-section (Fig. 3, K), tube white

Renee aes PE oS onan ease asec ans cca ase(lsi S. plaiae sp. nov. p.67

Accessory ridges lateral. Dorsal ridge a simple plate, at most

wedge to faintly T-shaped in cross-section (Fig. 3, L), tube pink

or with pink longitudinal bands ...................:seceeeeeeeeeeees 14

Bayonet chaetae dagger-shaped, with short blunt blades. Oper-

culum absent. Maximum number of radioles 6 pairs

Fee eee tet enc nce ter stoessterageecs S. caribensis sp. nov. p.68

Bayonet chaetae with moderately long blades and tapering tips.

Operculum present. Maximum number of radioles 5 pairs;

weet ah ateteca's deat dete cee S. lineatuba (Straughan, 1967) p.91

15. Tube with transparent discs attached to the wall externally and

mtemally (Eig. 35M)! .ccz.ss0.-ccs.te ee S. discifera sp. nov. p.94

ube withoutisuch dises'(Fig.3;,N’&,O)) foe .cce.-teneer> seer 16

1657 Opercolumtpresentir........-0e---8- S. karpatensis sp. nov. p.64

Operculiimabseutt site a bx. suasaaen tess ctoreepeece sence 17

17. Bayonet chaetae with 5-7 teeth on basal boss; maximum

number of radioles 4 pairs, abdominal segments about 55

rd Soe Sena MER ae cece seek evap deaseet ete S. zibrowii sp. nov. p.67

Bayonet chaetae with 3-4 (rarely 5) teeth on basal boss;

maximum number of radioles 6 pairs, abdominal segments

ADOUL SO oe oats chretys se decnlan oaSau te ty esos S. minuta sp. nov. p94

Although the ITS are very distinctive, the states of the

characters need to be used with caution. In two species S.

lineatuba (Straughan, 1967) and S. caribensis sp. nov., for

example, 25-40 tube fragments had to be examined before

the full extent of the development of dorsal and accessory

ridges could be established; the latter are missing in most

cross-sections. The shape of the distinctive inverted V, as in

the dorsal ITS of S. singularis sp. nov. is only found in a small

section in the earlier formed part of the tube; elsewhere, the

ridge is a smooth plate only. Along this ridge, the rounded

edge gradually becomes indented, gutter-shaped, and finally

widening to form a V. This would apply to certain other

characters as well. In S. massiliensis (Zibrowius, 1968) part of

the sample from Marseille was operculate and part had

rudimentary opercula only. However, all the specimens from

a large sample from Portman had rudimentary opercula only.

It may thus be expected that species which, on the basis of

relatively few specimens, have been described as non-

operculate, may turn out to be operculate when more mate-

rial becomes available. As another example, two samples

from Indonesia and Lizard Island (Queensland) initially

appeared to belong to two distinct species, on the basis of

differences in six character states. Additional material, how-

ever, yielded specimens with a full range of intermediate

states, showing that they belong to one and the same species.

DESCRIPTION OF SPECIES

Spiraserpula massiliensis (Zibrowius, 1968)

(Figs.4, A-O; 3, D; PI.2, A-D)

SYNONYMY. Serpula massiliensis Zibrowius, 1968: 102-105,

Pl.1, figs.24-37; Pl.14, fig.d.

Serpula massiliensis: Bianchi, 1981 : pp.51-S2, fig.16. Serpula

massiliensis: ten Hove & Aarts, 1986: 35 [not the tropical E.

Atlantic record, see S. ypsilon].

MATERIAL EXAMINED. Unless otherwise mentioned, the

material was collected and/or determined by Zibrowius.

Mediterranean:

France: Marseille: 1. Anse des Cuivres; below SME, over-

hang 6m, 21.vii.1987 (10 out of several specimens, BMNH

ZB 1989, 43-53). 2. fle Plane; submarine cave, 6m, legit G.

Harmelin, vi.1987 (4 specimens, SME). 3. fle Plane; 1987 (5

out of several specimens, BMNH 1989 101-150).4. fle Plane;

50 T.G. PILLAI AND H.A. TEN HOVE

)

0.01mm| /j

M N O

Fig. 4 Spiraserpula massiliensis (Zibrowius, 1968). A-O, From Marseille, Anse des Cuivres, BM(NH). ZB1989.43-53. A, Aggregation of

tubes with fractured ends showing the serrated dorsal ridge along the convex inner wall, and granular overlay. B—C, Erect parts of tube

with four-lobed peristome. D, Anterior part of operculate worm. E, Same specimen showing end of thoracic membrane. F, Thorax with

pair of prostomial ocellar clusters, also enlarged. G-J, Four bayonet chaetae from the same fascicle. K, Row of thoracic uncini. L, Anterior

abdominal uncini. M & N, Middle abdominal uncini. O, Posterior abdominal uncini.

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961

submarine cave, 6m, legit G. Harmelin, vii.1971 (7 speci-

mens, ZMA V. Pol.3159). 5. Friocil Harbour (5 out of

several specimens, BMNH ZB 1989 54-100). 6. Grand Con-

glu; 1987 (5 out of several specimens, BMNH ZB 1989

151-200). 7. Martigues; ca. 50km W of Marseille, Ponteau

Electric Plant, under stones, 1.3m, 5.iv.1977 (2 out of several

specimens, SME). 8. La Ciotat; ca 30km E. of Marseille, Bec

de l’Aigle, on concretions of sand, 40m, 111.1970 (3 speci-

mens, SME). 9. Canyon de la Cassidaigne; about 20km E of

Marseille, off Cassis, from 170-270m by dredging, 15.vi.1974

(tubes, 1 specimen, SME).

NW Corsica: 10. Revellata, 15m, calcareous algal masses,

8.iv.1978 (2 specimens, SME).

Italy: 11. S coast of Sorrento peninsula, ‘Grotto Zaffiro’,

10m, 29.v.1974 (3 specimens, SME). 12. Bari; 10m, cave,

legit T.M.Griessinger, 8.vii.1968 (5 specimens, SME).

Greece: 13. Gulf of Corinth, Aspra Spitia, 5m, 26.ix.1977 (3

specimens, SME).

Malta: 14. Oxford University Underwater Exploration Group

1965, scrapings from roof of cave, det. Pillai (2 specimens,

BMNH ZB1989 32-36).

Tunisia: 15. Tabarka; algal concretions, 31—36m, 24.iv.1969

(1 specimen, SME). 16. Zembra Islands, concretions, 35m,

30.iv.1969 (few tubes with portions of worms, SME). 17.

‘Dauphin’ Stn.24, 35°12’N 11°25’E, 73m, on Arca, legit

Bane, Medit. Mar. Sorting Center, 28.viii.1967 (1 specimen,

SME). 18. Gulf of Gabes; ‘Calypso’, 34°05’N 10°48’E, 23m,

muddy sand with Caulerpa meadow, on shelly material,

20.iv.1965 (1 specimen, SME1887). 19. Gulf of Gabes;

‘Calypso’, 34°13’N 10°31.9'E, 31m, Caulerpa meadow,

27.1v.1965 (1 specimen, SME 1910).

SE Spain: 20. Cabo de Palos; ca S50km E of Cartagena, 6m,

legit A. Ramos, 4.iv.1982 (3 specimens, SME). 21. Portman;

20km E of Cartagena, small overhang, 0.5—1.0m, on rock

covered by dark brown sediment, the latter retained on the

tube surfaces by oil pollution, 5.iv.1984 (20 out of several

specimens, SME).

Portugal: 22. From a submarine cave near Sagres, Algarve,

legit H. Zibrowius Sept. 1986 (BM(NH) 1992. 181-255).

NE Atlantic: 23. Gorringe Bank; ‘Meteor’ M9c, Stn.95, AT

29, 36°29.9'N 11°33.0’W, 150-430m, 24.vi.1967 (some empty

tubes, SME). 24. Madeira Archipelago; Jean Charcot,

Stn.42, SW of Porto Santo, approx. 33°0.4'N 16°24.5’W,

125-145m, 17.vii.1966 (empty tubes, information pers.

comm. H.Zibrowius). 25. Canary Islands; W coast of Palma,

Tijarafe, 28°42’N 17°58'W, 20m, CANCAP 4.D14, det. M.

Aarts (5 out of several specimens, RMNH 18465, ZMA

V.Pol.3739, BM(NH) 175-180). 26. NW Africa; off Point

Elbow, ex Spanish Sahara, ‘Tenace’ D16, Stn. 23, 24°13'N

16°17’'W, 50-60m, legit Marche-Marchard, 13.iv.1967 (4

specimens, SME).

TYPE LOCALITY. Marseille (France).

DESCRIPTION. In order to follow the variations within the

genus Spiraserpula it was considered useful to have as com-

plete a description as possible of one member of the group, S.

massiliensis was selected because of the large amount of

material available from various sources. The following

updated species description is based on the original account

as well as additional data obtained from a study of the above

collections, which include much of Zibrowius’ original mate-

rial.

According to Zibrowius (1968), the tubes are white, circu-

lar in cross-section and, although difficult to measure because

of their coiling, may attain a length of 50.0 mm for a diameter

of about 0.5—1.0 mm. Their coiling is highly irregular and the

direction may reverse. Sometimes many tubes are joined

together, coiling in the same direction. They are relatively

thick, except in their erect portions which are cylindrical. At

intervals along the latter, there may be one to a few out-

wardly directed expansions, generally referred to as peris-

tomes. They are sometimes in the form of four, somewhat

symmetrically placed lobes. In dense populations, the erect

tubes may form a kind of uniform meadow in submarine

caves of the Mediterranean (Zibrowius, pers. comm.). The

surface of the tube is covered by faint granulations which,

very rarely, may form short longitudinal ridges.

It is difficult to remove the worms from their tubes. When

removed, quite a number of specimens lacked their radiolar

crowns. An operculum may be present or absent. When

absent there is a rudimentary operculum on each side. When

present, the operculum is small, and its diameter does not

correspond with that of the tube. Its distal end is flattened to

slightly funnel-shaped, bearing 13 to 23 obtuse lobes. The

peduncle is more slender than the pinnulate radioles, and its

attachment to the operculum is central and constricted. The

corresponding radiole of the opposite side is reduced to a

filamentous rudimentary operculum, about one-third the

length of the radioles, and lacks pinnules.

The collar consists of a large ventro-median lobe and a

smaller one on either side of it, all of which are rounded. The

thoracic membranes are broad and well developed up to

about the fourth thoracic chaetiger, and reduced posteriorly.

They are not united to form a post-thoracic ventral flap or

apron. The number of thoracic segments may exceed the

usual seven found in many other species of Serpula. Each

collar fascicle generally possesses four bayonet-shaped

chaetae and a similar number of simple bladed chaetae. Each

bayonet chaeta has a striated blade distally, and several teeth

on the basal boss. Thoracic uncini bear 3—5 teeth. Anterior

abdominal bundles consist of 2-3 flat trumpet shaped

chaetae. Uncini possess 2-5 teeth in a single row. The

posterior abdominal segments bear long capillary chaetae,

and rasp-shaped uncini with several rows of teeth.

Additional data obtained during the present study are as

follows:

TUBES: White to faintly creamish, and may occur in closely

intertwined masses of a few to several individuals (Fig. 4, A);

sometimes solitary. Except for their free erect portions, they

are mutually bonded to various extents, particularly at their

bases. Their ‘granular overlay’ is shown in Fig. 4 A, and the

four-lobed peristomes in Fig. 4, B & C.

An important character which had not been reported

relates to the tube, which bears ITS. In its first formed

portion, which is normally coiled, there is a serrated longitu-

dinal ridge. Careful removal of numerous specimens from

their tubes has shown that this serrated ridge is always on the

convex side of the coils, as also found in masses consisting of

several individuals (Fig. 4, A). The orientation of the worm

within the tube is such that the posterior dorsal part of its

abdomen is always applied to this serrated ‘dorsal ridge’

(Fig. 3, D). This, in addition to its tight coiling, accounts for

the difficulties encountered in extracting complete worms

from their tubes by Zibrowius (1968) and in the present

study.

The numerous specimens in the collection from Portman

show an apparent exception in lacking ITS. However, confir-

52 T.G. PILLAI AND H.A. TEN HOVE

Se a :

fi 6 oa Cd, :

Renmei Oe

» ogee?

BSc

0.01imm

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961

mation that the specimens belong to this species came from a

very small number of old and empty tubes at the bottom of

the aggregations having the characteristic serrated internal

dorsal ridge. The population had been subjected to oil

pollution, as evidenced by a thick deposit of it which covered

the tubes externally, and even lined them thinly internally.

WORMS: The longest operculate specimen (from fle Plane,

Marseille, BMNH ZB 1989 101-150) is 19.0 mm long; its

thoracic width 1.1 mm. It has 5 pairs of radioles, in addition

to an operculum on one side and a rudimentary operculum on

the other. It also has the longest operculum and peduncle,

3.5 mm; its operculum is 1.0 mm long, 1.2 mm in diameter,

bell-shaped, and with 11 rounded marginal lobes. The abdo-

men is 13.5 mm long and has 51 segments, the last four with

capillaries. The longest non-operculate specimen, from the

same locality, has a total length of 20.0 mm. Its abdomen is

15.5 mm long and has 43 segments, the last ten with capillar-

ies. The two specimens indicate variations in abdominal

length due to extent of contraction during fixation, and that

the number of segments in the abdomen may not be always

proportional to its length. In the other specimens studied, the

length of the operculum together with the stalk ranges from

1.0-2.0 mm; the operculum from 0.5—0.6 mm in length and

0.3-1.5 mm in diameter.

The shape of the operculum varies from an elongate funnel

in the majority of cases to a narrow bell in the others (Fig. 4,

D, E).The opercular radii end in 10-16 rounded marginal

lobes. The width of the peduncle, just before the constriction,

is 3/Sths to 4/Sths that of the proximal end of the operculum.

The number of radioles observed is 4-6 pairs, and their

pinnule-free tips vary in length from 1/Sth to2/3rd the total

length of the radiole (i.e. radiole plus pinnule-free filament).

Zibrowius (1968) reported a higher maximum number of up

to 23 marginal lobes on the operculum. This high number, in

one specimen, is apparently not representative of the species

(Zibrowius, pers. comm.).

A total of 67 specimens from different localities (including

21 from the abnormal Portman material, vide below) were

examined for various characters. All the Portman specimens

possess only rudimentary opercula. Out of 46 specimens from

the other collections 25 possess an operculum, 12 lack one,

and the rest are indeterminate since they lack radiolar

crowns. The majority of specimens from normal populations,

therefore, possess an operculum.

Another character, hitherto not reported, is the presence

of two light red to reddish-brown clusters of prostomial ocelli

(Fig. 4, F). They can be seen when a worm with its radioles

removed is viewed from the anterior end, or through the

collar in mounted specimens. Each ocellus consists of a

pigmented cup-shaped part, and a transparent anteriorly or

antero-laterally directed lens-shaped part within it (Fig. 4, F).

Thoracic glands, as found in other species of the genus, are

absent.

The numbers of thoracic chaetal tufts, 6 to 9 on each side,

may be symmetrical or asymmetrical. The condition in 26

Table 1 Spiraserpula massiliensis (Zibrowius). Number of thoracic

chaetal tufts on each side.

No. of individuals (n=26) 1 4 jj 11 3

No. of thoracic chaetal tufts 6/7 TI SM ASISm 8/9

Table 2 Spiraserpula massiliensis (Zibrowius). Extent of the

thoracic membranes of the two sides.

No. of individuals (n=21) 1 2 10 2 3 3

Thoracic membrane ends 4/5 4/6 =5/5 5/4 «66/6 6/7

specimens is summarized in Table 1. Likewise, the thoracic

membranes may end symmetrically or asymmetrically, but

always anterior to the last thoracic chaetiger; an apron is,

therefore, absent (Fig. 4, E). The condition in 21 specimens

is given in Table 2.

Collar fascicles bear 3 to 5 bayonet chaetae each. Bayonet

chaetae consist of a long serrated blade, an unserrated notch

of moderate length, and a basal boss with several teeth of

variable size (Fig. 3, G—J; Pl. 2, A). Thoracic and anterior

abdominal uncini usually have 4 or 5 teeth arranged in a

single row (edge saw-shaped), (Fig. 4, K, L; Pl.2, B & C). In

the intermediate abdominal region, the edge of each uncinus

is saw-shaped anteriorly whereas several teeth are placed side

by side (edge rasp-shaped) posteriorly. The number of teeth

in a single row decreases and the rasp-shaped posterior

portion increases as the posterior end of the abdomen is

reached (Fig. 4, M—O). Although the posterior abdominal

uncini are rasp-shaped, they have a single large tooth anteri-

orly (Fig. 4, O).

LIVE MATERIAL. (Vide Zibrowius, 1968)

HABITAT AND DISTRIBUTION. This species is commonly

found in submarine caves and at depths accessible by diving

(Zibrowius, 1968). The original description mentioned a

depth of 10-22m, but subsequently the species was found to

occur in shallower and deeper water (see list of material

examined). Deeper water collections came from depths of

31-36m (Tunisia) and 58-60m (off Point Elbow, Western

Sahara), the latter consisting of operculate and non-

operculate specimens. Empty tubes of this and other serpulid

species typical of shallow water have been obtained along the

the steep slope of the Gorringe Bank at 150-430m. This

occurrence may be due to slumping from shallower depths

(Zibrowius, pers. comm.). The Madeira Archipelago mate-

rial (125—145m) also consisted of dead material. The empty

tubes and single specimen from Canyon de la Cassidaigne

(170-200m) is also exceptional. In general, therefore, the

species commonly occurs in depths to about 60m, rarely down

to about 200m.

S. massiliensis is common in the Mediterranean (Greece,

Fig. 5 Spiraserpula capeverdensis sp. nov. A—-P, From type locality (SW of Sao Vicente), CANCAP 6.148 & 6.146; A-L, From 6.148, M-P,

from 6.146. A, Opened tube showing serrated internal dorsal ridge along the convex wall of coiled part; and granular overlay in places. B,

Coiled part of tube with fine transverse growth wrinkles externally, and its fractured end showing dorsal ridge on convex inner wall, and

two ventro-lateral longitudinal rows of smooth tubercles on opposite wall. C & D, Cross-section of two tubes, both with serrated internal

dorsal ridge, and one with ventro-lateral rows of tubercles. E-G, Three views of same worm showing rudimentary opercula (F), condition

of the thoracic membranes (G), and dorsal longitudinal groove (E & G), and ventral abdominal groove posteriorly (E). H, Anterior end of

younger specimen. I-L, Bayonet collar chaetae, all from same fascicle. M, Thoracic uncini. N, Anterior abdominal uncini. O, Uncini from

transitional region of abdomen. P, Posterior abdominal uncini.

Italy, France, Spain, Malta, and Tunisia). In the eastern

Atlantic it is abundant in submarine caves, and has been

recognized on Gorringe Bank, the Madeira Archipelago,

Portugal and the coast of Sahara.

S. massiliensis has been erroneously reported from the Red

Sea (Amoureux et al., 1978). Examination of the specimens

(HUJ) showed that their tubes lack ITS and they do not,

therefore, belong to the genus Spiraserpula.

Spiraserpula capeverdensis sp. nov.

(Figs.5, A—P; 3, E)

MATERIAL EXAMINED.

Cape Verde Islands: All CANCAP stations. Off Sao Vicente:

1. 6.134; 110-120m, (2 PARATYPES and some empty tubes,

RMNH_ 18197). 2. 6.135; 110-150m, (1 PARATYPE,

BM(NH) 1992.8). 3. 6.137; 75-90m, (1 PARATYPE,

BM(NH) 1992.9). 4. 6.146; 75m, (1 specimen, BMNH). 5.

6.148; 100-200m, (HOLOTYPE, 2 PARATYPES & 3 empty

tubes (residual material) ZMA VPol. 3651). 6. 6.166;

78-85m, (1 PARATYPE, USNM 130995). Off Razo: 7

.7.117, 100-120m, (some empty tubes, RMNH 18198). 8.

7.123; 120m, (5 specimens, RMNH 18199, ZMA V.Pol.3733.

Scuba diving station : Boa Vista: 9. 7.D06; down to 12m, (3

questionable specimens, ZMA V.Pol. 3871).

TYPE LOCALITY. Cape Verde Islands, Sao Vicente.

DESCRIPTION.

TUBES: White, nearly circular in cross-section, and occurring

in aggregations of a few individuals, occasionally solitary.

They are closely coiled amongst or upon themselves (Fig. 5,

A), and mutually bonded by a granular overlay. Erect

portions, when present, are very short, hardly rising above

the rest of the tube, and may end in four lobes. Faint growth

rings are sometimes present (Fig.5, B), and anterior

uncoiled portions may sometimes show a few transverse

thickenings, representing peristomes. In their first formed

parts, they possess an internal serrated dorsal ridge (Fig. 5,

A, D) and, often, a short ventro-lateral longitudinal row of

small smooth knob-shaped processes on each side (Figs.5, B,

C; 3, E). A mid-dorsal longitudinal groove in the posterior

part of the abdomen (Fig. 5, E, G) is applied to the serrated

dorsal ridge when the worm is withdrawn into the tube. The

maximum external diameter of the tube varies from 0.6 mm

in a juvenile to 1.4 mm in older specimens.

WORMS: (Fig. 5, E-H). An operculum is absent in all the

specimens examined. Instead, a filamentous rudimentary

operculum is present on each side. The number of radioles in

the larger specimens is often 7 or 8 per side, 4 in the smallest.

They are about 2.0 mm long in the larger specimens, and

have transverse specks at intervals. Their pinnule-free tips

are slender, 1/5 to 1/6 the total length of the radioles. Two

T.G. PILLAI AND H.A. TEN HOVE .

Table 3 S. capeverdensis sp. nov. Measurements and meristic data.

Stn.No. TL Width No. of Length No. of | Caps.

(mm) of radiol. of abdom. on

thorax abdom. segs.

(mm) (mm)

6.137 Ay) AVES) 8/8 PPL 138 ay

6.148 20.6 0.5 7/8 is) 145 =

6.148 Tale Os 7/8 6.9 96 12

7.123 229) 08 4/4 2.6 49 -

Tel23 2.4 0.3 4/4 ip) 29 9

Table 4 S. capeverdensis sp. nov. Numbers of thoracic chaetal tufts

and extent of thoracic membranes.

No. examined (n=12) 2 1 2 1 aie * 2

No. of thoracic chaetal tufts 9/8 8/8 8/7 8/5 7/7 7/6

No. examined (n=9) i 4 lee

Thoracic membrane ends 6/6 5/5 S/? 4/4

clusters of reddish to reddish-brown prostomial ocelli are

present.

Measurements and other data from the two longest and

three juvenile worms are presented in Table 3. The numbers

of thoracic chaetal tufts and the extent of the thoracic

membranes on the two sides is variable, as shown in Table 4.

The thoracic membranes do not extend to the last thoracic

chaetigers (Fig. 5, F), and apparently end symmetrically, but

further study of additional material is necessary for confirma-

tion of the latter. Ventral thoracic glands are absent.

Each collar fascicle bears up to about 5 bayonet chaetae

(Fig. 5, I-L). They have a long serrated blade, a short

unserrated notch and several moderately large teeth on the

basal boss. Thoracic uncini (Fig. 5, M) usually have 4 teeth in

a single row. Anterior abdominal uncini are similar, with 4-6

teeth (Fig. 5, N). The posterior abdominal uncini are rasp-

shaped, except for the single anterior tooth (Fig. 5, P). There

is a transition (Fig. 5, O) between the condition found in the

anterior and posterior abdominal uncini.

The differences between S. capeverdensis sp. nov. and S.

massiliensis are as follows: The former has only rudimentary

opercula, and higher maximum numbers of radioles (8 pairs)

and abdominal segments (145). Its tubes do not form tall

erect portions, and usually possess two ventro-lateral rows of

knob-shaped tubercles internally, in addition to the serrated

dorsal ridge. In S. massiliensis, however, an operculum may

or may not be present, the maximum number of radioles is 6

per side, and of abdominal segments observed 51. There is

also strong indication of an ecological difference (see below).

Fig. 6 Spiraserpula ypsilon sp. nov. From type locality material (SW coast of Island of Brava), CANCAP 6.D03. A, Aggregation of

fractured tubes showing ITS, consisting of a Y-shaped ventral ridge along the concave wall and a serrated dorsal ridge along the convex

wall. An oblique section (bottom right) shows the tapering anterior end of the ventral ridge. B, Erect portion of tube showing four-lobed

peristome. C-E, Different views of complete worm showing showing rudimentary opercula, pinnule-free tips of the radioles (E), and dorsal

and ventral longitudinal abdominal grooves. F-H, Same anterior end showing four-lobed collar (H), thoracic chaetigers and membrane (F),

and ventral longitudinal groove. I-K, Three views of larger specimen, showing dorsal and ventral longitudinal abdominal grooves. Note the

longitudinal cord-shaped structure within the ventral groove of the abdomen which fits into the gutter-shaped part of the Y-shaped ventral

ridge of the tube. L, Anterior portion of of worm accidentally fixed outside its tube, showing filamentous rudimentary opercula and

thoracic membranes. M-O, Three views of anterior part of another worm fixed outside its tube. Its thorax is considerably wider than those

of specimens fixed within their tubes, and the longitudinal grooves may be stretched and shallow (N).

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961

REMARKS. Although the few damaged specimens from Stn.

7.D06 are very similar to S. capeverdensis in most respects,

only 2 tube parts (out of 10 recognizable fragments) showed a

serrated dorsal ridge and possibly latero-ventral knobs. As

opposed to all material of S. capeverdensis studied so far, a

specimen in a tube without visible internal structures showed

a bell-shaped operculum with 13 radii. Therefore the identifi-

cation of this lot is left at ? capeverdensis.

LIVE MATERIAL. No records.

ETYMOLOGY. Named after the type locality.

HABITAT AND DISTRIBUTION. S. capeverdensis sp. nov. is

known only from the Cape Verde Islands, mainly from

depths of 75—200m where the bottom consists of various

combinations of coarse sand, shell gravel, calcareous stones,

calcareous nodules, calcareous algae and sponges, on which it

occurs among the epifauna.

Scuba diving to 15—20m, in a total of 28 different stations

off the Cape Verde Islands during CANCAP-VI and CAN-

CAP-VII, did not yield this species from the shallower

coastal waters. However, the dives yielded a different spe-

cies, S. ypsilon sp. nov., from these depths which, in the

Mediterranean, are typical for S. massiliensis.

Spiraserpula ypsilon sp. nov.

Figs. 6, A-K; 7, A-T; 34, G; Pl. 1, B)

SYNONYMY. Serpula massiliensis: ten Hove & Aarts, 1986:

35 (tropical E. Atlantic record only).

MATERIAL EXAMINED.

Cape Verde Islands: CANCAP stations. Scuba diving sta-

tions: 1. 6.D01; S coast of Sao Tiago, SE of Porto Praia, 15m

(1 specimen, RMNH 18177). 2. 6.D02; S coast of Sao Tiago,

Baia de Santa Clara, 20m, caves in rock (2 out of several

specimens; RMNH 18187; BM(NH) 1992.85-115; FSBC I

39197 (1); AM W 20339 (1); NSMT (1)). 3. 6.D03; SW coast

of Brava, Porto dos Ferreiros, 15m (30 specimens: HOLO-

TYPE & 5 PARATYPES, RMNH 18176. Other

PARATYPES: ZMA V. Pol. 3650 (10); USNM 130993 (6)

and BM(NH) 1992.73-82 (10)). 4. 6.D06; SW coast of Sao

Nicolau, Baia do Tarrafal, 15m (4 specimens, RMNH 18188).

5. 6.D10; S. coast of Sao Vicente, 15m (5 out of several

specimens, RMNH 18189). Coastal stations: 6. 6.K13; SW

coast of Ilha Razo, (14 out of several specimens, RMNH

18190, ZMA V.Pol.3726, USNM). 7. 6.K15; SW coast of Ilha

de Santa Luzia, (1 out of several specimens; bulk RMNH

18191; clusters of 10-15 tubes each BM(NH), 1992.116—120;

ZMA V.Pol.3727; HUJ; Dr M. Jager). 8. 6.K21; NE coast of

Sao Vicente, Baia das Gatas, (3 specimens, RMNH 18192).

Scuba diving stations: 9. 7.D03; Cima, SE coast, (1 specimen,

RMNH 18193). 10. 7.D05; Maio, SW coast of Ponta Preta (2

out of few specimens, RMNH 18194). 11. 7.D06; Boa Vista,

Ilhéu de Sal Rei, 12m (1 out of few specimens, ZMA

V.Pol.3728). 12. 7.D10; Razo, S coast, 20m (1 out of few

specimens RMNH 18195; BM(NH) 1992. 121-131; ZMH).

Dredging station:13. 6.148: off Sao Vicente, 100—200m (1

empty eroded tube; RMNH 18196). Tropical Western Atlan-

tic, Gulf of Mexico: 14. Florida, Stn. EJ66—460, 26°24'N

T.G. PILLAI AND H.A. TEN HOVE

82°28’W, 18m, 6.xii.1966, ‘Hourglass’ Stn J, (20 out of

several specimens, FSBC I, ZMA V.Pol.3729, BM(NH)

1992. 132-147). 15. Florida: Stn. EJ 67-76, 27°37'N 83°28'W,

39m, 2.111.1967, ‘Hourglass’ Stn.C, (few specimens, FSBC I,

ZMA_ V.Pol1.3730. 16. Florida: Stn. EJ67-328, 27°37N

83°07W, 18m, 11.ix.1967, ‘Hourglass’ Stn. B (4 out of several

specimens, ZMA V.Pol.3731). Caribbean: 17. Aruba: Andi-

curi, cape W of beach, windward side, rockpool, exuberant

coral growth, strong wave action, 0.5m, legit H.A. ten Hove,

28.vili. 1970, Stn.2034B (together with S. caribensis sp. nov.;

ZMA V.Pol.3732). 18. Colombia: Santa Marta area, Cabo

and Ojo del Aguja, 8-27m, legit J. W. Dulfer and M. J. C.

Rozenmeyer 1986, ident. as S. massiliensis (1 damaged

specimen, tube; ZMA V.Pol.3778). Bermuda: 19. Stn.14,

legit Reed, with a note by Zibrowius in 1970 indicating that it

is a new species (3 specimens, USNM 43244).

TYPE LOCALITY. Cape Verde Islands, Brava.

DESCRIPTION.

TUBES: Faintly pinkish, often with a more pronounced shade

of light pink or light mauve in the granular overlay towards

their anterior ends. They normally occur in mutually bonded

highly coiled aggregations, occasionally also singly, adjacent

to the aggregations. The granular overlay is fine, somewhat

translucent and nearly uniform (partly shown in the top left

portion of Fig. 6, A). External longitudinal ridges are nor-

mally absent, but up to three may be faintly developed on the

less coiled tubes of solitary specimens. Their anterior por-

tions are generally attached, often with their lateral borders

extending somewhat over the substratum. Occasionally, they

possess erect ends which attain an external diameter of up to

1.25mm, and may bear a few peristomes which are usually

four-lobed and outwardly directed (Fig. 6, B).

The ITS are more complicated than those of the other

known species of the genus, with the exception of the closely

related species S. paraypsilon sp. nov. As seen in carefully

opened tubes or through their fractured ends in an aggrega-

tion (Fig. 6, A), they consist of a serrated dorsal ridge, and a

thin, very fragile, Y-shaped ventral ridge (Fig. 3, G; Pl.1, B).

The gutter-shaped part and the stem of the latter gradually

decrease anteriorly until they are represented only by a

simple ventral ridge, which itself decreases in height and

gradually disappears (Fig. 6, A, bottom right corner). These

ridges commence in the first formed portions of the tube, but

usually extend more anteriorly than in most of the other

species of the genus. The inner translucent layer of the tube is

faintly pinkish, as is the Y-shaped ventral ridge. In addition,

the latter may possess one or two thin dark pink longitudinal

stripes on the outside of the gutter-shaped part, one along the

top and the other along the bottom.

WORMS: Some measurements and counts are provided in

Table 5-8.

The worms attain a total length of about 27.5mm, a

thoracic width of 0.6mm, a maximum of about 131 abdominal

segments, with capillaries on the last 10 segments or so. The

maximum number of radioles is 8 pairs. The two specimens

with 4 pairs of radioles are juveniles. The pinnule-free tips

are short to moderately long, up to about 1/5 the entire length

Fig. 7 Spiraserpula ypsilon sp. nov. A-0, From type locality, CANCAP 6.D03. P-T, From Florida Stn. EJ 66-460. A-E, Bayonet chaetae

from same fascicle. F-I, Same, from second specimen. J, Thoracic uncini. K, Uncini from first abdominal torus. L, Uncini from third

abdominal torus. M, Anterior abdominal uncini from another specimen. N, Uncini from mid-abdominal torus (transitional region). O,

Posterior abdominal uncini. P-T, Bayonet chaetae of one fascicle.

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961

\ \ \ \ \ ‘ [o- 01mm

\ \\ ‘i

\ NY \ SS BEESELSID

A Wh Bit L

uy GER AGH

———

2<V?/"

i's

ees

es SO

FES

——_ S 1S) o

es °

= EEU_UAAZZZ™" a?

0 ——_——E

SSS SSS

Table5 SS. ypsilon sp. nov. Measurements (mm) and counts.

Total length DiS S6) 14 EOS LO) 8426

Thoracic width - 0:6) 0:6) 4 04 05s 05 04

No. of radioles - 77 8/8 3/5 66/6 =~(6/6 =4/4

Abdominal length WS Sy MLK nS Se AEG ALS 2G

No. of abdom. segs. iss Gi Ah OSE So 3/

Capillaries from 103 80 124 99 24 44 30

Table 6 S. ypsilon sp. nov. Numbers of radioles.

No. of specimens (n=37) 1 6 15 13 2

No. of radioles (L/R) 8/8 77 6/6 £S/S 4/4

Table 7 S. ypsilon sp. nov. Numbers of thoracic chaetal tufts in 69

specimens.

No. of specim. oS oe le, 2G) 23003: Soe ll

Nos. of. tufts. 10/9 10/8 9/9 9/8 9/7 9/6 8/8 8/7 8/6 7/7 7/6 7/5

Table 8S. ypsilon sp. nov. Extent of thoracic membranes in 43

specimens.

Number of specimens 3) 31 6 1

Thor. membranes end on 6/6 6/5 5/4 5/3

of the radioles (Fig. 6, E). Live material from Stns. 7.D03

and 7.D05 showed a pair of pigmented ocelli at the base of

each radiole, externally. An operculum is absent; there is a

filamentous rudimentary operculumon each side (Fig. 6, E,

LO).

Two clusters of reddish to reddish-brown prostomial ocelli

are present. Although the width of the thorax ranges from

(0.40.6 mm in specimens preserved within their tubes, it can

be wider in anterior portions of worms accidentally preserved

outside their tubes (Fig. 6, F-H, L-O). The median lobe of

the collar is sub-rectangular, rounded laterally and with a

smooth mid-ventral notch, giving the entire collar a four-

lobed appearance (Fig. 6, F-H & L-O).

The numbers of thoracic chaetal tufts on the two sides

range from 5 to 10, and may be symmetrical or asymmetrical,

as are the endings of the thoracic membranes (Tables 7 & 8).

Paired thoracic glands are absent.

Collar fascicles bear up to about four fully developed

bayonet chaetae and two more being formed deep within the

fascicle, with a similar number of simple bladed chaetae.

Juveniles possess fewer, often two fully developed bayonets

and two more being formed within the fascicle. Each bayonet

chaeta has a long serrated blade, a short unserrated notch,

and few to several moderately large somewhat conical teeth

with smooth tips on its basal boss (Fig. 7, A-E, F-I). The

serrations of the blade are short towards its proximal part,

T.G. PILLAI AND H.A. TEN HOVE

Table 9S. ypsilon sp. nov. from Florida (EJ— 66-460).

Measurements and meristic data from two longest specimens out

of 22 measured.

Total Thoracic No. of Abdomen

length width radioles Length No. of Caps

(mm) (mm) (mm) segs. from

Specimen 1 22 0.6 7/7 18.5 110 97

Specimen 2 20.7 0.8 6/6 16.7 130) 11

Table 10 Meristic and other data on S. ypsilon sp. nov. from

Florida (EJ— 66-460).

No. of specimens 4 2 AD fl 324 i

(n = 14)

No. ofradioles 7/7 7/6 6/6 6/5 S/5 3/5

No. of specim. 1 1 LY 4 Gi} 9) ee ae 1

(n = 29)

No. of th. chaetal 10/9

tufts

No. of specimens 1 1 LO cael

(n = 20)

Thor.membrane 7/S 6/5 S/S S/4 4/4

ends

10/7 9/9” 9/8) S18 ST TA T6e eri

but somewhat pilose distally. Thoracic uncini are mostly with

6 teeth, but some have 4 or 5 (Fig. 7, J). Anteriorly there are

up to 4 flat trumpet chaetae in each bundle, posteriorly there

are 1 or two capillaries instead. Anterior abdominal uncini

usually have 4 or 5 teeth arranged in a single row (Fig. 7,

K-M). The posterior abdominal uncini are rasp-shaped, with

a single anterior tooth and several (4-6) rows of teeth

posterior to it (Fig. 7, O). In between, there is a progressive

reduction in the number of teeth in a single row, and a

corresponding increase in the rasp-shaped area (Fig. 7, N).

The special adaptations of the body of the worm in relation to

the internal structures of the tube are as follows: A narrow

longitudinal groove extends along the mid-dorsal line of the

abdomen and thorax (Fig. 6, C, D, H-K, O). The abdomen

and thorax are also grooved ventrally, and within this longitu-

dinal groove, forms a cord-shaped longitudinal ridge (Fig. 6,

C, D, L-L). The orientation of the worms within their highly

coiled tubes is such that the dorsal groove is applied to the

serrated dorsal ridge of the tube, and the cord-shaped ventral

abdominal ridge fits into the gutter-shaped part of the

Y-shaped ventral ridge of the tube. The latter, in turn, fits

into the ventral groove of the body.

COLLECTIONS FROM THE WESTERN ATLANTIC. S. ypsilon has

also been collected from Florida, Bermuda and Aruba.

Study of material from Florida (EJ— 66-460) provided the

following data: The external diameter of the tubes attains

1.1 mm. A granular overlay is present. The external coloura-

tion varies from faintly creamish to faintly pinkish. Their

internal colouration and structures are similar to those from

Fig. 8 Spiraserpula paraypsilon sp. nov. A, Tube from Curagao, NA, Cornelisbaai, showing granular overlay and longitudinal ridges. B-N,

From Klein Bonaire Stn. 2105A. B, Tube with indistinct longitudinal ridges. C-N, From holotype. C, Tube fragment showing serrated

dorsal ridge along convex wall. D, Tube fragment with Y-shaped ventral ridge along opposite wall. E & F, Adult worm; E, Radioles of

both sides, with very long pinnule-free tips and lacking rudimentary opercula. F, Body showing dorsal and ventral longitudinal abdominal

grooves and ventral pigment patches. G-L, Bayonet collar chaetae. M, Thoracic uncini of holotype showing lateral denticles. N, Anterior

abdominal uncini, with more prominent denticles.

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961

(

; =3°0

| =

Sitka

—~

KC)

Pouttsle sane

the Cape Verde Islands. The colouration of the worms in

alcohol shows a difference. The radioles and body have an

overall fleshy to dark reddish-brown colour, with blackish

pigment clusters ventro-laterally in the abdominal segments.

Whether this colour difference is due to different methods of

fixation or not needs to be verified. Maximum sizes encoun-

tered have been given in Table 9.

An operculum is absent, but a pair of rudimentary opercula

is present. Up to 10 thoracic chaetal tufts per side were

counted, and the extent of the thoracic membranes is vari-

able, however, never reaching the last thoracic chaetiger.

Meristic and other data are given in Table 10.

The collar chaetae (Fig.6, P-T), are larger than those of

the specimens from the type locality, but otherwise similar. In

the abdomen, up to 10 flat trumpet shaped chaetae per

fascicle were counted.

The other two Florida samples are similar. However, in

sample EJ 67-76, one specimen lacking radioles has 12

thoracic chaetigers on the left and 11 on the right, with the

thoracic membranes ending on the 6th and Sth chaetiger,

respectively. The samples from Bermuda and Aruba are

similar to those from Florida.

LIVE MATERIAL. There are some intra-specific colour varia-

tions inS. ypsilon sp. nov., as observed in collections from

different stations in the Cape Verde Islands:

Stn.6.D02: Branchial radioles distally orange, proximally

pinkish orange, except for a bright red spot where the

radioles meet. Thorax is transparently reddish, with two

subcutaneous brown spots laterally. Abdomen is pink or

orange, with brown sides.

Stn.7.D03: Distal half of the short radioles banded white

and yellowish orange, basal half bright red. Basal radiole

parts with oval lens-shaped structures, apparently ocelli.

However, after preservation no lenses could be found in this

material, not even after staining in methylene blue. Thorax

bright red, abdomen orange, with brownish-green granules

laterally.

Stn.7.D05: Radioles transparent, hyaline, with a single row

of pigmented spots at the base. No lenses visible, even with a

compound microscope. Thorax and abdomen orange-brown.

Sides of abdomen show brown granules (in four specimens).

In two other specimens the radioles are hyaline, with trans-

verse orange bands and red pigment spots at their bases. The

rest of the body is red, otherwise similar to the other four

specimens.

ETYMOLOGY. The specific name refers to the Y-shaped

internal ridge.

HABITAT AND DISTRIBUTION. As revealed by several scuba

dives and littoral surveys in the Cape Verde Islands, S.

ypsilon sp. nov. occurs on various hard substrata in depths

occupied by S. massiliensis (Zibrowius) in the Mediterra-

nean. At Stn. 6.D02, for instance, the ceiling of a half metre

deep cave, at a diving depth of 14m, was covered with crusts

of partially erect tubes of S. ypsilon. However, S. massiliensis

T.G. PILLAI AND H.A. TEN HOVE

did not turn up in any of the collections from the Cape Verde

Islands. The single, eroded tube of S. ypsilon sp. nov. from

the dredging station 6.148 (100-200 m) was probably trans-

ported down the slope.

Although the Western Atlantic and Cape Verde Islands

material show some differences, they appear inadequate to

separate them into distinct species. The Western Atlantic

range is from Bermuda to Aruba. A species from the

Caribbean which has similar internal tube structures, but

differs in other respects, is described next.

Spiraserpula paraypsilon sp. nov.

(Figs.8, A-N; 9, A-R; 3, G)

MATERIAL EXAMINED.

Bonaire (Neth. Ant.): 1. Klein Bonaire, N, half mile E of

Westpunt, reef, little sand, corals, 38 m, legit H.A. ten Hove,

1.vii.1970, Stn. 2105A, HOLOTYPE & PARATYPE 2,

ZMA V. Pol.3714; PARATYPE 1, BM(NH) 1992.156. 2.

Lac, dam, pool in wash of plunging breakers, Diploria,

Millepora, Porites, cobble in coarse sand, 50 cm, from corals,

legit JH AS” ten) Hove, #15. viiel970% este 2127

(PARATYPES 6-8, ZMA V.Pol.3717). 3. Plaja Frans, on

dead coral covered with calcareous algae, little sand,

1.0-1.5 m, legit H. A. ten Hove, 16.vii.1970, Stn. 2110A

(portions of tube, 1 incomplete worm, BM(NH) 1992.157). 4.

Karpata, steep reef, drop off and flat above, 15-4 m, from

living corals, legit H.A. ten Hove, 19.v.1987, Stn. 87-5

(PARATYPES 4-5, USNM 130992).

Curacao (Neth. Ant.): 5. Cornelisbaai, sandy reef, from the

underside of dead plate-shaped coral, 15 m, legit H. A. ten

Hove, 15.xi.1988 (PARATYPE 3 USNM 130991) and 17.1.

1990 (4 specimens, AM W20338). 6. Piscaderabaai, outer bay

in front of Carmabi, rubbish on reef, 10 m, legit H. A. ten

Hove, 10-12.1.1990 (19 specimens, ZMA _ V.Pol.3718,

BM(NH) 1992.158-165, FSBC I 39196. 7. Salinja Fuik, reef

in front, 20-25 m, legit H. A. ten Hove, 18.1.1990 (25

specimens, NSMT).

TYPE LOCALITY. Klein Bonaire, Curacao.

DESCRIPTION.

TUBES: White, flattened, and with a granular overlay. The

maximum external diameter of the tube of the holotype is

2.0 mm. A median and about 3 pairs of lateral longitudinal

ridges can be observed (Fig.8, A). In an empty tube from the

type locality with a diameter of 1.5 mm,the number of ridges

is less distinct (Fig.8, B). ITS, located within the first formed

coiled parts, are translucent white, and very similar to those

S. ypsilon. They consist of a serrated dorsal ridge along the

convex wall (Fig.8, C) and a Y-shaped ventral ridge along the

opposite side (Figs.8, D; 3, G). The serrations of the dorsal

ridge are pointed and directed somewhat posteriorly. Tubes

found on asbestos plates (Piscaderabaai) were clearly branch-

ing, as described in detail for S. caribensis (Fig.16, A & B).

WORMS: The total length of the holotype is 16.4 mm. With a

thoracic width of 0.8mm, it is stouter than S. ypsilon. The

Fig.9 Spiraserpula paraypsilon sp. nov. A-N, From Klein Bonaire Stn. 2105A; O-R, from Bonaire, Karpata Stn.87.5. A-J, From juvenile

paratype. A, Tube showing start of external longitudinal ridge and shallow transverse growth markings. B, Posterior tube fragment with

serrated dorsal ridge. C, Radioles with long pinnule-free tips, a rudimentary operculum on the left and none on the right. D & E, Two

views of body showing pigment patches in both, dorsal an ventral longitudinal abdominal grooves (D), and extent of thoracic membrane

(E). F-J, Bayonet chaetae, including one newly formed within the fascicle (J). K-N, From holotype. K & L, Tube fragments (K) and the

other from a more posterior coil, with serrated dorsal ridge. M & N, Two views of body showing pigment patches, dorsal and ventral

longitudinal abdominal grooves (M), and extent of thoracic membrane (N). O-R, Bayonet chaetae of one specimen.

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961

abdomen is 9.9mm long, with about 98 segments, and capil-

laries in the last 20.

The pinnule-free tips (Fig.8, E) are quite conspicuous and

much longer than those of S. ypsilon, being up to about

1.9 mm. They constitute nearly half to more than half the

length of the radioles (about 3.7 mm). The number of

radioles, 11 on each side, is higher than that of S. ypsilon

(maximum 8). They bear at intervals what appear to be

narrow, transverse, lenticular lacunae. Two pigmented pros-

tomial ocellar clusters are present.

The rest of the body (Fig.8, F) is similar to that of S. ypsilon

in many respects. In alcohol, the abdomen has an overall

pinkish colour, with clusters of reddish-brown pigmented

cells ventro-laterally.

Thoracic chaetigers number 7 on both sides. Thoracic

membranes end on the third thoracic chaetiger (second

uncinigerous segment) on both sides of the thorax. Paired

ventral thoracic glands were not seen.

A collar fascicle of the holotype has five fully formed

bayonet chaetae, and a developing one deep within. A

paratype from Karpata has 6 fully formed bayonets and one

newly forming one within the fascicle. Each bayonet chaeta

consists of a long, narrow serrated blade, and a considerably

expanded basal boss bearing several moderately large, some-

what pointed teeth (Fig.8, G—L). The serrations are short and

fine proximally, but longer and pilose distally. The unser-

rated notch may be very short, up to about twice the length of

the longest teeth, or almost lacking (Fig.8, I). The thoracic

and anterior abdominal uncini bear 5 teeth in a single row.

They differ from all the other known species of the subgenus

in possessing minute denticles on their sides (Fig.8, M,N).

The adaptations of the worm in relation to ITS are similar

to those of S. ypsilon. The dorsal longitudinal abdominal

groove is applied to the serrated dorsal ridge of the tube, and

the Y-shaped ventral ridge is enclosed within a ventral

abdominal groove. Within the latter, a cord-shaped abdomi-

nal ridge fits into the gutter-shaped part of the Y.

The paratype from the type locality is a juvenile. Its tube

(Fig.9, A) shows faint transverse grooves mainly, but the

beginnings of a granular overlay and longitudinal ridges can

also be seen. ITS and adaptations of the body are identical to

those of the holotype (Fig.9, B,K,L). Measurements and

counts of the worm are as follows: Length 6.2 mm, thoracic

width 0.5 mm, radiolar length 2.1 mm, pinnules 1.1 mm,

abdomen 4.2 mm, 53 segments, with capillaries on the last

10. The number of thoracic chaetigers, ending of the thoracic

membranes (Fig.9,E), and colour, are the same as in the

holotype.

The 7 pairs of radioles already approach the maximum

number in other material, except the holotype, and their long

pinnule-free tips are similar to those of the holotype (Fig.9,

C). However, there is a very short and slender rudimentary

operculum on one side, while it is lacking on the other (Fig.9,

C), indicating that both may become completely lost in older

specimens (holotype). A similar condition is found in one of

the specimens from Karpata (below).

Collar fascicle with four fully formed bayonet chaetae

(Fig.9, F-I) and a developing one deep within (Fig.9, J).

Their basal bosses are not as expanded as in the holotype and

the blades are shorter. The uncini are similar to those of the

holotype.

The tubes of the three specimens from Karpata agree with

those from the type locality in being white externally, pinkish

internally,and bearing the Y-shaped ventral ridge and ser-

T.G. PILLAI AND H.A. TEN HOVE

rated dorsal ridge. The serrations of the latter bear posteri-

orly directed tapered tips (Fig.9, K,L). The radioles of all

specimens are detached, highly contracted, and do not clearly

show the extent of the pinnule-free tips. One crown has a

short rudimentary operculum on each side, the second has a

rudimentary operculum on one side but lacks it on the other,

and the third half crown has a rudimentary operculum which

is very reduced and filamentous. The thoracic width ranges

between 0.5 mm and 0.7 mm.The abdomen of the longest

specimen is 10.8 mm long and has about 59 segments, with

capillaries on the last six; that of the shortest is 7.3 mm, but

has about 86 segments, with capillaries on the last six. The

numbers of radioles, thoracic chaetal tufts and the extent of

the thoracic membranes in the three specimens is provided in

Table 11.

In two specimens the broad thoracic membranes are folded

outwards against the sides of the thorax (Fig.9, M,N).

Bayonet collar chaetae (Fig.9, O—-R) are similar to those of

the holotype, but lack an unserrated notch. Thoracic and

anterior abdominal uncini are also similar to those of the

holotype, with 4 or 5 teeth in a single row. Flat trumpet

chaetae number 6-8 in a bundle. Preserved in alcohol, the

abdominal segments show clumps of reddish-brown pig-

mented cells ventrolaterally, and of larger yellowish or orang-

ish cells ventrally (Fig.9, N).

The tube of the single juvenile paratype from Curacao

resembles that of the holotype; its diameter is 1.5 mm. The

worm has 6 radioles on the left and 5 on the right. As in the

specimens from Bonaire, the pinnule-free tips are very long.

However, both rudimentary opercula have already been lost

at this stage. The thorax is 0.5 mm wide and has 7 pairs of

chaetigers. The thoracic membranes end on the third thoracic

chaetiger. Two clusters of prostomial ocelli are present.

Thoracic glands were not seen. The chaetae also agree with

those of the specimens from Bonaire. In recently collected

material (Curacao, Piscaderabaai, 10.1.1990), thoracic mem-

branes end at 4/5, 4/4 respectively; the pinnule-free tips

generally are very long, rarely short; rudimentary opercula

are present in four specimens, absent in three.

LIVE MATERIAL. According to the field notes, rudimentary

opercula could not always be found, even in living specimens

from Curacao with radioles extended. The colouration of the

radioles is somewhat variable, often (transparently) whitish

to creamish, rarely yellowish to slightly orange or even

completely hyaline. At the base of the radioles there is a

series of up to six pairs of reddish spots, absent however in

the dorsal- and ventralmost radioles. Body predominantly

orange-brownish with up to 15 greenish-brown granules per

segment ventro-laterally in the abdomen and dorsally in the

thorax.

ETYMOLOGY. The specific name paraypsilon indicates the

close resemblance of the species to S. ypsilon.

HABITAT AND DISTRIBUTION. Occurs in shallow, clear, oce-

Table 11S. paraypsilon sp. nov. Meristic and other data from

specimens.

Specimen nos. 1 2 3

No. of radioles 8/? 8/8 8/7

No. of thoracic chaetal tufts 2/? 8/9 CHE

Thoracic membrane ends ?/? 3/3 3/3

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 63

Fig. 10 Spiraserpula singularis sp. nov. From type series. A & B, Two tubes, the second one younger. C, Substratal view of first formed coil

showing V-shaped dorsal ridge; a ventral ridge is absent. D, Same, smaller specimen. E, Paratype. F & G, Holotype and radioles. H-L,

From whole mount of another paratype: H & I, Two views of worm; J, only available bayonet chaeta; K, L, thoracic uncini; M, anterior

abdominal uncini. M-R, From whole mount of another specimen: N, anterior abdominal uncini and flat trumpet chaetae; O-R, Bayonet

collar chaetae, two present per side. S-V, Bayonet chaetae from whole mount of third specimen.

anic waters, at depths of up to 40 m on coral reefs. Hitherto

collected only from Bonaire, and Curacao, in the Caribbean.

Spiraserpula singularis sp. nov.

(Figs.10, A-V; 3, B; P1.2, E & F)

MATERIAL EXAMINED.

Puerto Rico : 1. Isla Matei, near buoy of Marine Institute,

vertical reef with surge channels, no sand, from living corals,

29-33 m, legit H. A. ten Hove, 2.x.1970, Stn. 2136A,

(HOLOTYPE & 3 PARATYPES, ZMAV.Pol.3710).

Curacao, (Neth. Ant.): 2. Salinja Fuik, reef in front, marine

park, 20-25 m, legit H. A. ten Hove, 18.1.1990 (2 specimens

BM(NH) 1992.166-167). 3. Piscaderabaai, outer bay, W of

entrance, sandy reef, 20 m, from underside of coral debris,

not in sediment, legit H. A. ten Hove, 12.1.1990 (3 speci-

mens, BM(NH) 1992.168-170).

TYPE LOCALITY. Puerto Rico.

DESCRIPTION.

TUBES: White, very tiny, one of the smallest species in the

genus. They may occur in mutually bonded aggregations of a

few individuals, or singly. Their coil diameters range from

1.2-1.3 mm. A granular overlay is present (Fig.10, A,B),

which makes the external diameters of the tubes

(0.5—0.6 mm) considerably larger than their internal diam-

eters (0.2-0.25 mm). Their apertures bear small, somewhat

lobed, peristome-shaped extensions (Fig.10, A,B), similar to

those found in S. massiliensis.

ITS consist of a V-shaped dorsal ridge, actually an inverted

V, along the convex side of the first formed coil (Figs.10,

C,D; 3, B). The two arms of the V are broader and outwardly

curved posteriorly, and their edges are smooth. Anteriorly

the dorsal ridge is a smooth plate only. A ventral ridge is

absent. When the worm is withdrawn into the tube, the

posterior, mid-dorsal part of the abdomen is applied to the

dorsal ridge.

WORMS: Four specimens were taken out of their tubes

(Fig.10, E-I). The holotype (Fig.10, F), which is the largest,

is only 5.7 mm long, 0.2 mm wide in the thorax, and its

abdomen is 4.6 mm long. There are four pairs of radioles

which, including the short and slender pinnule-free tips

(Fig.10, G), are about 0.55 mm long. There is a rudimentary

operculum on each side. Radioles are missing in the other

three specimens. However, a detached operculum was found

in the vial containing the specimens, and it is not certain

whether it belongs to one of them or another species.

Two clusters of prostomial ocelli are present. Five or six

globular ventral thoracic glands are present, more or less

arranged in a V. The numbers of thoracic chaetigers on the

two sides in the four specimens are: 9/9, 9/8, 8/8, and 7/7. It

was not possible to establish the extent of the thoracic

membranes due to the extremely small size of the worms. An

apron is, however, absent. One paratype with an abdominal

length of 1.95 mm has 29 segments, with capillaries on the

last 5, and another 3.0 mm long with 39 segments, but the

capillaries cannot be seen, having probably been damaged.

There are two bayonet chaetae in each collar fascicle. They

have moderately long serrated blades and 24 teeth on the

basal boss and some accessory ones (Fig.10, J, O-V; P1.2, E).

The unserrated notch is 1/5 the length of the blade. Thoracic

uncini (Fig.10, K,L) and anterior abdominal uncini (Fig.10,

M,N) have 6 and 4-6 teeth, respectively, all in a single row.

The middle abdominal uncini are rasp-shaped (P1.2, F), with

T.G. PILLAI AND H.A. TEN HOVE

up to 3 transverse rows of teeth above the single anterior

tooth. The abdominal segments bear 1 or 2 flat trumpet

chaetae in each bundle; one side is thickened into a claw-

shaped process (Fig.10, N).

REMARKS. In the comparison with other Caribbean species,

S. singularis would key out mainly on the absence of a ventral

longitudinal ridge/row of teeth and probably also the absence

of an operculum. So far, the presence of an operculum has

been observed only in a doubtful field identification. The

form of thoracic glands, shape of dorsal ridge and collar

chaetae are similar to those in S. plaiae.

ETYMOLOGY. singularis (Latin)= unique; referring to the

unique ITS.

HABITAT AND DISTRIBUTION. S. singularis sp. nov. appears

to be a shallow water coral reef dweller. It has hitherto been

collected only from Puerto Rico and Curagao.

Spiraserpula karpatensis sp. nov.

(Figs.11, A-K; 3, N)

MATERIAL EXAMINED.

Bonaire (Neth. Ant.): 1. Karpata, reef, 10 m, cryptic, legit H.

A. ten Hove, 9.xi.1988 (HOLOTYPE, ZMA V.Pol.3712;

PARATYPE, BM(NH) 1992.171).

Curacao (Neth. Ant.): 2. Reef in front of Salinja Fuik, buoy

13 of Marine Part, 20-30 m, corals and sandy/silty areas in

equal amounts. From under side of coral debris, not in

sediment, legit H. A. ten Hove, 18.1.1990 (1 specimen, ZMA

V. Pol. 3875).

TYPE LOCALITY. Bonaire (Netherlands Antilles).

DESCRIPTION.

TUBES: Pink, quite small, and lack longitudinal ridges. A pink

translucent granular overlay is present (Fig.11, A). The tubes

of both the types are coiled upon themselves, one much more

than the others (Fig.11, D). One has an erect part 2.0 mm

long, and a funnel-shaped, outwardly curved peristome

(Fig.11, A), while the other has a somewhat thickened

anterior end (Fig.11, D,E). The pink colouration gradually

fades to white towards the anterior end. The diameter of the

tubes is 0.6-0.7 mm in the attached parts, 0.4-0.6 mm in the

erect parts.

ITS consist of a serrated ventral ridge and an unserrated

dorsal ridge, with a sharp edge in cross-section (Fig.3, N).

The dorsal ridge may be absent (Fig.11, B,C), or greatly

reduced (Fig.11, D). In the latter it can be seen as a short

crescentic ridge through the broken end of one of the coils.

The serrated ventral ridge is regularly present (Fig.11, B, C,

and bottom left of D).

WORMS: The holotype is incomplete posteriorly (Fig.11, F),

broken in three parts, with a total length of 4.3 mm. An

operculum and 4 radioles are present on the left side; the

radioles on the right are missing. The length of the radioles is

approximately 0.8 mm, with a pinnule-free tip of 0.1 mm.

The operculum (Fig.11, G,H), is 0.26 mm long, and 0.28 mm

in diameter; inclusive of peduncle it is about 1 mm long.

Although bell-shaped, it is slightly zygomorphic, and has

numerous fine lobes, similar to that of S. plaiae described in

this paper. Branchial eyes have not been observed in the

fresh material. Two clusters of prostomial ocelli are present.

The thorax has 8 chaetigers on each side. The thoracic

membranes extend to the third chaetiger on the left (Fig.11,

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961

Fig. 11 Serpula karpatensis sp. nov. A, Tube with granular overlay, erect part and funnel-shaped peristome. B & C, Substratal view of two

tubes, opened to show internal serrated ventral ridge along convex wall, but absence of dorsal ridge. D, Aggregation of tubes, some

sections showing a very short crescentic dorsal ridge. E, Erect part from same aggregation showing somewhat thickened distal end; granular

overlay. F, Anterior end of holotype showing collar and thoracic membrane. G & H, Two views of zygomorph operculum. I-K, Bayonet

chaetae.

F) and the fourth on the right. It is not certain whether

ventral thoracic glands are present, but see note on live

material below.

Each collar fascicle bears 3 bayonet chaetae (Fig.11, I-K),

with moderately long, finely serrated blades, a moderately

long unserrated notch, and 3 teeth on the basal boss; the third

tooth may sometimes be difficult to observe and may be

reduced to a scar. Thoracic uncini have 6 (exceptionally 7)

teeth, anterior abdominal uncini 5, arranged in a single row.

The middle abdominal uncini are rasp-shaped, with 3-5 teeth

above the single anterior tooth. At least 35 abdominal

chaetigers are present, the last 7 with capillary chaetae.

Abdominal flat trumpet chaetae number 2-3 per bundle.

The specimen from Curagao agrees in most details with the

type material. Its numbers of radioles are 5/5, a long filamen-

tous rudimentary operculum is present opposite the opercu-

lum, and it has 38 abdominal chaetigers.

LIVE MATERIAL. As observed in material collected in 1990,

radioles are transparently lemon. Thorax ventrally with 5

bright red globules arranged in a V, presumably thoracic

glands.

ETYMOLOGY. named after the type locality, the coral reef in

front of the Sentro Ekologiko, Karpata.

HABITAT AND DISTRIBUTION. A shallow water cryptic reef

dweller. Has hitherto been recorded only from its type

locality in Bonaire, and Curagao.

66 T.G. PILLAI AND H.A. TEN HOVE

Fig. 12 Spiraserpula zibrowii sp. nov. From type specimens. A & B, Substratal view of two tubes opened to show the unserrated dorsal

ridge. The serrated ventral ridge consists here of a row of isolated teeth, but is a continuous ridge in the remaining material. C, Juvenile

paratype. D, Older specimen (holotype) broken in two. Anterior part with right rudimentary operculum; posterior abdomen with part of

dorsal ridge attached to mid-dorsal groove. E, Paratype. F & G, Two bayonet chaetae from holotype; H & I, two bayonet chaetae from

paratype. J—O from paratype. J, Thoracic uncini of small specimen with single row of teeth anteriorly and a cluster of more than one row

posteriorly. K, Similar uncini from first abdominal torus. L, Uncini from second abdominal torus; there are less teeth in a single row. M &

N, Anterior abdominal uncini, and flat trumpet chaetae with large lateral tooth. O, Posterior abdominal uncini; except for a single anterior

tooth, the uncini are rasp-shaped, with teeth in several rows.

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961

Spiraserpula zibrowii sp. nov.

(Figs.12, A—-N; 3, O; Pl.4, A-D)

MATERIAL EXAMINED.

Curagao (Neth. Ant.): 1. Lagoon of San Juan, E, raised reef,

lagoon side, Halimeda present, limestone cobbles, 10-15 cm;

from up to 20 cm deep crevices between cobbles, legit H. A.

ten Hove, 29.vi.1970, Stn. 2043 [HOLOTYPE &

PARATYPES 3-5: ZMA V.Pol.3707; PARATYPES 1

(slide), 2 (worm & tube fragments), and 7 (empty tube:

BM(NH) 1992.148-150); PARATYPE 6: USNM 130980

(unopened tube)].

Bonaire (Neth. Ant.): 2. Kralendijk, Flamingo Beach Hotel,

from corals, partly in sand, 45 m, legit H. A. ten Hove,

27.vii.1970, Stn. 2115D, (4 empty tubes, BM(NH)

1992.151-155). 3. 250 m N of Witte Pan, sandflat below reef,

47 m, mainly from the side of boulders, partly buried in sand,

legit H. A. ten Hove, 3.vii.1970, Stn. 2117B (4 empty tubes,

ZMAV. Pol. 3708).

TYPE LOCALITY. Curacao (Netherlands Antilles).

DESCRIPTION.

TUBES: Whitish, very tiny, and coiled upon themselves like

spirorbids, either individually (Fig.12, B), or in mutually

bonded aggregations of a few individuals. The direction of

coiling may reverse (see below). A fine granular overlay is

present. Longitudinal ridges are absent, but fine, smooth,

transverse growth markings are present. Juvenile tubes are

white. Although older tubes are white posteriorly, they have

a greyish-brown overlay anteriorly. The diameter of an

individual coil is 0.73 mm, with a tube diameter of 0.18 mm.

The maximum tube diameter is only 0.44 mm, which is the

smallest among the known species of the genus.

ITS consist of a serrated ventral ridge and an unserrated

dorsal ridge (Fig.12, A,B). The ventral ridge may consist

either of a continuous row of serrations, or only of a short

row of small separate teeth (Figs.12, A,B; 3, O). The dorsal

ridge is colourless and transparent, wedge- to Y-shaped in

cross-section, with its edges curved in places; it is spiral on a

columella-shaped axis when the tube is coiled upon itself.

Lateral ridges have not been found. The interior of the tube

may have a creamish lining.

One tube is coiled in one direction proximally, and in the

opposite direction distally. In the proximal coil the ITS are

similar to those described above. However, the distal coil has

only a columella-shaped axis with a dorsal ridge, which

became detached from the tube and is shown in situ (Fig. 12,

E); a serrated ventral ridge is absent here.

The mid-dorsal and mid-ventral longitudinal grooves of the

abdomen are applied to the unserrated dorsal ridge (Fig. 12,

D) and serrated ventral ridge of the tube, respectively.

WORMS: Measurements and meristic data are presented in

Table 12.

The right branchial half of the holotype (left missing)

shows a rudimentary operculum. The latter is present on both

| sides in the first paratype, but not in the second which is a

juvenile. The numbers of radioles on both sides are ?/3, 4/4

and 3/4, respectively. The pinnule-free tips are about 1/S—1/7

of their total length.

Two clusters of prostomial ocelli are present. The numbers

of thoracic chaetal tufts on both sides in the three specimens

are: 7/7, 8/7 and 8/7, respectively. The thoracic membranes

end on chaetigers 3/3 in the holotype and 4/4 in the juvenile

paratype; they are damaged in the second paratype. Two

Table 12S. zibrowii sp. nov. Measurements and meristic data

from Holotype and two paratypes.

Abdomen

Total Thoracic

length width Length Number of capillaries

(mm) (mm) (mm) segments on

Holotype 9.7 0.23 8.5 54 a

Paratype 1 7.0 0.18 5.8 43 7

Paratype 2 3.4 0.18 2.1 oi 9

translucent ventral thoracic glands are present, although not

as easily discerned as in some of the other species.

There are 2 or 3 fully developed bayonet chaetae per side

in the juvenile, 4 in the older specimens. They have moder-

ately long serrated blades, an unserrated notch which is about

1/4-1/5 the length of the blade, and 4 or 5 somewhat large

teeth and some accessory ones on the basal boss (Fig.12, F,I;

P1.4, A). The thoracic uncini have a single row of 6~7 teeth

(P1.4, B). Anterior abdominal uncini bear a cluster of small

teeth in two to seven rows at their posterior ends, and a single

row of larger teeth anteriorly; this type of uncini may occur in

juvenile specimens also (Fig.12, J-M). The posterior abdomi-

nal uncini are, however, similar to those of the other species

in being rasp-shaped, with 6 transverse rows of 2-5 teeth

each, except for the single anterior tooth (Fig.12, O; P1.4, C).

The abdominal flat trumpet chaetae number about 5 per

bundle. Their somewhat triangular, curved distal ends are

thickened and hooked at one end, and drawn out into an

acute angle at the other (Fig.12, N; Pl.4, D). Up to 54

abdominal segments are present, the last 4-9 with capillary

chaetae.

REMARKS. The collections from Bonaire, a mere 50 km from

Curacao, consist of a total of 8 empty tubes whose ITS are

identical with those of the present species. The largest tube

from Witte Pan has a coil diameter of 2.2 mm; two tubes have

erect portions with peristomes. Three tubes are white. The

fourth is creamish in colour, with a creamish interior lining.

Fine transverse growth markings are present on all. The

serrations of the ventral ridge are arranged on a low longitu-

dinal ridge in some of them.

In the absence of worms, and the markedly different

habitat from which they were collected (at a depth of

45-47 m), these tubes cannot be conclusively identified as S.

zibrowii.

ETYMOLOGY. named after H. Zibrowius, who recognized

some of these small species as being new.

HABITAT AND DISTRIBUTION. Appears to be a shallow water

species inhabiting crevices between boulders and their under-

sides in sandy areas close to coral reefs. Hitherto collected

from Curacao. Two uncertain records from Bonaire.

Spiraserpula plaiae sp. nov.

(Figs. 13, A-T; 3, K)

MATERIAL EXAMINED.

Curacao (Neth. Ant.): 1. Salinja Fuik, near Ceru Preekstul,

open reef, coral debris, 33 m, from limestone boulder on

sand, legit H. A. ten Hove, 18.ix.1970, Stn. 2088A (HOLO-

TYPE & PARATYPES 1 & 5: ZMA V. Pol.3713;

PARATYPES 2, 4, & 6: BM(NH) 1992.173-174;

PARATYPE 3: USNM 130990). 2. Reef in front of Salinja

Fuik, buoy 13 of marine park, coral debris, 18-27 m, legit H.

A. ten Hove, 18.i.1990 (4 specimens, ZMA V. Pol.3874). 3.

Cornelisbaai, E, steep reef, coral debris, 18-26 m, legit H.

A.ten Hove, 17.1.1990 (6 specimens, ZMA V. Pol.3873). 4.

Piscaderabaai, outer bay W of entrance, sandy reef, coral

debris, legit H. A. ten Hove, 12.1.1990 (5 specimens, ZMA

V. Pol.3872).

TYPE LOCALITY. Curacao (Netherlands Antilles).

DESCRIPTION.

TUBES: White to greyish-brown, occurring either individually

coiled upon themselves (Fig.13, A), or in mutually bonded

aggregations of a few individuals (Fig.13, B). They are

sub-circular in cross-section, with faint lateral ridges, and

bear fine smooth transverse ridges, and often have erect

anterior ends (Fig.13, A,B). A fine opaque granular overlay

is present (Fig.13, B), which can be seen under special

illumination only. Their external diameter attains 1.0 mm,

their erect portions somewhat smaller. The inside of one tube

has a light caramel coloured lining.

ITS consist of a serrated ventral ridge along the concave

side of the tube (Fig.13, C-E, H), which may not be well

developed and represented only by a few isolated or coa-

lesced teeth slanting backwards (Fig.13, F), and a smooth

dorsal ridge on the convex side (Figs.13, F,G; 3,K). The

latter is wedge-, T to Y-shaped in cross-section. In specimens

coiled upon themselves, the dorsal ridge occurs spirally on a

columella-shaped axis (Fig.13, I). A short accessory latero-

dorsal ridge, which tapers anteriorly and posteriorly, may

also be present on either side (Fig.13, F). Their edges are

unthickened.

In life, the mid-ventral and mid-dorsal longitudinal grooves

of the abdomen are applied to the serrated ventral and

smooth dorsal ridges, respectively, of the tube (Fig.13, G).

WORMS: Six specimens were taken out of their tubes. The

abdomen is complete in only one. Even though preserved in

alcohol, the abdominal segments still show clusters of pig-

mented specks laterally, light yellowish in one specimen, light

to bright orange in two, light to dark brown in two, and

uniformly caramel coloured in another.

Five specimens have an operculum on one side and a

rudimentary operculum on the other; the branchial crown is

partly missing in the sixth. The length of the operculum and

peduncle varies from 1.0 mm in a juvenile paratype to

1.8 mm in the holotype. The operculum itself is 0.4-0.5 mm

long, 0.3-0.5 mm wide. It is zygomorph, attached to the

peduncle eccentrically, and bears numerous (up to 50) radii

(Fig.13, K-M). The distal diameter of the peduncle is 1/3 to

2/3 that of the opercular base. The numbers of radioles on

both sides are 6/5, 5/4, 4/5, 4/4 and 4/3. They end in short

slender pinnule-free tips, which are about 1/5 to 1/7 the length

of the radioles (Fig.13, K). The only complete specimen

(Fig.13, J), is 4.9 mm long, with 42 abdominal segments, the

last 10 with capillaries. However, in another specimen, which

T.G. PILLAI AND H.A. TEN HOVE

is incomplete (Fig.13, M), 76 abdominal segments could be

counted.

Two clusters of prostomial ocelli are present. The num-

bers of thoracic chaetal tufts are 11/7, 9/9, 9/8, 8/8, 6.6.

Thoracic membranes extend to chaetigers 6/5, 4/4, 4/3, 4/?,

and they are damaged on both sides in the fifth. Two

groups of transparent to translucent ventral thoracic

glands, arranged in a V, and of unknown function, are

present. Further studies are needed to find out if they

could be responsible for secreting the caramel coloured

inner lining of the tube.

Collar fascicles bear 2 or 3 fully formed bayonet chaetae

each, and a newly formed one deep within the bundle. Each

bayonet chaeta consists of a moderately long blade, a moder-

ately long unserrated notch which is 1/3 to 1/4 the length of

the blade, and 24, seldom 5, teeth on the basal boss (Fig. 13,

N-T). The teeth are comparatively larger as their number

decreases (Fig.13, Q-T), and they may be accompanied by

one or two accessory teeth (Fig.13, O,P,S). Thoracic uncini

possess 5-7 teeth in a single row (Fig.13, U). Abdominal

uncini are similar, with 5—6 teeth; anteriorly saw- and rasp-

shaped uncini may occur in a single row, posteriorly all uncini

are rasp-shaped.

LIVE MATERIAL. As observed in material collected in 1990,

radioles are faintly yellow to lemon, operculum is transpar-

ent, almost colourless. Thorax ventrally with 24 bright red to

orange globules arranged in a V, presumably thoracic glands;

body transparent with yellow tinge, brownish gut.

ETYMOLOGY. Named after Gayle Plaia who, when working

at the Florida Marine Research Institute, first observed ITS

in one of the species, S. ypsilon, from the Gulf of Mexico.

HABITAT AND DISTRIBUTION. S. plaiae is a shallow water

species occurring in coral reefs, and has hitherto been col-

lected only from the type locality.

Spiraserpula caribensis sp. nov.

(Figs. 14, A-M; 15, A-Y; 16, A-K; 3, L; Pl.4, E & F;

P1.5,A-E)

MATERIAL EXAMINED.

Curacao (Neth. Ant.): 1. Awa Blancu, coral debris barrier,

20-30 cm, legit H.A. ten Hove, 15.ix.1975, Stn. 75-38

(HOLOTYPE & 3 PARATYPES: ZMA’ V-Bol-37l572

PARATYPES USNM 130987; 4 PARATYPES each: AM

W20157, NSMT, ZMK). 2. Awa Blancu, 3—4m, legit H. A.

ten Hove, 14.x.1975, Stn.75—37 (1 specimen, HUJ). 3. Awa

Blancu, coral debris, near Lagoen Blancu, 30-50 cm, legit

H. A. ten Hove, 30.vii.1970, Stn. 2090 (several subsamples

BM(NH) 1992.25-31, FSBC I 39195, ZMA V. Pol. 3716,

ZMB). 4. Lagoen Blancu, coral debris barrier, Halimeda,

20-30 cm, legit H. A. ten Hove, 15.ix.1975, Stn.75-36 (2

out of several specimens, RMNH 18174). 5. Awa di

Oostpunt, coral debris barrier, 30-50 cm, legit H. A. ten

Hove, 3.x.1975, Stn.75-77 (1 out of few specimens,

BM(NH) 1992.10-11. 6. St. Jorisbaai, Peninsula Groot St.

Fig. 13 Spiraserpula plaiae sp. nov. A, Aggregation of tubes, showing fine transverse growth markings, granular overlay in places. B,

Juvenile tube. C-E, Fragments of tubes showing serrated ventral ridge (C & D, paratype 3), (E, paratype 2). F, Holotype showing variant

form of ventral ridge with isolated teeth, and ventro-lateral ridge. G-I, Paratype 4: G, posterior end of tube and worm. H, Portion of tube

showing ventral ridge. I, Dorsal ridge on columella-shaped axis. J-L, Two views of paratype 4, and its operculum: J, (Operculum not seen),

ventral longitudinal abdominal groove, and dorsal groove within 2nd coil; K, showing operculum; L, Another view of operculum. M,

Paratype 2. N & O, Bayonet chaetae from holotype. P-T, Bayonet chaetae from paratype. U, Thoracic uncini from holotype.

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961

70 T.G. PILLAI AND H.A. TEN HOVE

Fig. 14 Spiraserpula caribensis sp. nov. A-J, From Florida, Stock Island, Stn. 7B. A, Tube with granular overlay and longitudinal ridges.

B-F, ITS seen in various tube fragments: B, bottom cross-section showing median dorsal ridge on convex wall, narrow lateral ridge on

either side, and serrated ventral ridge (barely visible); middle cross-section with serrated ventral ridge along concave wall; C, ventral ridge

in sectional view (barely visible), and dorsal and lateral ridges; D, substratal view of first formed coil of tube opened to show smooth dorsal

and serrated ventral ridge; E, smooth dorsal ridge in the first formed coil, on the right; F, uncoiled part of tube showing smooth dorsal

ridge tapering at both ends. G, Anterior end of specimen, with thoracic glands, and showing dorsal and ventral abdominal grooves. H,

Radioles of same specimen with left and right filamentous rudimentary opercula. I & J, anterior end of a worm with two views of collar.

K-M, Bayonet chaetae.

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961

Joris, muddy pebbles, Thalassia flat, few corals, from

limestone boulders, 30cm, legit H. A. ten Hove,

10.ix.1970, Stn.2096 (2 out of several specimens, BM(NH)

1992.20-24, RMNH 18175). 7. St. Jorisbaai, Koraal Tabak,

Punta Blanco, undersides of boulders, on rocky debris,

20-30 cm, legit H. A. ten Hove, 15.xi.1988, 9.1.1990, near

Stn.75-30 (5 out of several specimens, MCZ, ZMH). 8. St.

Jorisbaai, entrance channel, W, boulders and large metal

poles in surf; from undersides crusts of Spiraserpula, legit

H.A. ten Hove, 16.i.1990 (clusters, AM W20341, HUJ).

Aruba: 9. Spaans Lagoen, SE of bridge, rocks, etc., at

floodgate, mud, Rhizophora, 0-2.0 m, legit P. Wagenaar

Hummelinck, 24.iii.1970, Stn.1673 (2 specimens BM(NH)

1992.12-13). 10. Andicuri, cape W of beach, windward

side, rockpool, exuberant coral growth, strong wave

action, 0.5m, legit H. A. ten Hove, 20.viii.1970,

Stn.2034B (several fragments of tubes, 2 incomplete

worms, ZMA V. Pol. 3719).

Barbuda: 11. Great Lagoon, Lobster Point, N. of Palm

Beach, Thalassia and Halophila, O0-1.0 m, legit P.

Wagenaar Hummelinck, 23.vii.1967, Stn.1534 (3 out of

several specimens, ZMA V. Pol. 3725).

Bonaire (Neth. Ant.): 12. Lac, dam, beachrock in current

behind surf, 5-10 cm, from crevices in beachrock, legit H.

A. ten Hove, 15.vii.1970, Stn. 2123 (1 specimen, USNM

130986). 13. Lagun, N shore, 500 m from entrance, rock,

boulders, 0-50 cm, from undersides of boulders, legit H.

A. ten Hove, 23.vi.1970, Stn. 2129 (3 out of several

specimens, ZMAV. Pol. 3720). 14. Bonaire, Karpata, reef,

10 m, cryptic, legit H. A. ten Hove, 9.xi.88 (1 tube,

BM(NH) 1992.14).

Jamaica: 15. Drunkeman’s Key, sandy debris, 0-0.5 m,

legit P. Wagenaar Hummelinck, 15.vi.1973, Stn.1683,

(ZMA V. Pol. 3723).

Puerto Rico: 16. La Parguera, E, glade in mangroves,

Thalassia beds, muddy sand, from between boulders,

20-30 cm, legit H. A. ten Hove,1.x.1970, Stn.2135 (3

specimens, ZMA V.Pol.3724).

Panama: 17. Gatun Locks, walls of outer platform, lower

W chamber, Pan. Survey, 20.iii.1972, Pacific Stn. 81-1, M.

L. Jones coll., USNM No.58661 (2 specimens without their

tubes). 18. Same, Stn.81-2, M. L.Jones coll., USNM

No.58662, (1 specimen with its tube).

Florida: 19. Safe Harbour, Stock Island (near Key West),

Florida Keys, 5m, from chunks of calcareous materials

(shells, barnacles, etc.) cemented together and covered

with serpulids and small cirratulids, legit R. Chesher and

C. Hamlin, 17.vii.1970 and 1.vi.1971, Stn.7B, (22 out of

several specimens, USNM 130988, BM(NH) 1992. 15-19,

ZMAVYV.Pol.3721 (10+ specimens from 1.vi.1971). 20. Off

Egmont Key, 27. 0°37.0'N, 83°01.5'W, sea buoy, 18 m,

scarce sponges and corals, 2 cm of soft sludge on lime-

stone, many serpulids, legit H.A.ten Hove and T. Perkins,

2.1.1980, Stn.EJ.80002, (9 out of several specimens, ZMA

V. Pol. 3722, FSBC I 39202).

TYPE LOCALITY. Curagao (Netherlands Antilles).

| DESCRIPTION.

TUBES: Light to bright pink or rose coloured. They form

mutually bonded aggregations of a few to several individuals.

Their external diameter is generally about 1.0 mm, maxi-

mally 1.5mm. There are three longitudinal ridges, one

median and one along each lateral margin, which may be

indistinctly developed in places (Figs.14, A; 15, A). Narrow

transverse ridges may be developed to various extents

(Fig.15, A). Some of the tubes end anteriorly in 4 rounded,

anteriorly-directed lobes. A transparent to translucent granu-

lar overlay is present. The granulations are larger and more

densely laid along the ridges. The pink colour is faint along

the longitudinal ridges, as seen through the transparent

granules, but form of a pair of bright longitudinal bands

between the ridges. Branching tubes, difficult to observe

since they form dense aggregations, have been observed in

material from Curacao (Stn. 2090, 2096, 75-38), and from

Bonaire (Stn. 2123).

ITS consist of a serrated ventral ridge along the concave

wall (Figs.14, B,D; 15, P), and a smooth dorsal ridge

opposite (Figs.14, B, D-F; 15, O,P). The dorsal ridge is

nearly tongue-shaped in cross-section, with a gradual

decrease of its height, thickness and width of the widest part

both anteriorly and posteriorly. This is occasionally more

clearly seen in the non-coiled portions of tubes (Fig.14,F).

The dorsal ridge may be situated on a columella-shaped axis

in tubes coiled upon themselves (Fig.14, E). They usually

also possess a short accessory dorso-lateral ridge on either

side of the dorsal ridge (Figs.14, B & C; 3, L). The inside of

the tube may have a light caramel to light brown lining. The

mid-ventral and mid-dorsal longitudinal abdominal grooves

of the worm are applied to the serrated ventral and smooth

dorsal ridges, respectively.

WORMS: The longest available complete worm is from

Florida. It has a total length of 12.8 mm, thoracic width of

0.5 mm, abdominal length of 9.7 mm, and has 91 segments,

with capillaries commencing on the 80th. There are four

radioles and a rudimentary operculum on each side. Fully

developed opercula are absent in all the specimens, being

represented by a long and filamentous rudimentary opercu-

lum on each side (Fig.14, H). The highest number of radioles

is 6 pairs, the longest measure about 2.1 mm, and end in

slender pinnule-free tips which are 1/5-1/6 their entire length

(Fig.14, H). Radioles have up to 12 pairs of pinnules each, as

could be observed in living material. The smallest worm is a

juvenile from Curago (Stn. 75-77) which has a total length of

3.7 mm, a thoracic width of 0.45 mm, abdominal length of

2.0 mm, and has 20 segments, with capillaries in the last 5. It

has 4 radioles on the left and 5 on the right, in addition to the

rudimentary opercula.

Two reddish to reddish-brown clusters of prostomial ocelli

are present. The median lobe of the collar is sub-rectangular,

with rounded lateral borders and a smooth medial notch

(Fig.14, I & J). Five to seven globular ventral thoracic glands

are present (Fig.14, G), more or less arranged in a V.

Whether they are responsible for secreting the brownish

inner lining of the tube or not has to be further investigated.

A summary of data is presented in Table 13. Similar data

from the Florida material are provided in Table 14.

The bayonet collar chaetae, which number 3 or 4 fully

formed ones per fascicle and, usually, a developing one deep

within, are unique among the species of Spiraserpula and of

Serpula that have hitherto been described. Their blades are

conspicuously short, unserrated and dagger-shaped (Figs. 14,

K-M; 15, B-I, Q-W; P1.4, E & F). The number of large

conical teeth on the basal boss is usually 3 or 4. Often there

are 2 large teeth with 1 or 2 smaller ones in between (Fig.14,

K-L; 15, B-I). In the specimens from Gatun Locks, Panama,

the number of teeth is usually 4 or 5 (Fig.15, Q-W). These

dagger-shaped bayonet chaetae were noted and figured in the

Table 13 S. caribensis sp. nov. A summary of data from four

samples from Curagao (Stns. 75-38, 75-36 and 75-77 and 2096).

No. of specimens (n=10) Dr 1

No of radioles per side 6/5 S/S 5/4

No. of specimens (n=14) Deri 3.) Bae

9/8 9/7 8/8 8/7 7/7 7/6

No. of specimens (n=10) LL gr Sk Te a

Thoracic membrane ends S4 S/37 4/4 4/3, 3/3

No. of thoracic chaetal tufts

Table 14S. caribensis sp. nov. A summary of data from the

Florida material.

No of specimens (n=11) 2 5 4

No of radioles per side 6/6 5/5 4/4

No. of specim.(n=29) 1” 8 IRSISOY GON oy DIT De Ane

No. of thor. chaet. 10/6 9/8 9/7 8/8 8/7 8/6 7/7 7/6 7/5 6/6

No. of specimens (n=26) Sip Wee) whale SY

Thor. membranes end on 5/4 5/3 4/4 4/3 3/3

unpublished research of M. van Vliet and R. Fijn (see

acknowledgements).

The blades of developing bayonet chaetae deep within the

fascicle are similar to the fully formed dagger-shaped bayonet

chaetae, indicating that the latter have not resulted from wear

and tear of bayonets with tapered tips. Occasionally, a

developing chaeta with a truncated blade and tapered tip

(Fig.15, F,S), occurs deep within a fascicle, which provides a

clue to the origin of the former. Reduction in length of the

blade together with extension of the unserrated notch has

resulted in stout, truncated bayonet chaetae, with smooth

and dagger-shaped blades.

Thoracic uncini (Fig.15, J) usually possess 6 teeth, and

anterior abdominal uncini (Fig.15, K,X) 4 or 5, in a single

row. Posterior abdominal uncini are rasp-shaped (PI.5, A).

Flat trumpet chaetae number up to about 5 in each bundle,

and their triangular distal ends bear a hook-shaped process

on one side, and the other side is drawn out into an acute

angle (Fig.15, L-N, Y; P1.5, B).

COLLECTIONS FROM OTHER LOCALITIES. The specimens from

the other localities listed above agree with those from the

type locality. However, the smaller size of the tube and

chaetae of the specimens from Gatun Locks, Panama, and

the highly branched tubes of the specimens from Grenada,

are worth noting.

LIVE MATERIAL. As observed in material from Curacao,

radioles are colourless, transparent to transparently orange,

sometimes with reddish pinnules. Base of branchial lobes and

the collar may be tinged with purple. Branchial eyes not

T.G. PILLAI AND H.A. TEN HOVE

present. Body predominantly transparent orange, thorax

ventrally reddish.

ETYMOLOGY. The name acknowledges the fact that this

appears to be the most widely distributed species of Spiraser-

pula in the Caribbean.

HABITAT AND DISTRIBUTION. S. caribensis inhabits shallow

water, intertidally down to a few metres in the Caribbean, to

18 m in the E. Gulf of Mexico (temperature submerged ?). It

occurs in a variety of habitats, from rockpools to the under-

sides of boulders in mangrove glades. It is able to survive well

in somewhat muddy environments, always, however, cryptic

between piles of rock or similar hard substrata.

It appears to be widely distributed in the Caribbean and

Gulf of Mexico, from Florida to Barbuda and Panama.

A population from Grenada, with frequently branching

tubes and which is, for the present, regarded as belonging to

S. caribensis, is described below (Fig.16, A-K):

MATERIAL EXAMINED.

Grenada (Caribbean), Hog Island, near Pt. Salines, 0-1.5 m,

Rhizophora, mud, legit P. Wagenaar Hummelinck,

8.vii. 1967, Stn. 1550 (5 specimens and 4 tubes, ZMA V.Pol.

3706, USNM 130985, BM(NH) 1992.32).

DESCRIPTION.

TUBES: Dark pink to rose coloured. Except for their posterior

ends, they are all uncoiled, conspicuously branched, and

attached to the substratum throughout (Fig.16, A). A granu-

lar overlay is present, larger granules constituting a median

longitudinal ridge and a pair of lateral ridges (Fig.16, A,B).

The colouration is darker pink between the median and

lateral ridges. Fine transverse ridges may be present in

places. The lumen of the tube is continuous with that of the

branches.

ITS are similar to those of S. caribensis. However, the

tongue-shaped cross-section of the dorsal ridge is somewhat

more pronounced.

WORMS: Three worms were taken out of the tubes, of which

the longest (Fig.16, C,D), has a total length of 9.2 mm. There

are up to 5 pairs of radioles and a rudimentary operculum on

each side. The radioles are up to about 2.0 mm long, 1/6-1/8

of which constitute pinnule-free tips. Measurements and

meristic data are given in Table 15:

Two clusters of prostomial ocelli are present. All three

specimens bear 7 thoracic chaetal tufts on the left and 6 on

the right. The thoracic membranes end on the third thoracic

chaetiger on both sides in the first specimen, but are damaged

in the others. A pair of ventral thoracic glands is present

(Fig.16, C).

Each bayonet chaeta typically consists of a short, serrated

blade, and an unserrated notch and a tapered tip (Fig.16,

E-K). There are 2 or 3 large teeth on the basal boss, and a

few accessory teeth. Older chaetae in a fascicle which have

Fig. 15 Spiraserpula caribensis sp. nov. A-E & J-N, From Curagao, St. Jorisbaai, Stn. 2096. F-I, from Curagao, Lagoen Blancu, Stn. 75-36.

O-Y, From Panama, Gatun Locks: (O-S, from Stn. 81.1; T-Y, from Stn. 81.2). A, Tubes showing granular overlay, external ridges and

transverse wrinkles. B—E, Bayonet chaetae from the same fascicle with short dagger-shaped blades. F-I, Bayonet chaetae from fascicle of

another specimen: F, Newly formed, deep within fascicle; G-I, Older chaetae. J, Thoracic uncini. K, Anterior abdominal uncini, and L,

flat trumpet chaetae, from same segment. M & N, Flat abdominal trumpet chaetae from other specimens. O & P, Tube opened

substratally, viewed from two different angles, with worm in situ showing thoracic glands; O, with dorsal ridge only, and P, with both dorsal

and ventral ridges. Q-S and T—W, Bayonets from two different fascicles. Note much smaller size compared with those of Florida (Fig. 14,

K-L) and Curagao (Fig. 15, B—I) material, although drawn under same magnification. X & Y, Anterior abdominal uncini and flat trumpet

chaetae from same segment. Note much smaller size than in Curacao material (Fig. 15, L-N). «

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961

74 T.G. PILLAI AND H.A. TEN HOVE

Fig. 16 Spiraserpula caribensis sp. nov. A-K, From Grenada. A, Branched tube with granular overlay and longitudinal ridges. B, Branching

point marked Y in A, magnified. C & D, Two views of a worm showing rudimentary opercula, thoracic glands (C), and dorsal and ventral

longitudinal abdominal grooves. E-H, Four bayonets from a small specimen: A, Older chaeta with worn out tip; F-G, Chaetae with intact

tips. I-J, Bayonets from a larger specimen: the oldest (K) with a worn out tip, and the other two with intact tips.

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 iS

Gif PE SN

FA ew)

A Liza

| Gaz

Fig. 17 Spiraserpula nudicrista sp. nov. From Bonaire. A, Tube, with granular overlay and longitudinal ridges. B-D, Tube fragments

showing unserrated ventral ridge along concave wall and lack of a ridge on the opposite side. E-I, Other tube fragments showing a narrow

ventral ridge anteriorly, and a smooth and rounded edge to it posteriorly. J, First formed coil, lacking a dorsal ridge. K-L, Two different

views of holotype showing prostomial ocellar clusters, as seen through the collar, the ventral longitudinal groove of the abdomen, and

extent of thoracic membrane on the left side (K); L, Radioles with a pair of short club-shaped rudimentary opercula and moderately long

pinnule-free tips. M & N, Paratype: M, tube with granular overlay, prostominial ocellar cluster of one side, and longitudinal abdominal

grooves; N, branchial crown with a pair of short club-shaped rudimentary opercula, and long pinnule-free tips.

Table 15S. caribensis from Grenada. Measurements and counts.

Radioles Abdomen

Total Thoracic Number

Specimen length width Length Length of Capillaries

no. (mm) (mm) (mm) No.(mm) segments on

1 9.2 0.5 DAY SS OLS 50 44

2 8.1 0.5 De Aey| 30 7

3 3a 0.35 OG SIE) its 35 10

lost their tapered tips through abrasion may appear some-

what like the bayonets of S. caribensis from elsewhere

(Fig.16, E, K; Pl.5, C-E), but the newly formed bayonets,

within the fascicle, possess tapered tips.

HABITAT AND DISTRIBUTION. Appears to inhabit shallow

water and capable of withstanding the silty conditions found

in mangrove backwaters. It was found on the inside of a dead

oyster shell covered with much silt.

REMARKS. The extensively branching tubes and differences

in the collar chaetae initially led us to consider the Grenada

material as possibly belonging to a distinct species. However,

branching as such, although inconspicuous, was also subse-

quently observed in some specimens of S. caribensis from

Bonaire (Stn. 2123) and Curagao (Stns. 2090, 2096,75-38; see

above), in S. paraypsilon from Curacao (10.1.90). Moreover,

S. snellii, described later in this paper, revealed a schizont

with parent in one tube. By itself, therefore, branching

cannot be a good character to separate the Grenada material

as a distinct species. The fully formed bayonet chaetae,

including those within the fascicle, of S. caribensis proper,

have short dagger-shaped blades with blunt tips, while blades

of the Grenada material typically end in tapered tips.

Although the tip of a fully formed chaeta in the Grenada

material might be lost through abrasion (Fig.16, E-K), those

deep within the fascicle are tapered.

Further work on additional material is necessary to deter-

mine whether frequent branching of the tubes and the

features of the bayonet chaetae are consistent, and whether

there are other characters which would justify the separation

of the Grenada material into a distinct species or not.

Spiraserpula nudicrista sp. nov.

(Figs.17, A-N; 18, A—O; 3, F; Pl.3, A-D)

MATERIAL EXAMINED.

Bonaire (Neth. Ant.): 1. Karpata, reef, cryptic, 10 m, legit H.

A.ten Hove, 9.xi.1988, (HOLOTYPE & PARATYPE:

ZMA V. Pol.3711).

Curacao (Neth. Ant.): 2. Savonet, E of Boca Braun, reef, no

sand, about 22 m, from corals, some dead, legit H. A. ten

Hove, 28.xi.1970, Stn. 2101 (PARATYPES 2 & 3: BM(NH)

1992.61 & 62).

TYPE LOCALITY. Bonaire (Netherlands Antilles).

T.G. PILLAI AND H.A. TEN HOVE

DESCRIPTION.

TUBES: White to creamish white and have a conspicuous

granular overlay (Fig.17, A,E,M). They may be covered over

by encrusting calcareous organisms. They are trapezoidal in

cross-section, with two longitudinal ridges along the crest of

the tube and one along each flank (Fig.17, A). The maximum

external tube diameter of the holotype is 1.0 mm.

ITS consist of an unserrated ventral ridge which is rounded

and smooth towards its middle (Figs.17, H; 3, F), from where

it decreases in thickness and height both anteriorly and

posteriorly (Fig.17,C-E, F-G). A dorsal ridge is generally

absent, even on the convex pulley-shaped posterior end

(Fig.17, I). However, paratype 1 from Bonaire showed some

isolated dorsal teeth. The mid-ventral longitudinal groove of

the abdomen (Fig.17, M) is applied to the unserrated ventral

ridge.

WORMS: Only two worms were yielded by the tubes from

Bonaire. The complete holotype has a total length of

15.6 mm, a thoracic width 0.7 mm, an abdominal length

11.9 mm and about 101 segments, with capillaries on the last

7. The radioles are 2.5 mm in length, and their pinnule-free

tips of 0.6 mm are comparatively long (Fig.17, L, N). The

paratype is incomplete posteriorly.

The holotype has 9 pairs of radioles while the paratype has

8 pairs. Both specimens have a short filamentous rudimentary

operculum on each side (Fig.17, L). Two clusters of prosto-

mial ocelli are present, and are seen as conspicuous brown

patches through the collar (Fig.17, J,K,M). This is in contrast

to the other known members of the genus in which they can

be seen when viewed from the anterior end with the radioles

removed or when mounted.

Both specimens have 8 pairs of thoracic chaetal tufts, and

the thoracic membranes end on the fourth chaetiger on the

left (Fig.17, K) and the Sth on the right. Ventral thoracic

glands appeared to be absent.

The collar fascicles of the holotype possess four bayonet

chaetae with long serrated to pilose blades and several conical

teeth on the basal boss (Fig.18, A—D; P1.3, A). There may be

a number of accessory teeth arranged around the bases of the

larger teeth, which are lacking in the paratype (Fig.18, E-G).

The unserrated notch is short. Thoracic uncini (Fig.18, H;

P1.3, B) and anterior abdominal uncini (Fig.18, I; Pl.3, C)

possess 4 or 5 teeth arranged in a single row. There are about

4 flat trumpet chaetae in each abdominal fascicle (Fig.18,

J;P1.3, D). An anterior hook, as in most of the species of the

group, cannot be discerned, all distal teeth appearing more or

less equally developed.

Tubes of the specimens from Curagao are similar to those

from Bonaire with regard to colour, form and ITS (Fig.18,

K). Their maximum external diameters are 1.1-1.2 mm. Two

of them yielded worms which are incomplete posteriorly.

Some data from them are presented in Table 16.

Both specimens possess a rudimentary operculum on each

side (Fig.18, L,M). Bayonet collar chaetae (Fig.18, N,O), are

similar to those of the specimens from Bonaire, although

their basal bosses are somewhat stouter.

REMARKS. A small fragment from the inside of the coil of

Fig. 18 Spiraserpula nudicrista sp. nov. A-J, From Bonaire. A-D, Bayonet chaetae, holotype. E-G, Three, out of five, bayonet chaetae

from paratype. H—J, from paratype: H, Thoracic uncini; I & J, flat trumpet-shaped chaetae and uncini from anterior abdomen. K-O, From

Curaao. K, Tube, with granular overlay, external longitudinal ridges, and internal ventral longitudinal ridge seen through fractured end. L,

Branchial crown of older specimen with pair of rudimentary opercula. M, Branchial crown of younger specimen, with pair of shorter

rudimentary opercula. N & O, Two, out of four, bayonet chaetae.

| Lhe x 4 " 0.5 )

| x

) HAI L- 7 \ KZ {

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Ly K E

\ | : By

\\' \\ t —S = \ Gr 4 iy ~\ 0.5

( } ii PAS

\ | PN

NN LE A 1

\ yy hy

\ i Hr}

Fah

\ \ (hs Np

\ W\ N

‘ ‘ p 0.01mm | \

IY , AN )

PU, Mth ats) as

WY ’ \ M

\ 4 i

: NY : | \

\ \ G 0.01mm

\\ \ "

i \

N\\ (tS Wl

\ \ " \ 4

C \ |

Table 16 S. nudicrista sp. nov. Some data on two specimens from

Curacao.

Specimen1 Specimen 2

Width of thorax 0.5 mm 0.6 mm

Length of radioles 2.1mm 2.6 mm

Pinnule-free tips (Figs.17L,M) short short

No. of radioles (L/R) 7/8 9/8

No. of thoracic chaetal tufts (L/R) 9/7 8/6

Thoracic membrane ends 2/? 4/4

Length of abdomen ? 9.7 mm

No. of abdominal segments z, about 60

paratype 1 showed a slightly concave to asymmetrical cross-

sectional edge to the ventral ridge, and a few isolated teeth in

the location of the dorsal ridge, somewhat similar to the

condition in S. paraypsilon. There is also some similarity in

the collar chaetae.

ETYMOLOGY. nudus (L) = unadorned; crista = crest, ridge.

HABITAT AND DISTRIBUTION. S. nudicrista is a shallow water

cryptic species inhabiting coral reefs. It has hitherto been

collected from Bonaire and Curagao.

Spiraserpula sp.

(Fig.19, A—C)

MATERIAL EXAMINED.

Curacao (Neth. Ant.): Piscadera Baai, outer bay in front of

CARMABI, muddy reef, many sand spots, about 40 m, from

dead corals, legit H.A.ten Hove, 9.vi.1970, Stn. 2054B (3

empty tubes, and some abdominal fragments, ZMA V. Pol.

3883).

DESCRIPTION. Tubes are white, circular in cross-section. An

T.G. PILLAI AND H.A. TEN HOVE

erect portion shows a granular overlay, and an encrusting

sponge at its base (Fig.18,A). ITS characteristic of this genus

are present in the coiled parts, and consist of an unserrated

dorsal ridge and a serrated ventral ridge. The dorsal ridge is

transparent, somewhat high, and has a smooth, somewhat

T-shaped edge; it is spiral, on a columella-shaped axis in the

spiral proximal portions of the tube (Fig.18, B,C). The

available portions of the worms were inadequate to assign the

material to any of the other Caribbean species or a new

species.

Spiraserpula vasseuri sp. nov.

(Figs.20, A-H; 21, A-K; 3, J)

Helmut Zibrowius requested (pers. comm.) that the material

on which he based his preliminary description of the present

species in an unpublished manuscript be examined, and that

it be included in this paper if it belonged to the present group.

ITS are indeed present in this species, and its description

follows:

MATERIAL EXAMINED.

Europa Island (Mozambique Channel): North Reef, Gabriel

Cove grotto, 55 m, on oysters, legit Pierre Vasseur, scuba

diving, 28.xii.1965 (HOLOTYPE: USNM 46475, 6

PARATYPES USNM 46476).

TYPE LOCALITY. Europa Island.

DESCRIPTION.

TUBES: The colour is mostly whitish, with a very faint pinkish

to orangish tinge seen in places at certain angles of illumina-

tion. They are sinuous, coiled and bonded together, espe-

cially at their bases. A granular overlay is present. The

anterior portions are squarish to trapezoidal in cross-section

(Fig.20, A-C). The dorso-lateral angles may be incompletely

developed in places and represented by a pair of incomplete

longitudinal ridges; an additional incomplete ridge may be

Fig. 19 Spiraserpula sp. A-C, Three specimens from Curagao. A, Erect part of tube with encrusting sponge on its base. B, A tube opened to

show spiral dorsal ridge in its first formed coil. C, Aggregation of tubes opened to show variations of dorsal ridge.

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 79

Fig. 20 Spiraserpula vasseuri sp. nov. A-H, paratypes. A—-C, Anterior tube fragments showing granulations, transverse ridges and roughly

trapezoidal external outline. A, Shows incompletely formed external longitudinal ridges, and a thickened peristome towards the posterior

end of B. D & E, Two views of same posterior coil showing the unserrated dorsal ridge in both, and a serrated ventral ridge consisting of

separate teeth in D. F & G, Two views of same anterior tube fragment with an attached posterior coil opened (on the right). The anterior

tube fragment has two peristomes; the posterior coiled part (G)has been opened to expose the unserrated dorsal ridge and serrated ventral

ridge (G). H, Thorax showing prostomial ocelli, collar, thoracic membranes and chaetigers.

present along each flank (Fig.20, A). Transverse ridges are

present, which may be thickened in places, representing

peristomes (Fig.20, B,F,G). Although broken into fragments

during collection, total lengths appear to have been between

30-40 mm, and their maximum external diameters up to

about 3.0 mm. Their fractured ends show two concentric

layers of different consistency and thickness, an inner one

that is more vitreous and transparent than the outer which is

white and opaque. Their lateral margins are fragile and

chambered, with thin walls.

The posterior ends of the tubes are coiled. ITS consist of a

low unserrated dorsal ridge, and a serrated ventral ridge

(Fig.20, D-G). The latter may be represented by a row of

separate teeth (Figs.20, D; 3, J).

WORMS: The total length of the worms, based on the frag-

ments, exceeds 15.0 mm. The branchial crown is 4.0-5.0 mm

long, and each side bears 8-10 radioles and an operculum or a

rudimentary operculum. The opercular peduncle is long and

slender, of the same thickness as the radioles. One of the

specimens has a well-developed operculum on one side, and

another, much smaller, but similar operculum on the other.

The operculum is big and short, massive, bell-shaped, and

slightly concave distally. The radii end in large, rounded

marginal lobes, and range from 10 to 15 in number (10 in 1,

11 in 3, 12in 1, and 15 in 1). The number of thoracic segments

per side varies from 10 to 14. A pair of small ocellar clusters is

present. Collar large, roughly divided into three large ventral

lobes and a pair of latero-dorsal lobes. Thoracic membranes

are broad up to the third segment, after which they narrow,

and do not form an apron.

The longest opercular peduncle (holotype) together with

its operculum is 5.5mm long. The operculum (Fig.21,

A,C,D) is separated from the peduncle by a faint constric-

tion, where the peduncle is only 1/2—1/3 the diameter of the

base of the operculum. The variations in the dimensions of

opercula of the older specimens are as follows: length:

0.6-0.7 mm; width: 0.55—0.6 mm. They are bell-shaped, with

a small shallow concavity distally. They have a thick and

transparent cuticle (Fig.21, A,C,D). The second radiole of

the opposite side is modified into a rudimentary operculum

(Fig.21, B). The radioles end in short pinnule-free tips which

are about 1/10—1/15th the total length of the radioles (Fig.21,

A-C).

One of the specimens, a juvenile, provides an indication of

the possible ontogenetic changes in the operculum of this

species. Unlike in the adults, where peduncle and operculum

are markedly separated from each other, the slender

peduncle of the juvenile merges gradually into the base of the

operculum. In addition, the shape of the latter is an elongated

funnel, and its distal end is convex (Fig.21, E).

The collar fascicles may bear up to about 5 fully formed

bayonet chaetae and one developing deep within. Each

possesses a long serrated blade, a short unserrated notch, and

several moderately large teeth on the basal boss (Fig.21,

F-K). Thoracic uncini show 5-6 teeth in side view; however,

in oblique edge view it is evident that they are saw-rasp

shaped, with an anterior single row and a posterior cluster of

teeth (Fig.21, L). This is more clearly seen in the anterior

Fig. 21

T.G. PILLAI AND H.A. TEN HOVE

abdominal uncini (Fig.21, M). In side view, the number of

teeth in the latter vary from 4 or 5 towards the lateral end of

the torus to 7 at the dorsal end. Flat trumpet chaetae number

9-11 per bundle. Their distal ends terminate in a slender

hook-shaped process on one side and are drawn out into an

acute angle on the other (Fig.21, N).

ETYMOLOGY. As suggested by Zibrowius (pers. comm.), the

species is named after its collector, P. Vasseur.

HABITAT AND DISTRIBUTION. A reef dweller found on oyster

shells in submarine caves at depths of around 55 m. Hitherto

collected only from the Mozambique Channel.

Spiraserpula deltoides sp. nov.

(Figs.22, A—N; 3, C)

MATERIAL EXAMINED.

Lesser Sunda Islands, Sumba (Indonesia): Snellius II 4.051,

NE coast of Sumba, E. of Melolo 09°53.5’S 120°42.7’E,

75-90 m. (HOLOTYPE & 1 PARATYPE (empty tube):

RMNH 18296; 3 PARATYPES: ZMA V. Pol. 3736;

2 PARATYPES: BM(NH) 1992.37 & 38).

TYPE LOCALITY. Sumba (Indonesia).

DESCRIPTION.

TUBES: White, small, and spirally coiled upon themselves.

They are squarish in cross-section, smooth and rounded

dorso-laterally, and with a shallow longitudinal depression in

between (Fig.22, A). They have an extremely fine granular

overlay, which can only be seen at certain angles of illumina-

tion, and very fine transverse grooves. The coil diameter is

generally about 3 mm, maximally 9 mm; the maximum exter-

nal tube diameter is generally 0.7 mm, maximally 1.3 mm. In

two of the tubes an inner transparent lining was observed.

ITS consist only of a serrated dorsal ridge along the convex

wall of the tube (Figs.22, B,C; 3, C). The serrations are

delta-shaped, mostly separate, and opaquely white in colour.

WORMS: The holotype (Fig.22, D) is 5.0 mm long, 0.35 mm

wide in the thorax and its abdomen is 3.2 mm long. One

paratype is incomplete posteriorly, the other is 8.0 mm long,

with an abdomen of 3.5 mm. Some measurements and counts

are given Table 17:

The operculum is bell-shaped, with a shallow distal concav-

ity extending inwards as far as the inter-radial grooves. The

radii end in rounded marginal lobes, the constriction between

operculum and peduncle is sharp, and the diameter of the

distal end of the peduncle is about 1/2-3/4 that of the

proximal part of the operculum (Fig.22, D—G).The rudimen-

tary operculum is 1.5 mm long, thread-shaped. The radioles

end in short pinnule-free tips, about 1/7 the total length of the

radioles. Two clusters of prostomial ocelli are present. It is

difficult to determine whether ventral thoracic glands are

present. Thoracic membranes do not extend to the end of the

thorax, but exactly where they end cannot be located, it may

be at the 7th chaetiger in one paratype. The abdomen of the

holotype has about 67 segments, with capillaries on the last 8

or 9; the complete paratype has 85 abdominal segments, 24

with capillaries. The abdomen of the incomplete paratype is

Spiraserpula vasseuri sp. nov. A, Holotype. B-N, Paratypes. A, The left branchial crown and three views of the operculum and its

slender peduncle. B, Left branchial crown and rudimentary operculum from another specimen. C & D, branchial crowns and differrent

views of the opercula of two other specimens. E, Two views of the convex operculum of a juvenile. F-K, Bayonet collar chaetae bearing

several teeth on the basal boss, and a short unserrated notch. L, Thoracic uncini, with more than one row of teeth towards their posterior

ends. M, Anterior abdominal uncini. N, Bundle of anterior abdominal chaetae with flat trumpet-shaped ends.

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961

Table 17S. deltoides sp. nov. Some measurements and counts on

the holotype and 2 paratypes.

Holotype Paratype Paratype

1 6

Length of operculum and peduncle 0.94mm 1.2mm 3.2 mm

Length of operculum 0.38mm 0.52mm 0.48 mm

Diameter of operculum 0.44mm 0.35 mm _ 0.41 mm

No of opercular lobes 22 26 22

No. of radioles (L/R) 5/6 6/6 17

No. of thoracic chaetal tufts (L/R) 8/7 6/8 77

12.5 mm long, has about 75 segments, and the latter bear

reddish-brown granular material ventro-laterally.

The collar fascicles bear 2-6 bayonet chaetae. Each has a

long serrated blade, a short unserrated notch, and about 2-6

teeth on the basal boss (Fig.22, H—I, L—-N). Thoracic uncini

bear about 5 teeth in a single row (Fig.22, J,K); anterior

abdominal uncini are similar and bear 5-7 teeth.

ETYMOLOGY. The specific name refers to the delta-shaped

serrations of the internal dorsal ridge.

HABITAT AND DISTRIBUTION. Found on calcareous stones at

depths of 75-90 m. Hitherto collected only from Sumba

(Indonesia).

Spiraserpula sumbensis sp. nov.

(Figsa23,7A—U;; 3, Et)

MATERIAL EXAMINED.

Sumba (Indonesia): Snellius II 4.051, NE coast of Sumba, E

of Melolo, 09°53.5’S 120°42.7’E, 75-90 m, (HOLOTYPE:

RMNH 18297; 1 PARATYPE: ZMA V. Pol. 3737; 1

PARATYPE: BM(NH) 1992.72).

TYPE LOCALITY. Sumba (Indonesia).

DESCRIPTION.

TUBES: White to very faintly pinkish. A small species with

external tube diameter only up to about 0.5 mm, and a lumen

of about 0.25 mm wide. A granular overlay consisting of

extremely fine granules can be seen under special illumina-

tion. Tubes are circular in cross-section and bear faint trans-

verse wrinkles (Fig.23, A—C, O).

ITS consist of a dorsal ridge and a ventral ridge, which are

both unserrated, wedge-shaped in cross-section (Fig.3, H),

and partially divide the lumen into somewhat asymmetrical

left and right halves (Fig.23, C,D). The two ridges are light

Table 18S. sumbensis sp. nov. Measurements and counts.

Paratype Paratype

1 2

Left side Right side

Length of op. & peduncle (mm) 1.2 1.0 1.2

Length of operculum (mm) 0.36 0.36 0.35

No. of lobes 19 21 i19/

No. of radioles 5) 5) 4/4

No. of thoracic chaetal tufts 8 8 -

Thoracic membrane ends

(Fig.23,Q) 3 5 Ps

T.G. PILLAI AND H.A. TEN HOVE

pink and opaque. In cross-section they consist of a lens-

shaped whitish kernel in the inner hyaline tube layer; the

outer tube layer is opaque.

WORMs: The holotype (Fig.23, E-G), has a total length of

7.0 mm, thoracic width of 0.26 mm, an abdominal length of

5.1 mm and 66 segments, with capillaries on the last 17. The

length of the operculum plus peduncle is 1.3 mm, the length

and diameter of the operculum 0.38 mm and 0.26 mm,

respectively. The operculum is zygomorphic (Fig.23, E,F). It

has a distal concavity which extends as far as the inter-radial

grooves. The 15 radii end in somewhat acutely triangular

marginal lobes with smooth tips. The peduncle is slender, but

somewhat expanded before the constriction below the oper-

culum. There are 5 radioles on each side, with the operculum

on the left side and a short filamentous rudimentary opercu-

lum on the right (Fig.23, E). The short pinnule-free tips are

about 1/7-1/8 the total length of the radioles. Thoracic

chaetal tufts number 8 on each side. The thoracic membrane

ends on the fifth chaetiger on the left, but it is difficult to

determine its extent on the right. One tiny prostomial eye

appears to be present on the right side, the left side is

damaged. Thoracic glands could not be detected in the

material.

One paratype (Fig.23, O—Q) has an incomplete abdomen.

It is the first specimen encountered in this genus with two

equally well-developed opercula (Fig.23, O,P). The thorax of

the second paratype is missing, the remaining abdomen has

54 segments, 12 of them with capillaries. Some measurements

and other data are given in Table 18:

The paratypes agree with the holotype with regard to the

tube, operculum, radioles, and chaetae. The opercula are

somewhat zygomorphic.

Collar fascicles bear 4 fully formed bayonet chaetae in the

holotype; 3 fully formed bayonet chaetae and a newly formed

one deep within the fascicle in paratype 1. Each bayonet

chaeta (Fig.23,H-K, R—U) consists of a long serrated blade, a

moderately long unserrated notch, which is about 1/6-1/7 the

length of the blade, and several teeth on the basal boss.

Thoracic uncini appear to have a row of 7-9 teeth in side

view, but more than one row as seen in edge view (Fig.23,L).

Anterior abdominal uncini are similar, but appear to have

fewer teeth in side view (Fig.23, M). Flat trumpets number

four in each anterior bundle, their curved distal ends have a

poorly developed hook on one side and are comparatively

elongated on the other (Fig.23, N). Capillaries occur in the

posterior 12-17 chaetigers.

ETYMOLOGY. Named after the type locality.

HABITAT AND DISTRIBUTION. Found on calcareous stones at

depths of about 75-90 m. Hitherto collected only from

Sumba (Indonesia).

Spiraserpula iugoconvexa sp. nov.

(Figs. 24, A-K; 25, A—Q; 3, I)

MATERIAL EXAMINED.

NE Flores Sea to SW Banda Sea (Indonesia): 1. Taka Bone

Rate(Tiger Islands), Snellius II 4.139B, S of Tarupa Kecil,

06°30’S 121°8’E,depth -30 m, (HOLOTYPE: RMNH 18295;

PARATYPE I: ZMA V.Pol.3735). 2.Tukang Besi Island,

Binongko, Snellius II 4.044B, SW of Taipabu, Banda Sea,

5°56’S 123°58.5'E, down to 25 m, (PARATYPE II: BM(NH)

1992.39). Queensland (Australia): 3. Lizard Island, S. South

Island, sloping silty reef, little coral cover, legit H. A. ten

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 83

«¥

uncini. L-N, Bayonet chaetae.

Hove et al., Stn. 21,6.ii1.1986 (1 specimen, tube; AM,

W21676).

TYPE LOCALITY. NE Flores Sea (Indonesia).

ESCRIPTION.

TUBES: Bright rose, red in fresh material, with a translucent

granular overlay, coiled posteriorly but not anteriorly. All

three tubes of the type series were partially overgrown by

encrusting bryozoans, making their surfaces irregular. At

irregular intervals, there are also peculiar elongated struc-

Se Sy jem

Fig. 22 Spiraserpula deltoides sp. nov. A-E & H-K, From holotype; F-G & L-N, from paratype. A, Tube viewed from above. B & C, Two

views of same tube opened substratally to show deltoid dorsal ridge. D, Worm, showing operculum and dorsal abdominal groove. E,

Anterior view of operculum. F & G, Two views of operculum. H & I, L-N, Bayonet chaetae. J, Thoracic uncini. K, Anterior abdominal

tures with a semilunar opening, which appear to be a hydro-

zoan commensal, akin to Protulophila Rovereto, 1901 (vide

Scrutton, 1975). These sparsely occurring structures are

directed longitudinally or transversely as shown on the speci-

men from from Stn. 4044B (bottom end of Fig.24, A). The

anterior end of the tube from Tukang Besi Island is not

attached to the substratum. Viewed dorsally, it is squarish to

trapezoidal in cross section. It has two fairly distinct dorso-

lateral ridges and a faint median one in places, transverse

wrinkles which are occasionally thickened, and an expanded

peristome at its anterior end (Fig.24, A). The ventral side of

the unattached part, which commences from a swollen attach-

ment to the substratum, does not show the granular overlay

but only faint transverse wrinkles (Fig. 24, B). The peristome

consists of a broad triangular dorsal lobe which is continuous

with two narrow ventro-lateral lobes (Fig.24, A,B).The

inside of the tube is quite shiny. It attains a diameter of

1.6 mm at the peristome, and 1.5 mm at the swollen com-

mencement of the unattached part.

ITS, which are present only in the posterior part of the tube,

consist of an unserrated dorsal ridge (Figs.24, C; 25, B,C), anda

very short smooth ventral ridge (Fig.3, I), which is very short in

the holotype (Fig.25, D); in the specimen from Lizard Island the

ventral ridge is smooth to scalloped. The dorsal ridge may be

T-shaped in cross-section in places (Fig.25, C) but appears

irregular if damaged (Fig.24, C, middle). In the middle region of

the tube, tear-shaped depressions are present in the inner wall,

up to 0.2 mmin size.

WORMS: The holotype from Taka Bone Rate, broken in three

parts (Fig.25, E-G), has a total length of 31.5 mm, a thoracic

width of 0.5 mm, an abdominal length of 27.0 mm and 117

segments, with capillaries on the last 8. Its radioles are

2.7-3.0 mm long, of which the slender pinnule-free tips consti-

tute 0.3 mm. Paratype I lacks its branchial crown; it has a length

of 5.0 mm, a thoracic width of 0.4 mm, an abdominal length of

4.3 mm, and has 61 segments, with capillaries on the last 15 or16.

Paratype II, from Tukang Besi Island, lacks its radioles on the

right side, and its abdomen is in several parts. Its thorax,

however, is intact (Fig.24, G,H).

The operculum and peduncle measure 3.0 mm long in the

holotype, 4.1 mm in paratype II. Other measurements and

counts are given Table 19.

The operculum is zygomorphic, and its distal end is quite

different from that of other known species of the genus in being

markedly convex (Figs.24, D-F; 25, G—J). The cuticle is thick-

ened and transparent, particularly in its convex distal end, the

marginal lobes of the radii, and the asymmetrical projection at

the base of the operculum. The number of radial lobes reaches

about a dozen. There is a sharp constriction between the

operculum and the peduncle, the latter being slender, except for

a slight expansion before the constriction (Figs.24, E,F;25,J). A

filamentous rudimentary operculum is present on the side

opposite to that of the operculum (Fig.25, G).

The number of radioles per side reaches 14. Their short

pinnule-free tips are about 1/7—1/8 the entire length of the

radiole. Prostomial eyes were not found. Thoracic glands are

present, transparent in the holotype and paratype II, light

brown in paratype I. The number of thoracic segments per

side is 7-8, and the thoracic membranes do not reach the last

thoracic segment (Fig.24, G—I).

The abdomen of the holotype appears glandular ventrally,

packed with eggs, and bears peculiar swellings (Fig.25, E) which

fit into corresponding depressions in the tube. It was not possible

to find them in the damaged abdomen of paratype II, although

this is a mature specimen too, and the inner tube wall shows

tear-shaped depressions (0.32 x 0.22 mm); they are absent in

T.G. PILLAI AND H.A. TEN HOVE

Table 19 S. iugoconvexa sp. nov. Measurements and other data on

type specimens.

Holotype ParatypeI Paratype II

Length of operculum (mm) 0.7 ? 1.0

Diameter of operculum (mm) 0.5 i 0.7

No. of radii 12 ? 11

No. of radioles(L/R) 10 2 14/?

No. of thoracic chaetal tufts 8/7 7/7 7/7

Thoracic membrane ends 2/? 1/3 5/4

the juvenile paratype I. Possibly, these abdominal swellings are

developed in older worms only.

Collar fascicles of the holotype bear 4 bayonet chaetae

each (Fig.25, K-N). Each possesses a long serrated blade, a

short unserrated notch and two teeth on the basal boss, one

of which may be difficult to observe in side view since it lies

directly behind the other. The number of teeth is clearly seen

in one of the bayonets of paratype I which has its blade

broken off at its base (Fig.24, J), although it is difficult to

observe in a newly formed chaeta from within the same

fascicle (Fig.24,K). Thoracic uncini bear 5 or 6 teeth (Fig.25,

O), and anterior abdominal uncini 4 or 5 teeth in a single row

(Fig.25, P). There are up to about a dozen flat trumpet

chaetae in each bundle (Fig.25, Q). Their distal ends bear a

claw-shaped process on one side and are drawn out into an

acute angle on the other.

ETYMOLOGY. Iugum (L) = yoke; convexus (L) = bulbous;

refers to the zygomorphic, convex operculum.

MATERIAL FROM OTHER LOCALITY. The material from Liz-

ard Island agrees closely with that of the type series with

regard to collar chaetae, operculum and tear-shaped depres-

sions in the inner tube wall. However, the ventral internal

ridge has a scalloped edge, not smooth as in the Indonesian

material.

HABITAT AND DISTRIBUTION. A reef dweller occurring at

depths of about 25 m. Hitherto collected from Indonesia

(Flores Sea and Banda Sea) and Australia (Queensland).

Spiraserpula snellii sp. nov.

(Eigs.26, A-X: 27, ASE: 28, Aq Ve 3.F)

MATERIAL EXAMINED.

Flores Sea, (Indonesia): 1. Taka Bone Rate (Tiger Island),

Snellius II 4.139B, S. of Tarupa Kecil, 06°30'S 121°8’E, edge

of reef flat, 30 m, (HOLOTYPE & 4 PARATYPES: RMNH

18298; 4 PARATYPES (+ one abdomen & internal tube

ridge): BM(NH) 1992.66-71; 5 PARATYPES & tube mate-

rial: ZMA V. Pol. 3738; 3 PARATYPES & fragmentary tube

material: USNM 130983 & 130984).

Fig. 23 Spiraserpula sumbensis sp. nov. A-M, Holotype. O-U, Paratype. A-D, O, Tubes showing granular overlay and faint transverse

growth ridges; A, An erect part; B, also showing body, operculum & radioles in situ; C & D, two views of same tube fragment showing

wedge-shaped dorsal and ventral ridges, both unserrated. E, Operculum. F & G, Holotype. F, Radioles showing zygomorph operculum on

left, rudimentary operculum on right. G, Worm showing extent of thoracic membrane. H-K, Bayonet collar chaetae. L, Thoracic uncini.

M, Anterior abdominal uncini. N, Bundle of anterior abdominal flat trumpet chaetae. O, Tube of paratype, also showing worm with its two

opercula in situ. P, Radioles with two well-developed opercula. Q, Two views of thorax showing extent of thoracic membranes. R-U,

Bayonet collar chaetae.

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961

T.G. PILLAI AND H.A. TEN HOVE

- mm

a 3 5) 1

Vane

Wes = “H

{ een i)

L rife Nake CS

\ fe i ay

”

Fig. 24 Spiraserpula iugoconvexa sp. nov. A-K, Paratypes. A, Dorsal view of erect tube part showing longitudinal external ridges, granular

overlay, transverse ridges, triangular dorsal lobe of its aperture, and an unidentified transverse epibiont at its base. B, Ventral view of same

tube showing the two small ventral lobes of the aperture, rounded ventral side and fine transverse growth ridges. C, Posterior coil of tube

showing a damaged dorsal ridge. D, Radioles and operculum of the right side. E & F, Same operculum showing its zygomorphy and convex

distal end. G & E, Dorsal and ventral views of thorax showing the collar and extent of the thoracic membranes. I, Juvenile paratype. J,

Bayonet collar chaeta, lacking blade, but showing two teeth on the basal boss. K, Newly formed chaeta from within the fascicle.

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961

| Fig. 25 Spiraserpula iugoconvexa sp. nov. Holotype. A, Tube fragment showing irregular surface and peculiar structure with a semilunar

opening (? Protulophila). B & C, Views of the opened tube showing the unserrated dorsal ridge, with a somewhat flatened ridge (C,

bottom left). D, Tube fragment showing ventral ridge. E-G, Entire holotype, in three parts. E & F, Body showing the dorsal longitudinal

groove, the apparently glandular ventral side of the abdomen and its peculiar outpouchings. G, Radioles with operculum on the left and

rudimentary operculum on the right. G-J, Four views of the zygomorph operculum with convex distal end. K-N, Bayonet collar chaetae

with long slender blade, short unserrated notch and two teeth (seemingly one tooth) on the basal boss. O, Thoracic uncini. P, Anterior

abdominal uncini. Q, Bundle of anterior abdominal flat trumpet chaetae.

88 T.G. PILLAI AND H.A. TEN HOVE

Fig. 26 Spiraserpula snellii sp. nov. A, JI-K, N-Q, V-W, Holotype. B—L, R-U, Paratypes. M, Juvenile. A & C, Tubes showing longitudinal

pigment bands, transverse bands and thickenings (A). B, Tube fragment showing unserrated ventral ridge. E-H, tube fragments in their

relative positions to the unopened tube (D) showing the unserrated ventral ridge which is thickened towards the middle of the tube (G,H).

I, Tube fragment showing T-shaped ventral ridge. J-K, Holotype, showing operculum, radioles, and dorsal longitudinal groove along its

body. L, Paratype, juvenile without operculum. M, Smaller juvenile worm, also without operculum. N—-U, Bayonet collar chaetae. V & W,

Thoracic uncini. X, Anterior abdominal uncini.

Queensland (Australia): 2. Lizard Island, N of South Island, Island, S. of light-house; coral heads on sandy bottom, 7 m,

14.4°S 145.3°E, reef front, sloping reef outside of lagoon and legit H. A. ten Hove, 2.iii.1986, Stn.17 (1 specimen in four

sandy bottom below, 10-17m, legit H. A. ten Hove, P. fragments, BM(NH) 1992.65). 4. Lizard Island, S. South

Hutchings and M. Reid, 5.iii.1986, Stn.20 (3 specimens, Island; sloping silty reef, little coral cover, legit H. A. ten

ZMA V. Pol. 3734, AM W20342). 3.Lizard Island, Palfrey Hove et al, Stn.21, 6.iii1.1986 (8+ specimens, ZMA V. Pol.

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961

3830, BM(NH) 1993.17, AM W21677). 5. Boulton Reef, on

scleractinian coral (Thecopsammia regularis Gardiner 1899),

USNM 78572, dry material, legit J. C. Lang, 31.vii.1973. H.

Zibrowius, who identified this, kindly drew our attention to

its serpulid epifauna.

Loyalty Islands, E. of New Caledonia: 6. SW Pacific Lagoon

of Beautemps-Beaupré Atoll; overhang 8 m, on heavily

encrusted dendrophylliid scleractinian coral, scuba diving,

dry material, MUSORSTOM 6 cruise, legit H. Zibrowius,

17.11.1989.

Okinawa (Japan): 7. W. side of Sesoko Island, 2-3 m, on

cliffside, in caves and grooves, scuba diving, on unidentified

coral, legit S. Nakamura, 10.i.1989, dry material, USNM . H.

Zibrowius kindly drew our attention to the serpulid epifauna.

Sinai (Egypt): 8. Strait of Tiran, at Sharks Observatory,

20-25 m; Nos. 210-213, legit H. A. ten Hove, 8.vi.1990 (2

specimens, tubes, HUJ, ZMA V. Pol. 3886).

Elat (Israel): 9. In front of Marine Biological Laboratory,

20-25 m, coral rubble; Nos. 154, a-d, legit H. A. ten Hove,

4.vi.1990. 10. Oil port, S. pier, 6-25 m, coral rubble and

pillars of pier; Nos. 181,244, 311, 339, 340, legit H. A. ten

Hove, 6.vi.1990 (3 specimens, several tubes, HUJ).

TYPE LOCALITY. Taka Bone Rate (Flores Sea, Indonesia).

DESCRIPTION.

TUBES: Mustard coloured, with a pair of darker longitudinal

bands in places along each flank, joined by transverse bands,

especially just anterior to the thickenings found at intervals

(Fig.26, A,C). They may be coiled more or less parallel to

one another in the horizontal plane, mutually bonded

together or spread out on the substratum and branched in

places. Their external diameter is quite small, only up to

about 0.6 mm. Earlier formed portions of tubes may show

narrow transverse wrinkles (Fig.26, B,D). In fresh material

the colour of the tube may be more brownish, and appears to

fade to mustard after a few months in alcohol.

ITS consist of an unserrated ventral ridge only (Fig.26,

B,E-1), which is T-shaped in cross-section towards its middle

(Figs.26,G,I; 3, F), and becomes progressively less thickened

both anteriorly and posteriorly (Fig.26, E,I).

WORMS: The total length of the worms ranges from 2.2 mm in

the case of a juvenile, to a little more that 12.3 mm in an

older individual which lacks its radioles. The complete holo-

type (Fig.26, J) is only 5.8 mm long. The thoracic width in all

the specimens is around 0.3 mm.

An operculum may or may not be present. Younger

specimens have radioles but lack opercula (Fig.26, L,M);

apparently opercula appear only in older worms (Fig.26,

A,J,K). The length of the operculum and peduncle in the

holotype is 1.5mm, the operculum 0.3 mm long and its

diameter 0.2mm. Its distal part is nearly globular

(Fig.26,A,J,K) and, unlike the opercula of the other known

members of the group, its margin is not divided into lobes,

but shows about four pseudo-lobes, apparently caused by

contraction in alcohol. Its proximal part is shaped like a

narrow funnel, separated by a sharp constriction from the

slender peduncle. A short filamentous rudimentary opercu-

lum was observed in one specimen only. It appears likely

that, like the operculum, they are developed in older worms.

Pinnule-free tips of radioles short. Thoracic glands were not

found. Some counts and meristic data are given in Table 20:

The abdominal length in eight specimens ranged between

11.2 and 1.0 mm, and the number of segments between 48

Table 20S. snellii sp. nov. Some meristic and other data of type

series.

No. of specimens (n=6) 3 3

No. of radioles S/S 4/4

No. of specimens (n=8) 2 1 2 De il

No. of thoracic chaetal tufts o/s Omen Tee HON TIS

No. of specimens (n=3) 1 1 1

Thoracic membrane ends 4/4 4/3 3/3

and 22, respectively, with capillaries on the last 6 or 7.

Collar fascicles of older specimens bear about four fully

formed bayonet chaetae and a developing one deep within.

Each bayonet chaeta possesses a long serrated blade, a

moderately long unserrated notch (1/3-1/4 the length of the

entire blade), and several teeth on the basal boss (Fig.26,

N-U; PI.5, F). Thoracic uncini (Fig.26, V,W) and anterior

abdominal uncini (Fig.26, X; PI.5, G) bear 4-6 and 4-5 teeth,

respectively, in a single row. Flat trumpet-shaped chaetae are

typical (P1.5, H).

REMARKS. One single tube revealed 2 specimens: a parent

with schizont closely appressed to its posterior end. Posteri-

orly, the abdomen of the parent was abruptly tapering

(dorso-ventrally), with long capillaries. Lying between those,

the three pairs of radioles of the schizont could be found. It

had a narrow, still not fully developed thorax with 7/6 chaetal

tufts, followed by a well-formed abdomen with 17 chaetigers

(the last 7 with capillaries). The entire schizont was folded

over the ventral internal ridge.

COLLECTIONS FROM OTHER LOCALITIES. The specimens in

sample 2 from Queensland agree with those in the Indonesian

sample with regard to the overall mustard colour. Against

this background colouration there are darker mustard to

brown longitudinal bands, which are variable. One of the

three available tubes has a pair of lateral longitudinal bands,

lacking in places. The second tube has a thin median longitu-

dinal stripe in addition. The third has a pair of mustard yellow

longitudinal bands laterally, and a broad brownish median

band which is partially divided into two bands by a narrow,

yellow longitudinal band.

They are coiled upon themselves either individually or

mutually bonded together. The coils are more or less concen-

tric, low, flattened against the substratum, and bonded

together (Fig.27, A). The maximum external tube diameter is

1.2 mm. The granular overlay consists of a median longitudi-

nal band made up of broad, transverse, forwardly-directed

scutes, and a narrow band of smaller granules laterally

(Fig.27, A). At irregular intervals there are wavy, thickened,

peristome-shaped transverse ridges.

ITS agree with those of the Indonesian specimens. They

consist of only an unserrated ventral ridge. Its edge is smooth

and, in cross-sectional appearance, varies from being wedge-

shaped to thickened and T-shaped at its maximal develop-

ment (Fig.27, B). The cross-bar of the T may also be curved

outwards and bear a shallow longitudinal depression. The

mid-ventral longitudinal abdominal groove is applied to this

ridge.

Three worms were removed from the tubes. One has a

damaged thorax and an incomplete abdomen, while the

second lacks the radioles of both sides, the third is broken in 4

fragments. The former (Fig.27,D), is 0.4 mm wide in the

T.G. PILLAI AND H.A. TEN HOVE

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961

thorax, and has 6 radioles and a short club-shaped rudimen-

tary operculum on each side. The radioles are about 0.4 mm

long and end in short pinnule-free tips. Two clusters of dark

brown prostomial ocelli are present. There are 7 thoracic

chaetal tufts on one side, but the number on the other side

and the extent of the thoracic membranes cannot be deter-

mined due to the damaged thorax.

The first worm is spirally coiled along the substratal plane

(Fig.27, E). The length of the thorax and abdomen is

13.8 mm, of which the posterior portion of about 1.18 mm is

abruptly narrower than the rest of the abdomen. The entire

length of the third worm is about 7.0 mm, radioles 1.3 mm,

thorax and abdomen 5.7 mm; it has 7/6 radioles, and thoracic

width of 0.35 mm.

There are 7 thoracic chaetal tufts on each side in one worm,

6/7 in the other. The thoracic membranes end between

chaetal tufts 6 and 7 on the left, and 5 and 6 on the right

(Fig.27, E,F). Thoracic glands are absent. The number of

abdominal segments is 102, with capillaries on the last 6. A

mid-ventral longitudinal groove traverses the entire abdomen

and thorax (Fig.27, E, F).

Each collar fascicle bears about 4 bayonet chaetae. They

have long serrated blades, a short unserrated notch, and

several teeth on the basal boss (Fig.27, G—J). Thoracic uncini

usually bear 4 teeth in a single row (Fig.27, K). The anterior

abdominal uncini also possess 4 teeth in a single row, but the

3 posterior teeth are not closely appressed, as in the Indone-

sian specimens (Fig.27, L). Flat trumpet chaetae number

about 3 per bundle. Since their edge is curved, details on the

anterior tooth cannot be observed.

The third sample from Lizard Island, Queensland agrees

with the first and the Indonesian specimens in all important

characters (Fig. 27, A-J). The fully formed operculum is an

unlobed funnel with « shallow distal concavity (Fig.28, C,D),

while in the earlier stages it is spherical or nearly spherical

(Fig.28, E-G).

The dry tubes from Loyalty Islands (Fig.28, K-—Q)

appeared to be brownish, but regained the typical mustard

colouration when immersed in alcohol. One fragment shows

branching (Fig.28, K). Internally there is only a smooth

ventral ridge (Fig.28, N-Q), which is clearly T-shaped in

places (Fig.28, Q); it is markedly so and occupies a larger part

of the lumen in some tubes which are comparatively very

thick-walled (Fig.28, N).

The collection from Ras Mohammed, Sinai, consists of

tubes with fragments of worms (Fig.28, R-V). The tubes

have an overall mustard colour, but the anterior portions

(Fig.28, R,S) have pinkish peristomes, and a conspicuous

granular overlay along the lateral borders of the attached

portions (Fig.28, S). The medial overlay is scute-shaped, but

not as prominent as in the first Lizard Island sample. A

smooth ventral ridge is present (Fig.28, T—W), which is

T-shaped in its fully formed condition (Fig.28, T,U,W).

Those details of the worm that still could be observed (collar

chaetae, ends of thoracic membranes) agree with the data

given above.

The samples from Elat, Israel, agree with regard to tube

colouration, the smooth ventral longitudinal ridge and other

important characters. No.154 is a single specimen on a piece

of coral rubble. The granular overlay is translucent in places;

transverse scutes are not seen medially, but this may be

because it is a juvenile. The worm has a total length of

3.6 mm; reddish prostomial ocellar clusters are present; its

thorax is 0.2 mm wide; gills 1.0 mm long, with short pinnule-

free tips; the number of thoracic chaetal tufts L?/R7; its

abdomen 1.9 mm long, with 24 segments and capillaries on

the last 4. There are 3 bayonet chaetae per fascicle, each with

an elongated blade, a short unserrated notch and several

teeth on the basal boss. Anterior abdominal uncini bear 4

teeth in a single row. An operculum had yet to be developed.

However, an operculum was observed in sample 311. A

schizont was separated from sample 244.

ETYMOLOGY. Named after the Indonesian-Dutch Snellius II

Expedition which enabled the second author to collect exten-

sively in Indonesian waters.

HABITAT AND DISTRIBUTION. A reef dweller occurring at

depths of about 15-30 m. Appears to be the most widely

distributed species of the genus. Hitherto collected from the

northern Red Sea, Indonesia (Flores Sea), Australia (Great

Barrier Reef) and W. Pacific (S. Japan to New Caledonia).

Spiraserpula lineatuba (Straughan, 1967)

(Figs.29, A-O; 30, A-M; 3, L; Pl.1, A, C & D, PI1.3,

E-G)

SYNONYMY. Serpula lineatuba Straughan, 1967, pp. 211-212,

Fig.5a-g.

MATERIAL EXAMINED.

New South Wales: 1. Sydney, Long Reef, underside of rocks,

LWS,27.11.1965, legit D. Straughan (HOLOTYPE,

AM4018). 2. Sydney, Long Reef, rocks just below LWS,

Colloroy, Stn. 30, 27.i1.1964 (Topotypical material, 2 speci-

mens and several tubes, AM4019, ZMA V. Pol. 3450,

BM(NH) 1992.51). 3. Norah Head, at foot of light house,

from bottom of tidal pools at low-tide, from undersides of

boulders, legit H. A. ten Hove, 12.iv.1986, Stn.31 (5 out of

several specimens, AM W20340). 4. Split Solitary Island,

rocky island area with corals, algae and little sand, from

ceiling of small cave, 12-19 m, legit H. A. ten Hove, P.

Hutchings and R. Phipps, 26.iv.1986, Stn.36 (18 out of

several specimens, ZMA V.Pol. 3709, USNM_ 130996,

BM(NH) 1992.40-S0, AM W20163, QM, NSMT). 5. South

Solitary Island, S of light house, rocky area, cobbles and

corals, little sand, 12-20 m, legit H. A. ten Hove, P. Hutch-

ings and R. Phipps, 27.iv.1986, Stn.37 (3 out of several

specimens, BM(NH) 1992.52-60).

TYPE LOCALITY. Sydney, Long Reef (Australia).

DESCRIPTION.

According to the original description (Straughan, 1967), the

tube is circular in cross-section, white, with a pair of dark

pink lateral longitudinal stripes, pale pink dorsal surface. The

Fig. 27 Spiraserpula snellii sp. nov. From Stn. 20, Lizard Island, Australia: A, Adult tube showing flattened coil form, granular overlay,

which is scutate medially, granular laterally and has a transverse thickened peristome. B, Aggregation of tube fragments with unserrated

ventral ridge, T-shaped in cross-section. C, Scutate juvenile tube with some transverse thickenings; granular overlay not yet developed. D,

worm showing radioles, rudimentary operculum and collar. E, worm from tube figured in A, showing thoracic membrane, ventral

longitudinal groove. F, Anterior part of latter, showing lack of apron and thoracic membrane ending on the 6th chaetiger on the left side.

G-J, Bayonet chaetae, all from same fascicle. K, Thoracic uncini. L, Anterior abdominal uncini.

total length of the worm ranges from 4.5-6.5 mm. There are

4-6 pairs of branchiae with pinnule-free tips. A hollow

operculum with about 22 lobes is present on one side, with a

pseudoperculum on the other. The collar has a pair of lateral

elongations on the median lobe. The thorax has 9 or 10

segments on each side, and the bayonet collar chaetae have 2

conical processes at the base of the blade.

The holotype (AM W4018) is in very poor condition. When

it was examined by the second author in 1979, the poorly

preserved worm, still within its tube, lacked both an opercu-

lum and a rudimentary operculum, although there appeared

to have been one on one side and none on the other. Other

observations were as follows: A cluster of pigmented ocelli

present at the base of each branchial lobe; bayonet collar

chaetae possess 2 conical teeth, with 1-3 accessory conical

teeth; the anterior abdominal uncini of two types: some with

a single row of teeth, others in which the posterior tooth is

split into two; middle abdominal uncini appeared to possess 7

simple teeth in side view; in edge view, however, four

anterior teeth are single and the rest are rows of 3 minute

teeth each.

However, examination of topotypical material collected on

the same date as the holotype and determined by Straughan

(AM 4019, ZMA V.Pol. 3450) yielded the following addi-

tional data: The tube has a pair of light pink longitudinal

bands (Fig.29, A), not clearly defined dark pink stripes as

mentioned and figured in the original description. It is coiled,

somewhat flattened against the substratum, but the free

surface is rounded. The coils are bonded together. A granular

overlay is present, but it is extremely fine and can only be

seen in places, under special illumination (Fig.29, F). A short

erect portion is present (Fig.29, A,B), with a four-lobed

peristome similar to that of S. massiliensis.

The most important data obtained during the present study

of this topotypical material is that S. lineatuba has ITS. They

consist of an unserrated dorsal ridge along the convex wall

(Fig.29, B,C,E,F), and a serrated ventral ridge along the

opposite wall (Figs.29, D,F; 3,L). The former may be high in

the first formed coil (Fig.29, E, F, bottom left), or low

anteriorly (Fig.29, B,C), and is wedge-, tongue- to somewhat

T-shaped in cross-section.

The worm is 6.5 mm long, its thorax is 0.5 mm wide, and

its abdomen is 4.5mm long. There are 5 radioles and a

slender rudimentary operculum on one side. The median lobe

of the collar has only one forwardly directed process, in

contrast with the original description, indicating that this is a

variable feature. There are 7 pairs of thoracic chaetal tufts,

and the abdomen has 49 segments, with capillaries on the last

19. Two clusters of prostomial ocelli are present and the

thoracic membranes do not extend to the last thoracic chaeti-

gers, but end on the fourth and fifth.

There are 5 bayonet chaetae in each collar fascicle, each

with a moderately long serrated blade, a moderately long

unserrated notch which is 1/3—1/4 the length of the blade, and

2 or 3 conical teeth on the basal boss (Fig.29, G—J; P1.3, E).

In bayonets with two large teeth, a single accessory tooth may

T.G. PILLAI AND H.A. TEN HOVE

be present between them (Fig.29, G,H,J). Thoracic uncini

bear 5 or 6 teeth. As seen in edge view, in the outermost

uncini of the row, 3 to 5 of the anterior teeth are single, while

the remaining teeth are subdivided into 2 or more smaller

teeth which form a short, rasp-shaped posterior cluster

(Fig.29,L). Anterior abdominal uncini are similar (Fig.29,

N). However, SEM of anterior abdominal uncini of another

specimen showed a single row of teeth in edge view (P1.3, F).

It appears, therefore, that both types of uncini may some-

times be present. Posterior abdominal uncini are rasp-

shaped, except for the single anterior tooth. The uncini of the

intermediate region show a transition between the two types.

Flat trumpet chaetae number about 5-7 per fascicle (Fig.29,

N; P1.3, G).

A more complete account of the species, however, was

obtained from numerous well-preserved specimens collected

in 1986 from Split Solitary Island.

TUBES: Have the colouration described above, including the

pair of light to somewhat darker pink lateral longitudinal

bands. They occur in aggregations of a few to numerous

individuals, highly coiled amongst themselves and mutually

bonded together, particularly at their bases (Fig.29, O).

Erect parts are sometimes present, and they may bear

four-lobed peristomes (Fig.30, A). The uncoiled part of one

of the longest tubes measures 26.7 mm; together with its

coiled part it is approximately 30.0 mm long, and its maxi-

mum external width is 1.1 mm.

ITS consist of an unserrated dorsal ridge, a serrated ventral

ridge and, usually, a pair of accessory dorso-lateral ridges

(Figs.29,0,, middle left; 30, B,C;.3, L; Pl.1, Ag@D). The

dorsal and ventral ridges of the tube are applied to corre-

sponding longitudinal mid-dorsal and mid-ventral abdominal

grooves (Figs.30, D-F).

Eighteen worms from Split Solitary Island provided impor-

tant additional data. Measurements and other meristic data

from 8 complete specimens of total lengths ranging between

15.9 mm and 1.3 mm presented in Table 21 show that the

worms can attain two and a half times the length mentioned

in the original description. The maximum number of abdomi-

nal segments counted is 89:

Thirteen complete anterior ends all possess an operculum

on one side, a rudimentary operculum on the other, and 5

pairs of radioles. The pinnule-free tips are about 1/4 the

length of the radioles and are as thick as the pinnules (Fig.30,

D,E). The length of the operculum together with its peduncle

ranges between 0.8 mm in the smallest specimen to 1.6 mm

in the largest; the length of the operculum itself from 0.3 mm

to 0.7 mm, and its diameter from 0.4 mm to 0.6 mm, respec-

tively. All the opercula are zygomorph (Fig.30, D,F), their

distal ends are concave and the radii end in somewhat pointed

lobes. Many of the latter are actually double, the sub-dividing

grooves being only about 1/3 the length of the main interra-

dial grooves which extend to about half the opercular length.

Thus the total number of about 17-23 radii end in about

double the number of marginal lobes (Fig.30, D-F). The

constriction between the peduncle and the operculum is

Fig. 28 Spiraserpula snellii sp. nov. A—J, from Stn. 21, Lizard Island, Australia. K-Q, From Loyalty Is. R-W, from Egypt. A-B, Tube

lacking scutes and granular overlay, but with faint transverse grooves between transverse areas (representing scutes ?). D-E, Same worm

with fully formed operculum. E-G, worms showing early vesicular operculum. H, Bayonet chaetae. I, Thoracic uncini. J, Anterior

abdominal uncini. K, Anterior fragment of a tube showing branching and a peristome. L, Another fragment showing transverse ridges. M,

Juvenile tube. N, Fractured end of a tube showing a thick wall and a T-shaped ventral ridge occupying most of its lumen. O-Q, Tube

fragments with varying form and thickness of the T-shaped ventral ridge. R & S, Anterior tube fragments, R with peristomes. T-W, tube

aggregations with fractured ends showing the T-shaped ventral ridge. W, V, with longitudinal view of ventral ridge.

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961

SEN

=cwiteire

Table 21_ S. lineatuba (Straughan). Measurements and counts from

Split Solitary Island material.

Specimen Total Widthof Lengthof No. of Capillaries

no. length thorax abdomen abdominal on

(mm) (mm) segments

1 15.9 0.6 1Be2 89 13

2 IS.7/ 0.6 12.6 85 13

3 IS 7 0.5 27) 81 12

4 13.9 0.5 10.9 58 ly)

5 13.0 0.5 2) 7/ 81 13

6 10.1 0.5 10.9 58 17

I 9.3 0.5 6.8 78 16

8 eS 0.5 0.4 45 25

Table 22S. lineatuba (Straughan). Number of thoracic chaetal

tufts and extent of the thoracic membranes in specimens from

Split Solitary Island.

No. of specimens (18) 1 a SS Ue |

No. of thor. chaetal

tufts 10/10 10/910/8 9/9 9/8 8/8 8/7 7/7

No. of specimens

(15) el led fl ealig ee 2) al

Thoracic membr.

ends 5 6/4 S/S 5/4 5/3 4/4 4/3 4/2 3/3 3/2

sharp, and the diameter of the distal end of the former varies

from 1/3 to 2/3 that of the base of the latter.

The median lobe of the collar shows one or more anteriorly

directed processes in some specimens, none in others. Up to 8

bayonet chaetae have been counted in a collar fascicle. A pair

of ventral thoracic glands is present (Fig.30, E). The number

of thoracic chaetal tufts on each side varies from 7—10, and

the thoracic membranes end on the 3rd to 6th chaetigers, as

shown in Table 22.

The specimens from South Solitary Island and Norah Head

agree with the above description.

LIVE MATERIAL. No records.

HABITAT AND DISTRIBUTION. The species occurs from the

tidal zone down to about 20 m. It was very abundant on a

ceiling of a small cave at a depth of 12-19 m, forming

aggregations of up to 35mm thick, and _ superficially

resembles S. ypsilon from a similar habitat in the Cape Verde

Islands. It has hitherto been collected only from N.S.W.

Spiraserpula discifera sp. nov.

(Figs.31, A-M; 3, M)

MATERIAL EXAMINED.

New South Wales: Sydney, Long Reef, from undersides of

rocks in and bottom of tidal pools, mats of Serpula rubens

T.G. PILLAI AND H.A. TEN HOVE

Straughan, 29.11.1986, legit H. A. ten Hove and P. Hutch-

ings, Stn. 30 (HOLOTYPE, AM W20390).

TYPE LOCALITY. Sydney, Long Reef (Australia).

DESCRIPTION.

TUBE: Pink, with whitish lateral attachment areas and very

fine transverse wrinkles. The median tube parts are of a paler

pink colour than the medio-lateral parts, in fresh material. A

fine granular overlay is present, which can be seen at certain

angles of illumination. The lateral borders of the tube are

glassy and transparent. Irregularly laid along the outer sur-

face of the tube, and more or less perpendicular to it, are

small semilunar to crescentic discs (Fig.31, A-C,E). They are

very thin, pink, glassy and transparent, and their axes are at

various angles to the longitudinal axis of the tube. Some of

them are even attached to the substratum just outside the

tube (Fig.31, B). The maximum external diameter of the tube

is 0.85 mm.

ITS consist of a serrated ventral ridge along its concave

wall (Fig.31, E), and a smooth dorsal ridge. In addition, pink

disks are found on the inside too, on either side of the

serrated ventral ridge (Figs.31, D; 3, M). In some cases the

discs appear to be through and through the wall. The

mid-ventral longitudinal groove of the abdomen (Fig.31, F) is

applied to the serrated ventral ridge. The worm appears to

have a remarkable ability to adjust its abdominal segments in

relation to these sharp discs within the tube.

WORM: Although only one specimen is available, it is com-

plete (Fig.31, F). Its total length is 7.7 mm, thoracic width

0.56 mm; the abdomen is 6.6 mm long and has about 56

segments, the last 20 with capillaries. There are 6 radioles and

a rudimentary operculum on each side. A cluster of blackish

prostomial ocelli is present at the base of the radioles on each

side. There are 8 pairs of thoracic chaetal tufts. Where the

thoracic membranes of the two sides end precisely is not clear

since the thorax is highly contracted (Fig.31, F,G), but an

apron is absent. No thoracic glands were discernible.

The number of bayonet chaetae, 6 in each collar fascicle, is

high in relation to the size of the worm. Their serrated blades

are moderately long, the unserrated notch is about 1/3 the

length of the blade, and there are only 2-4 teeth on the basal

boss (Fig.31,H-M). A few small accessory teeth may be

present. Thoracic and anterior abdominal uncini bear about 6

and 5 teeth, respectively, in a single row.

ETYMOLOGY. diskos (Gr.) = discus; pherein (Gr.) = to

carry.

LIVE MATERIAL. Animal is orange in colour, with transpar-

ent branchiae.

HABITAT AND DISTRIBUTION. S. discifera occurs intertidally

on rocks. It has hitherto been collected only from Sydney.

Fig. 29 Spiraserpula lineatuba (Straughan, 1967). A-N, From topotypical material, Straughan’s original collection, NSW, Long Reef,

AM4019, ZMA V. Pol. 3450. O, From NSW, Split Solitary Island. A & B, Two views of same coiled tube with an erect part ending in

peristome, with longitudinal colour bands in A. B & C, Same tube with posterior coils opened to show the dorsal ridge. D & F, Tube

fragments with serrated ventral ridge. E, Posterior tube fragment with unserrated dorsal ridge. G-K, Bayonet collar chaetae. L, Thoracic

uncini with more than one row of teeth posteriorly. M, Anterior abdominal uncini. N, Bundle of flat trumpet chaetae from same abdominal

segment. O, Aggregation of tubes showing serrated ventral ridges along concave walls, unserrated dorsal ridges along convex walls, and

accessory dorso-lateral ridges (bottom left).

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961

96 T.G. PILLAI AND H.A. TEN HOVE

Fig. 30 Spiraserpula lineatuba (Straughan, 1967). A-M, From NSW, Split Solitary Island. A, Erect tube part showing fine transverse

wrinkles and longitudinal colour bands. B & C, Views of same posterior coil, opened, exposing unserrated dorsal ridge. D & E, Two views

of the same worm, and F, another worm, showing zygomorph operculum, extent of thoracic membranes, and dorsal and ventral abdominal

grooves. G-M, Bayonet chaetae showing variations in the teeth on the basal boss.

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 97

| Fig. 31 Spiraserpula discifera sp. nov. From holotype. A, Fractured tube showing transverse wrinkles, granular overlay, and characteristic

sharp crescentic discs on outer surface. B, Tube fragment, with one of the discs fixed to the substratum, adjacent to the tube. C & D,

Opposite halves of tube fragment split open to show the ITS: a serrated ventral ridge (in longitudinal view), and a lateral row of transparent

crescentic discs. E, Tube fragment with internal serrated ventral ridge and external crescentic discs. F, Worm, showing ventral abdominal

groove. G, Anterior part of worm showing thorax, collar and thoracic membrane. H—M, Bayonet chaetae, all from same fascicle. N,

Thoracic uncinus. O, Abdominal chaetae. P, Anterior abdominal uncini. a bl

98 T.G. PILLAI AND H.A. TEN HOVE

acids

Fig. 32 Spiraserpula minuta (Straughan, 1967). A-Q, From Port Douglas, N. Queensland. A, Erect part of tube showing granular overlay.

B & C, Substratal view of tubes opened to show ITS: an unserrated dorsal ridge along the convex wall, and a serrated ventral ridge

opposite. D & E, Complete worm showing filamentous rudimentary opercula, extent of thoracic membranes and dorsal and ventral

longitudinal abdominal grooves. F, Branchial crown and rudimentary operculum of right side from another worm. G, Two views of the

anterior part of a worm fixed outside the tube, collar and thoracic membranes. I-M, Five bayonet chaetae from one fascicle. N, Thoracic

uncini. O, Anterior abdominal uncini. P, Uncini from a torus middle abdominal region. Q, Posterior abdominal uncini.

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961

Spiraserpula minuta (Straughan, 1967)

(Figs.32, A—-Q; 3, N)

Synonymy: Pseudoserpula minuta Straughan, 1967, p.216,

Fig.6, h-l.

MATERIAL EXAMINED.

Queensland (Australia): 1. Port Douglas, sheltered side of

rocks, near LWM, legit D. Straughan, 17.i1.1963 (HOLO-

TYPE AM W4062). 2. Same locality and date, legit D.

Straughan: (25 studied out of numerous specimens, AM

W4059).

TYPE LOCALITY. Port Douglas, Queensland (Australia).

DESCRIPTION. According to the original description, the

tubes are white, round, and may have a pink tinge. An

operculum or pseudopercula (=rudimentary opercula) are

absent, there are 5 pairs of radioles, the number of thoracic

chaetigers is 7 or 8, and bayonet collar chaetae have 2 or 3

blunt teeth on the basal boss (Straughan, 1967). Vide discus-

sion on the taxonomy which follows this description.

The holotype (AM W4062) was examined by the second

author in 1979. Upon removal of the worm from its tube, it

lacked a branchial crown and its abdomen consisted of 48

segments, the last 6 with capillaries. However, the present

study revealed that AM W4059, which was collected from the

same locality and on the same date by Straughan, contains

several well-preserved specimens, and the following descrip-

tion is based on 25 of them.

TUBES: Whitish in juveniles, with a very faint overall pinkish

tinge in adults but, unlike S. /ineatuba (Straughan), lacking a

pair of pink longitudinal bands. They occur in aggregations of

a few to several individuals which are mutually bonded at

their bases (Fig.32, C). Their anterior ends are often free and

erect (Fig.32, A). A very fine granular overlay is present

(Fig.32, A).

ITS, located in the first formed parts of the tube, consist of

an unserrated dorsal ridge, which is somewhat T-shaped in

cross-section, along its convex wall, and a serrated ridge

along the opposite side (Figs.32, B,C; 3, N). The posterior

end of the abdomen often shows a short mid-dorsal longitudi-

nal groove, into which the unserrated dorsal ridge of the tube

fits (Fig.32, D). The serrated ventral ridge of the tube fits into

a mid-ventral longitudinal groove in the abdomen (Fig.32,

D).

WORMS: Out of the 25 specimens only 11 are complete. Three

have total lengths of over 10 mm, three between 8.0 and

10.0 mm, and five between 6.4 and 8.0 mm. Measurements

and meristic data of the longest and two smallest specimens

are presented in Table 23:

Quite in contrast to the original description, nineteen

specimens with complete anterior ends all possess a pair of

Table 23 S. minuta (Straughan). Measurements and meristic data

of three specimens.

| Specimen Total Thoracic Lengthof No. of Capillaries

no. length width abdomen segments on

(mm) (mm)

1 1347) 0.5 11.3 82 11

2 7.5 0.5 5.5 48 8

3 6.5 0.5 Sy 78 10

Table 24 S. minuta (Straughan). Meristic and other data.

No. of specimens (n=19) 2 14 3

No. of radioles GSS) Seas

No. of specimens (n=20) ee eee ae ee flee

No. of thor. chaetal tufts 9/8 9/7 8/8 8/7 8/6 7/7 7/6

No. of specimens (n=17) 2-3 Siw oneal

Thor. membranes end 6/5 6/4 5/4 4/4 4/3

rudimentary opercula (Fig.32, D-F). A fully formed opercu-

lum is absent. The length of the radioles,ranges between

1.0 mm and 1.3 mm, and they end in short pinnule-free tips

which are about as long and as thick as the pinnules (Fig.32,

D-F). Some meristic data on the population are given Table

24.

The thorax is somewhat wider in specimens that had been

accidentally removed from their tubes prior to fixation

(Fig.32, G,H). Two clusters of prostomial ocelli are present.

Ventral thoracic glands were not discernible.

The numbers of bayonet chaetae in 8 collar fascicles from

different specimens, including a developing one deep within

are: 4in 1,5 in 6, and 6 in 1. Their blades are moderately long

and faintly serrated. The unserrated notch is about 1/3 the

length of the blade. The tooth counts in the above 40 bayonet

chaetae are: 3 in 18 (Fig.32,1,L), 4 in 17 (Fig.32, J,M), 5 in 1

(Fig.32, K), and indeterminate in the remaining 4. The usual

number of teeth is, therefore, 3 or 4. In some chaetae two

teeth may be large, while the third is much smaller (Fig.32,I).

Thoracic uncini (Fig.32, N) bear 4-5 teeth in a single row.

Anterior abdominal uncini (Fig.32, O) are similar, with 4-6

teeth. Posterior abdominal uncini bear 1-3 teeth in a single

row anteriorly, followed by a rasp-shaped cluster of smaller

teeth posteriorly (Fig.32, Q). In the intermediate region

there is a gradual reduction of the number of anterior teeth in

the single row and a corresponding increase in the posterior

cluster (Fig.32, P).

ETYMOLOGY. Renamed after its discoverer, D. Straughan.

HABITAT AND DISTRIBUTION. S. minuta occurs in shallow

water, where it may form ‘dense mats on the sheltered side of

vertical rocks near L.W.M.’ (Straughan, 1967).

It has hitherto been reported only from the type locality,

Port Douglas, Queensland.

DISCUSSION

Spiraserpula Regenhardt 1961 differs from the remaining

genera of its clade, namely Serpula Linnaeus 1758, Hydroides

Gunnerus 1768 and Crucigera Benedict 1887, with regard to

an important character, namely, the presence of ITS. In

addition, the worm lacks an apron. The tubes of the other

three genera lack ITS and, with a few exceptions, their worms

possess an apron.

Straughan (1967) erected the the genus Pseudoserpula for

P. rugosa (type species) and P. minuta, believing an opercu-

lum to be absent in both, and distinguished between them on

the grounds that the former possessed a pair of pseudoper-

cula (= rudimentary opercula) which were said to be absent

in the latter. Ten Hove and Jansen-Jacobs (1984:162-165)

100

synonymized Pseudoserpula rugosa Straughan, 1967, with

Crucigera inconstans Straughan, 1967, stating that the type of

Pseudoserpula is a pseudoperculate individual of C. incon-

stans, but the evidence was incomplete. Moreover, there still

remained the problem of the actual generic identity of P.

minuta Straughan, 1967. Hence it was considered necessary

to re-examine the types of P. rugosa and P. minuta and

compare them with other collections, including those of

Crucigera inconstans.

The holotype of the nominal species Pseudoserpula rugosa

(AM W4027) yielded the following data: The tube is white,

2.0 mm in external diameter, and has conspicuous transverse

wrinkles (Fig.33, A—C). It lacks ITS. An operculum is absent, but

a rudimentary operculum is present on each side (Fig.33, C). An

apron is present (Fig.33,B). The bayonet chaetae typically

possess two conical teeth on the basal boss, as seen in developing

chaetae deep inside the fascicle (Fig.33,H,I); one of the conical

teeth may be smaller than the other or abraded in the older

chaetae (Fig.33, D-F). A short, unserrated notch is present; the

chaetal shaft is smooth below the teeth.

Additional material of Crucigera inconstans (NSW, Sandy

Beach, 21 km N. of Coffs Harbour, legit H. A. ten Hove,

27.iv.1986, Stn. 38 [8 specimens, ZMA V. Pol. 3741] and

Sydney, Long Reef, intertidal rockpools, legit H. A. ten

Hove and P. Hutchings, 29.iii.1986, Stn 30. [1specimen,

ZMA V. Pol. 3740]) gave the following data: The tube is

smooth and has transverse wrinkles (Fig.33, J). Only three

specimens, two from Sandy Beach and one from Long Reef,

possess opercula; the remaining five only rudimentary oper-

cula. The opercula (Fig.33, J-L, S—-V) agree fully with

Straughan’s description and figures of C. inconstans. The

bayonet collar chaetae (Fig.33, M—P) agree with those of the

holotype of Pseudoserpula rugosa. The rudimentary opercula

show different stages of development: both club-shaped, with

somewhat tapering ends in the holotype (Fig.33, C); one long

and tapering, the other more bulbous (Fig.33, W); and a

clearly developing operculum (Fig.34, A). While the holo-

type of Crucigera inconstans has 10 or 11 pairs of radioles

(Straughan, 1967), an operculate individual in our material

shows 15/16. The bayonet chaetae (Fig.34, D—M) are similar,

although they may occasionally possess a small third tooth

(Fig.34, J). A well developed apron is present in non-

operculate and operculate specimens (Fig.34, B), and in the

holotype of P. rugosa (Fig.33, B).

Some of the specimens bearing rudimentary opercula have

regenerating radioles and/or operculum, which appear to have

been nipped off on one side or the other (Figs.33, W; 34, C). In

one of the operculate specimens too some of the radioles are

regenerating (Fig.33, J). It appears, therefore, that opercula and

radioles in Crucigera inconstans are favoured as food by certain

predators. Whether this is the sole reason for a large number of

specimens possessing only rudimentary opercula or not, has to be

determined through further studies. It is worth noting, however,

that the radioles of both sides in the holotype of Pseudoserpula

rugosa are disproportionately small for the size of the worm, and

show every indication of being in a state of regeneration (Fig.33,

B,C).

T.G, PILLAI AND H.A. TEN HOVE

Meanwhile, Pseudoserpula minuta Straughan, 1967, lacks

an apron and has ITS and, therefore, belongs to the genus

Spiraserpula Regenhardt, 1961.

Another nominal genus, Protoserpula Uchida, 1978, needs

to be discussed. Its original description does not mention if

and where any material has deposited. It is not in the

National Science Museum, Tokyo, and other efforts to obtain

it were unsuccessful. Among the generic characters men-

tioned are the following: An operculum is absent, the number

of thoracic chaetigers is 9 or 10, and capillary chaetae are

absent towards the posterior end of the abdomen. The latter

is emphasized in the statement “All the species of Serpula and

its related genera have their long hair-like abdominal poste-

rior segments, but the new genus has not such kind of setae in

abdomen’ (Uchida, 1978: p.23).

The more important characters described under P. paci-

fica, the type species, are as follows: ‘Tube calcareous white

opaque, cylindrical form, attached throughout its length,

curved irregularly . . Operculum absent. Branchiae consist-

ing of 5 pairs of filaments and a pair of palpi. .. The

ventral-most one pair of branchiae are much reduced (0.3mm

long). . . The thoracic membrane developed in the anterior

region but suddenly reduced in width from the 5th segment,

and it becomes to continue to the abdominal body surface

without any structures in the last thoracic segment. Abdomen

. . consisting of about 20 setigerous segments, 2.5 mm long

and 0.2 mm wide. . . Bayonet-shaped seta has a basal process

with about 8 large teeth arranged into a circle... Each

abdominal segment has 1-3 spatulate setae and 8-11 uncini

on one side... The spatulate setae arranged to the last

setigerous segment, and without substitution to the long

hair-shaped setae as occurred in every species in Hydroides,

Serpula, Vermiliopsis, and other many genera’ (Uchida,

1978, p.23—24).

Protoserpula appears to be based on a juvenile serpulid

(ten Hove, 1984, p. 193). A juvenile specimen would not be

sound for erecting a genus since the adult characters could be

different, particularly with regard to the presence or absence

of an operculum.

In the development of operculate serpulids, the operculum

appears after a certain number of radioles have already been

formed (ten Hove 1984, p.183, and Fig. 3). This appears to be

in keeping with the greater importance of feeding and respi-

ration over closure of the tube against predators at this stage.

The dorsalmost pair of radioles remains simple and palp-

shaped, and forms the lateral appendages of the dorsal lip

(mouth palps). In this process they may decrease in size. The

operculum develops as a modification of the second most

dorsal radiole of one side.

As seen from the species of Spiraserpula described in this

account, some possess an operculum, some may or may not

possess one, and others lack one but possess a rudimentary

operculum on each side. Some agree with regard to the

number of radioles, but none of them show palps.

It appears unlikely that rudimentary opercula (=pseudo-

percula) were mistaken for ‘a pair of palpi’ since, in the

same paper, Uchida clearly distinguishes between pseudo-

Fig. 33 Crucigera inconstans Straughan, 1967. A-I, From holotype of Pseudoserpula rugosa Straughan, 1967, with only rudimentary

opercula. A, Anterior part of tube. B & C, Worm within posterior part of tube, showing apron (B), presence of rudimentary opercula

(=pseudopercula, C). D-I, Bayonet chaetae. J-W, Crucigera inconstans Straughan, 1967 from Long Reef, Sydney. J & K, Operculate

specimen within its tube and three views of its operculum. M-R, Bayonet chaetae from same fascicle, M & N newly formed deep within

fascicle. S & T, Two views of small operculum. U & V, Two views of large operculum. W, Anterior end of large non-operculate specimen

with two rudimentary opercula.

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961

101

102

Fig. 34

T.G. PILLAI AND H.A. TEN HOVE

Crucigera inconstans Straughan, 1967. A-M, From Sandy Beach, 21 km from Coff’s Harbour. A, Right radioles of non-operculate

specimen with rudimentary, but developing, operculum. B, Ventral view of large specimen showing apron. C, A large non-operculate

individual with rudimentary opercula. D-M, Bayonet chaetae from the same fascicle of a large non-operculate specimen.

percula and opercula while defining the genera Serpula,

Crucigera, Hydroides and Protohydroides. However, there

is some inconsistency in terminology since he explicitly

refers to the ‘much reduced ventral most one pair of

branchiae’, and branchiae are also referred to as ‘filaments’

(= radioles). There is, therefore, room for doubt, and one

might infer that his ‘palpi’ are located on the dorsal side,

and may actually be elongated rudimentary opercula.

The condition of the thoracic membranes in P. pacifica is not

clear from the description. They sharply decrease in width

posteriorly from the 5th chaetiger onwards and are insignificant

or lacking as they approach the end of the thorax.

ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961

A range of specimens, including very small ones (Table

25), were covered under the species of Spiraserpula described

in the present paper. The nominal species P. pacifica, there-

fore, has the same number of abdominal segments as the

smallest juvenile among the other three species. The anterior

abdominal chaetae of Serpula and related genera are fre-

quently described as being trumpet-shaped or flat trumpet-

shaped in serpulid literature. Uchida characterizes P. pacifica

as lacking capillaries, but having spatulate chaetae in the

terminal abdominal segments. However, all the species

described in the present paper, including juveniles, possess

capillary chaetae in the terminal abdominal segments,

although there may occasionally be an individual in which

they are lost (see condition in S. singularis sp. nov.).

Table 25 Some abdominal characters of the smallest juveniles of

three new species, compared with those of Protoserpula pacifica.

Species JUG Lengthof No.of Capillaries

(mm) abdomen Abdom. from

(mm) segs.

S. caribensis 367/ 2.0 20 16

S. zibrowii 3.4 ZA 27 19

S. capeverdensis 2.4 2, 29 21

Protoserpula pacifica 5.1 PPS) 20 ~

The bayonet collar chaetae are similar to those of some

species of Spiraserpula, but such chaetae are also found in

Serpula sensu stricto species, such as Serpula japonica Ima-

jima, 1979. It is not known whether Protoserpula possesses

ITS or not, but they had also been overlooked until now in S.

massiliensis (Zibrowius, 1968), S.lineatuba (Straughan,

1967), and S. minuta (Straughan, 1967).

It appears, therefore, that Protoserpula pacifica was a

juvenile serpulid, and its true identity can only be established

with more studies to determine the following: Whether it has

a pair of true palps and an apron; if so, it does not belong to

Spiraserpula. If, however, it has rudimentary opercula and

ITS, and lacks an apron, it is likely to belong to Spiraserpula

Regenhardt, 1961, which has priority over Protoserpula

Uchida, 1978.

PHYLOGENY

Spiraserpula Regenhardt, 1961, is a member of the Serpula/

Crucigera/Hydroides clade. A cladistic analysis of Spiraser-

pula Regenhardt, 1961, based on the above species (Hove &

Pillai) was presented at the Fourth Polychaete Conference,

and the paper is due to be published.

ACKNOWLEDGEMENTS. We wish to express our sincere gratitude to the

following: M. N. Ben-Eliahu (HUJ), P. B. Berents & P. Hutchings

(AM), S. D. Cairns, K. Fauchald & L. Ward (USNM), P. Wagenaar

Hummelinck (former ZLU), M. Jager, (Rohrbach Zement, Dottern-

hausen, Germany), A. Muir (NHM), T. H. Perkins (FMRI), and H.

Zibrowius (SME), for loaning or donating material; M. O. M. Aarts,

R. Fijn, G. van Ee, D. Makhan, C. Schénemann, H. B. Verkaart

jand M. van Vliet (all of former ZLU), G. R. Plaia (FMRI) and R.

van Praag-Sigaar (ITZ) for careful sorting of material, without which

103

we would not have been able to study so many specimens of these

tiny species; the Foundations and Institutions which funded the

second author’s participation in the various expeditions in which the

samples were collected: The Netherlands Marine Science Foundation

(CANCAP Expeditions) and the Indonesian-Dutch Snellius II Expe-

dition; the Netherlands Foundation for the Advancement of Tropical

Research (WOTRO), and Trustees of the Australian Museum,

Sydney, for further field trips. F. Hiemstra for making SEM photo-

graphs; H. Zibrowius (SME) for liberal exchange of data; P. Corne-

lius, A. Muir, and G. Paterson (NHM) for helpful discussions on the

taxonomy; H. Zibrowius (SME), E. W. Knight-Jones and P. Knight-

Jones (UCS) and J. D. George (NHM) for kindly reading through

the manuscript and providing various criticisms and suggestions;

finally, to J.D. George (formerly Head of the Annelida Section), and

C. Curds, Keeper of Zoology, (NHM), for kindly providing facilities

which enabled the first author to undertake studies on this group.

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(A)68: 3-219.

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Netherlands: K/T boundary extinction, survival, and origination patterns.

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Ocean (1976-1986). Part I. List of Stations. Zodlogische Verhandelingen,

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Uchida, H. 1978. Serpulid Tube Worms (Polychaeta.Sedentaria) from Japan

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CONTENTS

1. Anew subfamily and genus in Achatinidae (Pulmonata: Sigmurethra)

A.R. Mead

39 On Recent species of Spiraserpula Regenhardt, 1961, a serpulid polychaete genus hitherto

known only from Cretaceous and Tertiary fossils

T. Gottfried Pillai and H.A. Ten Hove

Bulletin of The Natural History Museum

ZOOLOGY SERIES

Vol. 60, No. 1, June 1994

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VOLUME 60 NUMBER 2 24 NOVEMBER 1994

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Bull. nat. Hist. Mus. Lond. (Zool.) 60(2): 105-172

Phylogenetic relationships between arietellid

genera (Copepoda: Calanoida), with the

establishment of three new genera

S. OHTSUKA

Fisheries Laboratory, Hiroshima University, 1294 Takehara, Hiroshima 725, Japan

G.A. BOXSHALL

Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 SBD, England

H.S.J. ROE

Institute of Oceanographic Sciences Deacon Laboratory, Brook Road, Wormley, Godalming, Surrey

GU8 SUB, England

CONTENTS

EEE NCIC EGET Mineo oe ance eae eee si ces occ an soe cata cheeks SIR eres wanes Matte Pe stoic aciaa.acls dea cinuiels onaeiteisdhcwos seccaciens 105

Matentals and Methods, «.:6.2-cmusteacasnceaesancatiod Meaass-sepr ke a of Cli = 2 ERE St Cee Se 108

AsretelidaeeSars: 1902) os, sash tidwareds cowwtawasetoaell eaheemeene H a Pes RAL a] is. Seca eee A ope tee ee 108

GAASSGRICLAIISIPCNE TOW aidat eee 8-20 c2 es cese oc dnce eee eboed HISTORY SARE Piet ad ON cB sarc clgaaetoieecsssssgescccuces 109

GA TNTERIMOENIE DOVN sass ross tdscus. tacks soe ores mR Uae dete ace auedauce Seta ca ae ntivcnonssicsil Mh eFnscsesecseacchensaeessinasenaee 119

POPAUpApRIOIdES PEM. NOV 2 2r5 6.2... .2sc0sc.seecescoe Porc eesreeces 1.6..DEC..1994 Faciec ccioaipa me cc eeeas eae ab cicaics twits gee 120

Paramisophria T. Scott, 1897 ........:::ccceeeeeeeelh LOG

Metacalanus.Cleve, 1901 gasz..ccssvesecessacesaneacee ee ee ME nae aciste ade isenidervs ance Paseacs geese

BALGULGD MIO OLLETIG OL gal OA er. sian drinnrnaiteisats Reiss alert SaaS eR Meats os niin dsl Joke oft aia ov icisls nase Saar cingimsiede Silesia nmetastecion ste

WAULORCRIN ESE TACIONC: (19 G9RET af <.seteawactas alge Peete eaadtee AEE oxo ssc o0d Bde bblecessnwectad dosoiaeale tment dosinesccdpecseese

APSA ICICIUSMOAMP AMOK, HOSA. sevcaccacsscsctee diet Secteemeccke ebee cana coacBecas hab dis orcas sesedjenstiubebeee Mudasstemeoectees

AE CHAU NV ALCZAMLIOD! ecaacchaite de aka cetians scent Messer seamed ts de te ceue ce nuance cade tembetene nase saa beniecadeaeSeeeReAN Ne ane ee ee

BYES CUSSION Pacers tees eee sec caesar clearest le ne oes atte AM ANIAD Silo te ce aloes eluciie a sinaies wa ie aie eics salar diowsisiaieiae dea teeeetaneltedelteniea ative

FA ORMO WICC PCIICIES aaas. caareatte tanta ance: cecaea re cdaarcena tet occas stiles seanioeweBittuemtdasaeasananeavedseteabeameretvcesctmnares

ERGO MGIC CS mr rane a atte sae aaa nchatandntnare nica valceralo cig en ainciaial ee tcp cecip tvicscles enlan atid le-ae oe inleieisa.walegeiusimnte a sett suaeaeceieraamene

Synopsis. Ten genera, including three new genera, Crassarietellus, Campaneria and Paraugaptiloides, of the family

Arietellidae (Copepoda: Calanoida) are newly defined or redefined with special reference to the genital systems of

the females and fusion patterns and armature elements of appendages. The present study revealed that the single

specimen reported as the male of Sarsarietellus abyssalis (Sars, 1905) represents a new genus, Crassarietellus, and

that Paraugaptilus mohri BjOrnberg, 1975 belongs to the genus Ariefellus. Ancestral and derived states and character

transformations of the genital systems and the appendages in the family are discussed. A cladistic analysis for all 10

genera except for Rhapidophorus revealed that the Arietellidae consists of two lineages, the Crassarietellus-

Paramisophria-Pilarella-Metacalanus-group and the Campaneria-Sarsarietellus-Paraugaptiloides-Scutogerulus-Par-

augaptilus-Arietellus-group. The deep-sea hyperbenthic genera Crassarietellus and Campaneria are the most

plesiomorphic in each group, and the shallow-water hyperbenthic/epipelagic/cave-living Metacalanus and the

bathypelagic Arietellus and Paraugaptilus are relatively apomorphic.

Issued 24 November 1994

INTRODUCTION

_ The family Arietellidae Sars, 1902 has been regarded as one of

the most primitive families in the Calanoida based on the

_ segmentation of appendages and the genital systems (Andronov,

_ 1974; Park, 1986; Huys & Boxshall, 1991). The Arietellidae had

hitherto accommodated the following eight genera: Rhapi-

dophorus Edwards, 1891, Arietellus Giesbrecht, 1892, Parami-

sophria T. Scott, 1897 (= Parapseudocyclops Campaner, 1977),

Metacalanus Cleve, 1901 (= Scottula Sars, 1902), Paraugaptilus

Wolfenden, 1904, Scutogerulus Bradford, 1969, Sarsarietellus

Campaner, 1984, and Pilarella Alvarez, 1985. The genus Phyllo-

pus Brady, 1883 was separated by Brodsky (1950) who proposed

placing it in a new subfamily; it was later removed from the

Arietellidae and placed in its own family, the Phyllopodidae

106

Table 1 Sampling date, locality, depth and gear used for arietellid collection.

S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

Species Sex Number of Date Locality Depth (m) Gear Remarks

specimens

Crassarietellus huysi' ie) 1 18 IV 1977 20°18.5'N, 21°41.2’W 3974-4036 RMT8 5-20 m off

20°20.8'N, 21°53.0'W bottom

fe) 2 18 IV 1977 20°19.7'N, 21°51.3"W 4008-4060 RMT8 5-20 m off

20°18.4’N, 21°40.5'W bottom

Crassarietellus sp.* C 1 24 VI 1908 38°02'N, 10°44’W 04800 Richard net

Campaneria latipes* (of 1 4 X 1968 ST eOHES, WTAE 1234-1260 Modified Menzies Some small

trawl stones

Paraugaptiloides magnus*' Cv 1 11 X 1968 34°38'S, 174°36’E 1697 Modified Menzies

trawl

Arietellus aculeatus Q 1 23 II 1979 15°00'-15°05'N 400 IKMWT

158°00'-158°01'W

of 2 23 II 1979 15°10’-31°14'S 400 IKMWT

71°56'—71°58'W

Arietellus mohri° fe) 1 24 VI 1962 31°10'-31°14'S 1932-3142 Phlenger corer

71°56'-71°58'W

Arietellus pavoninus® fe) 1 28 XI 1965 28°05'N, 14°06'W 600-720 N113H

Arietellus plumifer je) 1 28 VI 1985 31°20'N, 25°17'W 600-840 RMT1

SIP ISHN 25227 W.

Oe 1 26 XI 1965 28°07'N, 14°07'W 680-800 N113H

oy 1 13 XI 1965 28°04'N, 14°12’W 360-410 N113H

Arietellus setosus® C 1 29 XI 1965 28°05'N, 14°10'W 50-85 N113H

Arietellus simplex® Co 1 28 XI 1965 28°05'N, 14°06’W 750-900 N113H

Arietellus sp.° 2 1 11 XI 1965 28°04'N, 14°11'W 460-510 N113H

Metacalanus sp. 1 Q 4 23 V 1989 26°17.9'N, 126°54.2’E 167 Sledge-net

of 1 23 V 1989 26°17.9'N, 126°54.2'E 167 Sledge-net

Metacalanus sp. 2 Q 4 23 V 1989 26°17.9'N, 126°54.2'E 167 Sledge-net

Paramisophria giselae’ Q 2 3 IX 1970 23°19'S, 41°57'W 100 Plankton net adapted

to dredge

Paramisophria japonica® ie) 1 23 V 1989 26°17.9'N, 126°54.2'E 167 Sledge-net

Paramisophria reducta? os 1 25 II 1984 Jameos del Agua 10-28 Plankton net

Paraugaptilus buchani® Q 1 16 XI 1969 17°41'N, 25°18'W 410-500 RMT1

fe) 1 15 XI 1965 27°48'N, 13°55'W 450-510 N113H

Of 1 24 XI 1965 28°06'N, 14°08'W 775-830 N113H

Paraugaptilus similis Q 1 211 1978 04°02'S, 150°00'W 275 IKMWT

or 1 211 1978 04°02'S, 150°00’W 2s IKMWT

Pilarella longicornis'® Q 1 22 VI 1970 28°36'S, 47°55’W 135 Plankton net adapted

to dredge

Scutogerulus pelophilus* Q 1 10 X 1968 34°56'S, 175°23’E 1383-1397 Modified Menzies Globigerina

trawl Ooze

Sarsarietellus abyssalis” 2 1 4-5 VIII 1897 38°37'N, 28°14'W 1260 ‘Nasse’

Sampling data after: ' Boxshall & Roe (1980); 7 Sars (1925); * Bradford (1969); + Bradford (1974); > Bjérnberg (1975); © Currie et al. (1969); ’ Campaner (1977); *

Ohtsuka et al. (1991); ? Ohtsuka et al. (1993a); !” Alvarez (1985).

(Campaner, 1977; Bowman & Abele, 1982).

Arietellids are widely distributed from neritic to oceanic

waters and range vertically from the epipelagic zone to the

bathypelagic hyperbenthic layer (Campaner, 1984).

Recently, cave-living species of Metacalanus and Parami-

sophria have been discovered (Ohtsuka et al., 1993a). How-

ever, neither phylogenetic nor ecological studies on the

family have been carried out in detail, partly because of the

paucity of pelagic arietellids in the water column, and partly

because of the lack of intense sampling effort in the hyper-

benthic layers where many species are found.

Campaner (1984) first examined the relationships between

arietellid genera. He divided them into two morphologically and

ecologically different groups. The first group comprised Arietel-

lus, Paraugaptilus and Scutogerulus, which are characterized by a

reduced female leg 5 and complex male leg 5, and are distributed

in the bathypelagic or deep-sea hyperbenthic zones. The second

group consisted of Metacalanus, Paramisophria, Rhapidophorus

and Sarsarietellus and was diagnosed by characters such as the

relatively well developed leg 5 in the female (except for Metacala-

nus) and the simplified second exopod segments and reduction of

endopod of leg 5 in the male. These are highly adapted

hyperbenthic forms found in relatively shallow waters (<1000 m

deep) or in epipelagic waters in neritic regions. However,

Campaner’s (1984) classification relied solely on the structure of

the fifth legs although he recognized interspecific variation

between congeners in leg characters.

The present paper describes a new arietellid genus col-

lected from the deep-sea hyperbenthic community in the

northeastern Atlantic, and establishes two new genera to

accommodate previously known arietellids, the male of Scu-

togerulus pelophilus Bradford, 1969 and the male of

Paraugaptilus magnus Bradford, 1974. Revised diagnoses of

all known arietellid genera, except for Rhapidophorus, are

given here together with supplementary descriptions. Charac-

ter transformations of the genital systems and appendages of

these arietellids are considered in detail. A cladistic analysis

is employed to help clarify phylogenetic relationships

between the arietellid genera.

PHYLOGENY OF ARIETELLID COPEPODS 107

Table 2. Characters used in the cladistic analysis for genera of the family Arietellidae. Codes 0 to 2 refer to transformation series of

multi-state characters; 0: plesiomorphic state; 9: missing data.

ile Gonopore and copulatory pore sharing common opening yes/no 0/1

2s Right and left copulatory pores separate/fused 01

3: Lengths of right and left antennules of female equal/uneugal 0'1

4. Fusion of female antennulary segments I-III and IV separate/fused O/1

Ds Fusion between female antennulary segments XXIII and XXIV separate/fused 0/1

6. Aesthetasc present on female antennulary segment IV present/absent C/I

ie Aesthetasc present on female antennulary segment VI present/absent C1

8. Aesthetasc present on female antennulary segment VIII present/absent C1

9) Aesthetasc present on female antennulary segment X present/absent 0/1

10. Aesthetasc present on female antennulary segment XII present/absent C1

Ii. Modification of seta into process on male antennulary segment XV no/yes G1

12 Fusion of male antennulary segments XXI & XXII separate/fused 0/1

13. Seta adjacent to aesthetasc on male antennulary segments II present/absent 0/1

14. Seta adjacent to aesthetasc on male antennulary segment III present/absent 0/1

ile Modification of seta into process on male antennulary segment XXII no/yes 01

16. Process on male antennulary compound segment XXIV-XXV no/yes 0/1

17. Seta on first endopod segment of antenna present/absent 0/1

18. Proximal inner seta on mid-margin of second endopod segment of antenna present/absent 0/1

19, Vestigial element on second endopod segment of antenna present/absent C/I

20. l-segmented, rudimentary mandibular endopod with 1 or 2 setae present/absent C'1

PAA Outer terminal seta on fifth exopod segment of mandible normal/reduced O/1

22. Process on maxillulary praecoxal arthrite present/absent C'1

23) Inner basal enditic seta of maxillule present/absent C/1

24. Third seta of maxillulary endopod present/absent 0/1

25: Inner angle seta of maxillulary endopod present/absent 0/1

26. Distal seta on first on first praecoxal endite of maxilla present/absent C1

Zale Reduction of seta a on sixth endopod segment of maxilliped (length of seta

at most as long as the segment) no/yes 0/1

28. Reduction of seta b on sixth endopod segment of maxilliped (length of seta

at most as twice as long as segment) no/yes 0/1

29. Proximal spine on outer margin of third exopod segment of leg 1 present/absent C/1

30. Inner coxal seta of leg 4 present/absent @/1

31. Fusion between endopod and basis of female leg 5 separate/fused G1

B2. Inner margin of endopod of female leg 5 with proximal (seta A) and A+B present/ 0/1

distal (seta B) setae A or B absent

33. One seta (A or B) on inner margin of endopod of female leg 5 present/absent C1

34, Inner angle seta on distal margin of endopod of female leg 5 present/absent C/I

35), Exopod segment of female leg 5 partly defined/ (/1/2

unsegmented/absent

36. Spine (element d) on outer distal angle of exopod of female leg 5 present/absent /1

37. Left endopod of male leg 5 (including incomplete fusion) 2-segmented/ (/1/2

1-segmented/absent

38. Right endopod of male leg 5 l-segmented/absent = (1

59! Seta c on third exopod segment of left male leg 5 present/absent C1

40. Setae e and f of left male leg 5 transformed into bifid process no/yes C1

41. Third exopod segment of left male leg 5 rotated so that vestigial outer

margin elements now on inner surface no/yes 0/1

42. Seta f on third exopod segment of right male leg 5 well developed/minute (/1

43. Seta c on third exopod segment of right male leg 5 present/absent 0/1

44. Fusion between coxa and basis of right male leg 5 separate/fused 0/1

Table 3 Character matrix for analysis using PAUP 3.0.

Earamisopnra lito oroooo0o01 1? 10000 00 OY OO O OW O oil it OO, Oil 0 O&O O o

Metacalanus Dmg Om TiemsiLi (DL mee Lhe () ri (lh Lee Os Oats 1! (OOP Eee se OO eet Leelee 10! OFOMIO

Arietellus Ore a OOM, Ol TO eee ee St OR 1 Rl siO) SOOM ORS Ss On testis

Paraugaptilus Leeda Opae measles eel eile ee LATO alent (0 Tete Te teete Tat OO ee 2 2 SOF We Serer

Scutogerulus OOO OO WOO, OW GeO Te COO! O91 Oe he teh wily tk Oe ve PbO) IO) Os 8) as) 8) &

Sarsarietellus LileO OO, OOOO Y OOo Oi COO OOOO Ot OO LTO O00 0999899 9 Y

Pilarella Oy Oeil Oeil ah OG OP SOO Qe On OO TO OC On ih ale OO CeO. iO it eh it i i Be!) Deore gee &

Campaneria QE ODO DDD 9 DOOM O.0.0 LOO Od OO O Ore og? we Oo Oo Ol to

ULAR ApiMOMesm= omomo! 9 SOS OO sO 1 110) 1 170) (0) OMOMO MONI NONOM 1 Os ONON9, 99 FONO OSORNO! ON 1s se

108

MATERIALS AND METHODS

The present study is based on collections deposited in The

Natural History Museum, London, the New Zealand

Oceanographic Institute, the United States National

Museum, Smithsonian Institution, the Zoological Museum,

University of Oslo, the University of Sao Paulo, and

Hiroshima University. Sampling data and locality are summa-

rized in Table 1. Specimens, except for those previously

mounted, were dissected and mounted in Gum-chloral and

observed with a differential interference contrast microscope

(Olympus BH-2). The genital double-somites of females of

several species were observed with a scanning electron micro-

scope (Hitachi S-800). The morphological terminology is

based on Huys & Boxshall (1991). Type specimens of the new

genera are deposited in The Natural History Museum and the

New Zealand Oceanographic Institute.

Phylogenetic relationships between genera were analyzed

using PAUP version 3.0 prepared by D. Swofford, Illinois

Natural History Survey. The character matrix (Tables 2,3)

summarizes the character distributions among the 10 genera

available for study. A multistate scoring system was

employed and missing characters were scored 9. A hypotheti-

cal composite ancestor was included in the analysis which

scored 0 for all characters. The options employed in the

analysis were Branch and Bound, which guaranteed to find

all the most parsimonious trees, and the MINF optimisation,

which assigns character states so that the f-value is mini-

mized. All characters were set as irreversible using the

Camin-Sokal option.

The abbreviations used in the text and figures 1 to 37 are as

follows: cd: copulatory duct; cp: copulatory pore; g: gonop-

ore; 0: oviduct; rd: receptacle duct; s: spermatophore rem-

nant; sr: seminal receptacle.

SYSTEMATICS

Family Arietellidae Sars, 1902

DIAGNOSIS (emend.) Female. Body of variable size (from ca.

0.8 to 7 mm), relatively robust, rarely compressed.Cephalo-

some and first pedigerous somite separate or weakly fused;

fourth and fifth pedigerous somites completely fused. Cepha-

losome round or pointed at apex; rostrum produced ven-

trally, with pair of filaments. Posterior corner of prosome

sharply or weakly produced, with or without dorsolateral

and/or ventrolateral process. Urosome comparatively short,

4-segmented; genital double-somite with single or paired

gonopores and copulatory pores; gonopore(s) located ventro-

laterally or ventrally, with or without opercular plate; copula-

tory pore sharing common opening with gonopore or

separate from gonopore, located ventro-medially or

-posteriorly, rarely ventrally on right side; seminal recep-

tacles usually paired, rarely left receptacle entirely lacking.

Egg-sac present or absent. Caudal rami well defined, sym-

metrical or slightly asymmetrical, relatively short, with vesti-

gial seta I, well developed or reduced setae II-III, well

developed setae IV—VI and small seta VII.

Antennules symmetrical or asymmetrical, longer on left

side than on right, sometimes differing in fusion pattern and

S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

armature; 16- to 22-segmented; segments I to III, rarely up to

VI fused; segments X to XII more or less fused; segments

XXIII and XXIV separate or fused; segments XXV and

XXVI completely or incompletely fused; segments II, XXII-—

XXIV, XXVI and XXVII lacking aesthetasc; segment IV,

VI, VIII-X, XII and XIII with or without aesthetasc; seg-

ment XIII with 1 or 2 setae; compound segment XXVI-

XXVIII with 8 or 9 elements; posterior margin of proximal

segments fringed with row of setules or not. Antenna: basal

seta present; both rami separate from basis; endopod

2-segmented, first segment with 0-1 inner seta at midlength,

second segment elongate, with 1-3 inner setae medially and 5

or 6 setae terminally; exopod indistinctly 6- to 10-segmented,

ancestral segments I-III and IX unarmed. Mandible: gnatho-

base well chitinized, with 3 or 4 sharp teeth; endopod

rudimentary, 1-segmented with 1 or 2 setae terminally or

completely absent; first exopod segment with normal or

reduced seta, fifth segment carrying 2 setae, one of which

sometimes vestigial. Maxillule: praecoxal arthrite with 0-6

elements; coxal endite with 1 seta or unarmed; coxal epi-

podite carrying 5—9 setae; inner basal seta representing endite

vestigial or absent; endopod bulbous, 1-segmented, with 0-3

setae or completely incorporated to basis; exopod lobate,

bearing 3 long setae. Maxilla well developed; first praecoxal

endite with 1 or 2 setae and 1 vestigial element, second

praecoxal endite having 1 or 2 setae; first and second coxal

endites each with 2 setae; basal spine stout, spinulose or bare;

endopod 4-segmented, with chitinized long setae, setal for-

mula 1,3,2,2. Maxilliped elongate; syncoxa with 1 medial and

2 terminal setae; basis with patches of setules or spinules and

2 setae medially; endopod 6-segmented, first segment almost

fully incorporated into basis, setal formula 1,4,4,3 (rarely 2),3

(rarely 2),4, sixth segment with 2 outermost terminal setae

(setae ‘a’ and ‘b’, see Fig. SC) reduced or not.

Legs 1-4 with distinctly 3-segmented rami or, very rarely,

with endopod segments of leg 1 incompletely fused. Seta and

spine formula of legs 1-4 as shown in Table 4.

Leg 5 variable but not natatory, almost symmetrical; coxae

and intercoxal sclerite separate or fused; basis and endopod

separate or fused; endopod with 0-4 setae; exopod 1- to

3-segmented or completely fused with basis, carrying 0—S

elements.

Male. Body similar to that of female, but urosome

5-segmented. Left antennule geniculate, 16- to 20-segmented;

segments I to IV fused; segments XI to XV more or less

fused; segments XXI and XXII fused or rarely separate;

segments XXIII and XXIV separate; segments XXV and

XXVI completely or incompletely fused; segments II and III

with 1 or 2 setae; segments X, XII-XIV and XX with anterior

process; segments XIX and XXI with 2 processes; segment

XIII with 0-1 seta; segments XV, XXII and XXIV with or

without process; proximal segments often with row of setules

along posterior margins. Mouthparts and legs 1+ similar to

Table 4 Spine and seta formula of legs 1-4.

Coxa Basis Exopod segment Endopod segment

Leg 1 0-1 1-1 I-1;I-1;1V/1,1/1,4 0-1;0-2;1,2,2

Leg 2 0-1 0-0 I-1;I-1;1II,1,5 0-1;0-2;2,2,4/3

Leg 3 0-1 0-0 I-1;1-1;111,1,5 0-1;0-2;2,2,4/3

Leg 4 0-0/1 1-0 I-1;I-1;II1,1,5 0-1;0-2;2,2,3/2

PHYLOGENY OF ARIETELLID COPEPODS

those of female or slightly different in armature elements of

antennary second endopod segment and mandibular first

exopod segment.

Leg 5 variable, but not natatory, almost symmetrical to

strongly asymmetrical; coxae and intercoxal sclerite fused to

form common base or separate; right basis sometimes fused

with coxa; right endopod 1-segmented, bulbous or absent;

right exopod distinctly or indistinctly 3-segmented, first and

second segments each with seta on outer margin (rarely first

segment unarmed), second segment with tuft of setules on

inner distal angle of second segment, third segment with 0-3

elements terminally; left endopod 1- or 2-segmented,

unarmed or completely absent; left exopod distinctly or

indistinctly 3-segmented, first and second segments each with

seta on outer margin, third segment with 1-3 elements

terminally.

TYPE GENUS. Arietellus Giesbrecht, 1892.

REMARKS. The above diagnosis excludes Rhapidophorus

Edwards, 1891, which was inadequately described and has

never been redescribed. Although the family was briefly

defined by Sars (1902), Rose (1933), Brodsky (1950) and

Campaner (1977), the present amended definition includes

new information on the genital systems of females and the

armature elements on the appendages.

Genus Crassarietellus gen. nov.

DIAGNOsIs. Female. Body compact, prosome ovoid in dorsal

view; cephalosome separate from first pedigerous somite;

posterior corner of prosome produced posteriorly to form

rounded lobe. Urosome short, at most one-third as long as

prosome; genital double somite wider than long, with pair of

gonopores ventrolaterally and paired copulatory pores each

located beneath ventral projection; anal operculum not

developed; caudal rami symmetrical, longer than wide, with

vestigial seta I and normally developed seta II.

Antennule symmetrical reaching to posterior end of second

pedigerous somite, 22-segmented; segments I-III fused, with

7 setae and 2 aesthetascs; segments IV, VI, XII and XIII each

with 2 setae and 1 aesthetasc; segments XXIII and XXIV

separate; compound segment XXVI-XXVIII with 8 setae

and 1 aesthetasc; posterior margin of ancestral segments I to

XIII fringed with long setules; segments IV—VIII with trans-

verse row of long setules along distal end of segment.

Antenna: first endopod segment with medial inner seta;

second segment bearing 3 midlength and 5 terminal setae;

exopod indistinctly 10-segmented exopod. Mandibular gna-

thobase with tuft of setules at midlength and 3 teeth on

cutting edge. Mandibular palp: endopod rudimentary,

l-segmented, with 2 setae; seta on first exopod segment not

reduced; outer seta on fifth exopod segment relatively long.

Maxillule: praecoxal arthrite with 5 stout, serrate spines

and 1 process; coxal epipodite having 6 setae; coxal endite

bearing long seta; second basal endite with vestigial seta;

endopod rudimentary, 1-segmented with 2 setae. Maxilla:

first syncoxal endite with 2 setae and vestigial element;

second syncoxal endite with 2 setae; basal endite carrying

stout spine with row of spinules medially. Maxilliped with

second to sixth endopod segments bearing 4, 4, 3, 3 and 4

setae, respectively; innermost seta on fourth and fifth endo-

pod segments not reduced; setae a and b on sixth endopod

segment not reduced.

Leg 1 bearing 2 outer lateral spines on third exopod

109

segment. Leg 5 having distinctly 1-segmented, rudimentary

endopod with 2 setae and indistinctly 3-segmented exopod

with 3 outer lateral and 2 terminal spines.

Male. Left antennule geniculate, 19-segmented; segments

I-IV fused, with 9 setae and 4 aesthetascs; segments XXI and

XXII fused; segments I to X fringed with setules along

posterior margin; segments IV to VIII with transverse row of

setules as in female. Mouthparts and legs similar to those of

female.

Leg 5 with coxae and intercoxal sclerite incompletely fused

to form common base; coxa separate from basis. Right leg

lacking endopod; exopod, at least 2-segmented, first segment

with outer spine on distal corner.Left leg: endopod incom-

pletely 2-segmented, first segment expanded, second segment

small, semispherical; exopod distinctly 3-segmented, first

segment with spine on outer corner, second segment

expanded, bearing outer spine at midlength, third segment

small, having 2 small outer setules and chitinized, long

terminal seta.

TYPE SPECIES. Crassarietellus huysi, gen. et sp. nov.

Other species. Crassarietellus sp. based on a male which

was erroneously assigned to Scottula abyssalis Sars, 1905 by

Sars (1924, 1925).

REMARKS. Sars (1924, 1925) assigned one male collected

from off Lisbon to Scottula abyssalis Sars, 1905, the female of

which was captured off the Azores. However, this male

should be included in the new genus Crassarietellus based on

the similarities of the mouthparts: the indistinctly

10-segmented antennary exopod (compare Fig. 1F with Fig.

7D); 5 serrate spines and a process on the praecoxal arthrite

and 6 setae on the coxal epipodite of the maxillule (Figs 5A,

8A); 2 non-reduced setae on the sixth endopod segment of

the maxilliped (Figs 5C, 8E). Additionally, a transverse row

of setules is present, on each of the antennulary segments IV

to VIII in the male (Fig. 7A), that is found only in the genus

Crassarietellus. The ornamentation of the appendages of the

male, such as the many tiny spinules along the outer margin

of the mandibular palp and the stout, outer processes on the

exopod segments of legs 1 to 4, also supports the proposal to

place the male in Crassarietellus. The right leg 5 of the male

lacks distal exopod segment(s), a condition which Sars (1924,

1925) misinterpreted as ‘l-segmented left’ exopod.

ETYMOLOGY. The new generic name Crassarietellus (Latin

crassus meaning thick) refers to the ovoid, compact body

form of the new genus. The specific name is named in honour

of Mr. Rony Huys.

ECOLOGICAL NOTE. The type species of the new genus was

found in near-bottom samples taken at depths of 3974-4060

m. The plump body and the relatively short antennules

indicates that the new genus is hyperbenthic.

Crassarietellus huysi gen. et sp. nov. (Figs 1-6)

MATERIAL EXAMINED. 3 QQ.

Types. Holotype: 9, 18 IV 1977, North Atlantic (off western

Africa), 20°8.5'N, —21°1.2’W-20°20.8’N, = 21°53.0’W,

3974-4036 m in depth, dissected and mounted on slides,

prosome and urosome preserved in 70% ethanol, BM(NH)

1993. 424. Paratype 1: 9, 18 IV 1977, 20°19.7'N,

21°51.3’N—20°18.4'N, 21°40.5’W, 4008-4060 m in depth, dis-

sected and mounted on slides, prosome preserved in 70%

110 S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

Fig. 1. Crassarietellus huysi gen. et sp. nov., female (holotype: F,G; paratype: A-E). A, Habitus, dorsal view; B, Habitus, lateral view; C,

Urosome, ventral view; D, Genital double-somite, ventral view; E, Genital double-somite, lateral view, cd: copulatory duct; cp: copulatory

pore; g: gonopore; rd: receptacle duct; 0: oviduct; s: spermatophore remnant; sr: seminal receptacle; F, Antenna, one terminal seta on

second endopod segment missing; G, Terminal part of second endopod of other antenna. Scales in mm.

PHYLOGENY OF ARIETELLID COPEPODS 111

Fig. 2. Crassarietellus huysi gen. et sp. nov., female. SEM micrographs of genital double-somite of female. A, Genital double-somite,

ventral view, scale bar = 200 wm (arrows indicating positions of copulatory pores); B, Gonopore and copulatory pore (indicated by arrow),

scale bar = 100 wm; C, Right gonopore, scale bar = 30 wm; D, Left gonopore, scale bar = 30 wm.

S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

Fig. 3 Crassarietellus huysi gen. et sp. nov., female. SEM micro-graphs of remnant of spermatophore attached to genital double-somite of

female. A, Spermatophore remnant penetrating copulatory pore, scale bar = 20 pm; B, Spermatophore remnant, scale bar = 20 pm.

ethanol, BM(NH) 1993. 425. Paratype 2: 9, the same collec-

tion date and locality as in paratype 1, only legs 2 and 3

dissected and mounted on glass slides, urosome mounted on

stub for SEM examination, prosome preserved in 70% etha-

nol, BM(NH) 1993. 426.

BODY LENGTH. 3.88 mm _ (holotype);

(paratypes).

DESCRIPTION. Female. Body (Fig. 1A,B) oval in dorsal

view. Cephalosome and first pedigerous somite separate;

fourth and fifth pedigerous somites completely fused; poste-

rior corner of prosome produced posteriorly into rounded

lobe directed backwards, reaching half length of genital

double-somite. Urosome (Fig. 1C) 4-segmented, one-third as

long as prosome; genital double-somite (Figs 1D,E,2A,B)

wider than long; pair of medial gonopores (Fig. 2C,D)

located ventro-laterally near mid-level of double-somite;

paired copulatory pores posterior to gonopores, each con-

cealed beneath ventrolateral projection; remnants of diver-

gent fertilization tubes of spermatophore (Fig. 3) still

attached to genital double-somite of both paratypes, each

connecting through posteroventral groove with copulatory

pore beneath projection; copulatory duct swollen in ventro-

lateral projection, almost horizontal, extending to large semi-

nal receptacle; 1 medial and 2 pairs of lateral shallow

chitinized pits anteriorly; anal somite small, anal operculum

not developed; caudal ramus (Fig. 1C) longer than wide,

fringed with long setules along inner margin, with vestigial

seta I and developed setae II to VI, seta VII originating

dorsally near base of seta VI; inner margin near anus with

3.88, 3.85 mm

patch of minute spinules. Integument of body and append-

ages pitted.

Antennules (Fig. 4A-C) equal in length, distinctly

22-segmented, reaching to posterior end of second pediger-

ous somite; distal 2 segments incompletely fused; fusion

pattern and armature as follows: I-III-7 + 2 aesthetascs,

IV-—2 + aesthetasc, V-2 + aesthetasc, VI-2 + aesthetasc,

VII-2 + aesthetasc, VIII-2 + aesthetasc, [X—2 + aesthetasc,

X-2 + aesthetasc, XI-2 + aesthetasc, XII-2 + aesthetasc,

XIII-2 + aesthetasc, XIV—2 + aesthetasc, XV-—2 + aes-

thetasc, XVI-2 + aesthetasc, XVII-2 + aesthetasc, XVIII-2

+ aesthetasc, XIX-2 + aesthetasc, XX-2 + aesthetasc,

XXI-2 + aesthetasc, XXII-1, XXIII-1, XXIV—XXVIII-12

+ 2 aesthetascs. Segments I to XIII fringed with long setules

along posterior margin; segments IV to VIII each furnished

with transverse row of minute setules near posterior corner.

Sutures between segments I to III weakly visible.

Antenna (Fig. 1F,G): coxa unarmed; basis with spinulose

seta at inner angle; endopod 2-segmented, first segment with

minute seta at three quarters length, covered with minute

spinules distally, second with 3 setae of unequal lengths

medially and 5 setae distally and sparsely covered by spinules;

exopod indistinctly 10-segmented, second to fourth segments

almost fused; armature as follows: 0,0,0,1,1,1,1,1,0,3; ninth

segment sparsely ornamented with minute spinules.

Mandible (Fig. 4D): gnathobase heavily chitinized, ventro-

medial margin with dense fringe of long setules; cutting edge

with 3 acute teeth, dorsalmost of which bifid at tip; 2 patches

of dagger-like spinules present dorsally; tuft of long setules

present medially on knob; basis of palp with patches of

PHYLOGENY OF ARIETELLID COPEPODS 113

Fig. 4. Crassarietellus huysi gen. et sp. nov., female (holotype: E,F; paratype: A-D). A, Antennulary segments I to XV; B, Antennulary

segments XVI to XXVIII; C, Antennulary segments XXII to XXVIII; D, Mandible; E, Maxillulary praecoxal arthrite and coxal endite; F,

Proximal spine on praecoxal arthrite of maxillule. Scales in mm.

114 S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

Fig. 5. Crassarietellus huysi gen. et sp. nov., female (holotype: C; paratype: A,B). A, Maxillule, with arrowhead indicating enditic seta of

basis; B, Maxilla; C, Maxilliped. The armature elements on the sixth endopod segment of maxilliped are identified individually by the

letters a to d. Scales in mm.

PHYLOGENY OF ARIETELLID COPEPODS tS

Fig. 6. Crassarietellus huysi gen. et sp. nov., female (holotype: A-C,J-L; paratypes: D-I). A, Second endopod segment of maxilliped; B,

Third endopod segment of maxilliped, innermost seta indicated by arrowhead; C, Fourth endopod segment of maxilliped, innermost seta

indicated by arrowhead; D, Leg 1, anterior surface; E, Leg 2, posterior surface; F, Aberrant leg 3, anterior surface; G, Right endopod of

leg 3, anterior surface; H, Another aberrant leg 3, posterior surface; I, Extremely aberrant leg 3, anterior surface; J, Leg 4, posterior

surface; K, Left leg 5, anterior surface; L, Right leg 5, anterior surface. Scales in mm.

116

minute spinules and row of long setules proximally (almost

missing in Fig. 4D); endopod rudimentary, 1-segmented, with

2 plumose setae of unequal lengths; exopod 5-segmented,

almost completely separate, first to fourth segments each

bearing 1 seta, terminal segment with 2 setae, one of which

thinner and shorter than other; second segment with patch of

minute spinules.

Maxillule (Figs 4E,F,5A): praecoxal arthrite with 5 stout

spines, 2 of which (Fig. 4F) bearing 2 rows of strong spinules,

and 1 process, patch of long setules, and numerous minute

spinules of various sizes along inner margin and patch of fine

prominences along outer margin; coxal epipodite with 6

setae; coxal endite with elongate, spinulose seta terminally;

basis carrying minute enditic seta and row of long, fine setules

along inner margin; endopod rudimentary, 1-segmented,

bearing 2 spinulose setae of unequal lengths distally; exopod

lamellar, having 3 long, plumose setae distally.

Maxilla (Fig. 5B) stout; first praecoxal endite with 2

spinulose setae and vestigial element; second praecoxal and

both coxal endites each carrying 2 spinulose setae; basal

endite bearing long, subterminal spine with 2 rows of spinules

medially; endopod 4-segmented, first segment with 1 spinu-

lose seta, second to fourth segments having 3, 2 and 2 long,

spinulose setae, respectively.

Maxilliped (Figs SC,6A-C) elongate; syncoxa with 1 medial

and 2 subterminal setae and patch of fine spinules subtermi-

nally; basis bearing 2 patches of spinules proximally and

midway along inner margin and 2 spinulose subterminal

setae; endopod 6-segmented, first segment incompletely

fused with basis, first to sixth segments carrying 1, 4, 4, 3, 3,

and 4 setae, respectively; innermost seta on fourth and fifth

segments relatively long; sixth with setae a and b well

developed, seta c chitinized, bearing row of simple spinules

along inner margin, seta d long, with inner row of simple

spinules.

Leg 1 (Fig. 6D); second endopod segment produced at

outer angle; third endopod segment produced distally into

acute process, with 2 outer lateral spines and terminal plu-

mose seta; first exopod segment produced near outer angle;

second and third exopod segments produced at outer angle.

Leg 2 (Fig. 6E) and leg 3 (Fig. 6F-I) similar; outer angle of

second endopod segment acutely produced; third endopod

segment with 4 inner setae. Third legs with several aberra-

tions: extra spine present on each of first (Fig. 6F) and third

exopod segments (Fig. 6F,H); extra seta on first (Fig. 6H)

and second endopod segments (Fig. 6F); fewer seta on third

endopod segment (Fig. 6F); both rami extremely abnormal

(Fig. 61). Leg 4 (Fig. 6J): basis with small plumose seta near

base of exopod on posterior surface; terminal endopod

segment with 3 inner setae.

Leg 5 (Fig. 6K,L): both legs almost symmetrical; right and

left coxae incompletely separate from intercoxal sclerite;

basis with relatively narrow base, bearing plumose seta at

outer angle; endopod small, 1-segmented, distinctly separate

from basis, with inner medial and terminal plumose seta;

exopod indistinctly 3-segmented, each almost fused, first and

second segment with serrate spine at outer angle, third with 2

terminal and 1 lateral spines.

Male. Unknown.

VARIABILITY. The paratypic females have aberrant third legs

(Fig. 6F,H,I). Both paratypes have 4 outer spines on the third

exopodal segment of leg 3, but it is likely that the segment

normally has 3 outer spines, because the males of Crassari-

S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

etellus sp. and other arietellids carry only 3 spines on this

segment. An additional spine on the first exopodal segment

of leg 3 has also been reported in specimens of some

shallow-water hyperbenthic and cave-dwelling species of the

calanoid family Pseudocyclopiidae (Scott, 1894; Fosshagen &

Iliffe, 1985). Some females of Paracyclopia naessi Fosshagen,

1985 had 2 outer spines on the first exopodal segment of leg 3

(Fosshagen & Iliffe, 1985) and this segment of the same leg in

Pseudocyclopia crassicornis Scott, 1892 was figured with 2

spines (Scott, 1892).

It is interesting to note that it is the same segment of the

same leg which carried the extra spine in both Crassarietellus

and pseudocyclopiids. The presence of a seta on the outer

margin of the second endopodal segment of leg 3 (Fig. 6F) is

unique for the Calanoida. Elsewhere in the Copepoda such a

seta has only ever been found in the two superornatiremid

harpacticoids figured by Huys & Boxshall (1991).

REMARKS. The male of C. huysi is unknown. Crassarietellus

sp. described below, which was erroneously considered to be

the male of Sarsarietellus (= Scottula) abyssalis (Sars, 1905),

is similar to C. huysi except in sexual dimorphic characters,

but is smaller than C. huysi. Considering that the locality of

Crassarietellus sp. (38°02'N, 10°44’W) is near the type locality

of C. huysi (20°18.5'N, 21°41.2’W-20°20.8'N, 21°53.0’W), it

is possible that this male can be assigned to C. huysi.

Crassarietellus sp. (Figs 7-8)

of Oslo, Catalog No. F5445-5446, labeled as Scottula abyssa-

lis G.O. Sars.

BODY LENGTH. 2.8 mm (after Rose, 1933).

DESCRIPTION. Integument of urosome and appendages pit-

ted as in Crassarietellus huysi. Left antennule (Fig. 7A-C)

geniculate between ancestral segments XX and XXI, fringed

with setules along posterior margins of segments I-X, trans-

verse row of setules on each of segments IV to VIII; fusion

pattern and armature as follows: I-IV—9 + 4 aesthetascs, V—2

+ aesthetasc, VI-2 + aesthetasc, VII-2 + aesthetasc, VIII-2

+ aesthetasc, [IX-2 + aesthetasc, X-2 + aesthetasc, XI-2 +

aesthetasc, XIJ—2 + aesthetasc, XIII—2 + aesthetasc, XIV—2

+ aesthetasc, XV-2 + aesthetasc, XVI-2 + aesthetasc,

XVIU-2 + aesthetasc, XVIII-2 + aesthetasc, XIX-1 +

aesthetasc + 2 processes, XX-2 + process, XXI-—XXIII-1 +

aesthetasc + 2 processes, XXIV—XXVIII-12 + 2 aesthetascs;

segment XXV incompletely fused with XXVI. Sutures

between segment I to IV weakly visible. Right antennule as in

female of Crassarietellus huysi.

Antenna (Fig. 7D): basis with serrate inner seta; endopod

2-segmented, first segment with short, inner seta at three

quarters length and numerous spinules subterminally, second

segment with 3 inner setae of unequal lengths and 5 setae

terminally, covered almost entirely with spinules; exopod

indistinctly 10-segmented, eighth segment fringed with

minute spinules along both sides; setal formula of exopod as

follows: 0,0,0,1,1,1,1,1,0,3.

Mandibular gnathobase (Fig. 7F) with 3 stout teeth, dor-

salmost of which bifid at tip; tuft of long setules present near

base of palp. Mandibular palp (Fig. 7E): basis elongate,

furnished with numerous minute spinules and row of long

setules along inner margin; endopod _ rudimentary,

1-segmented, bearing 2 unequal setae; seta on first exopod

PHYLOGENY OF ARIETELLID COPEPODS

Fig. 7. Crassarietellus sp., male. A, Left antennule; B, Antennulary segments XIX to XXVIII, elements on segments XXIV-XXVIII

omitted except for outer seta; C, Antennulary segments XXIV—XXVIII; D, Antenna; E, Mandibular palp; F, Mandibular gnathobasic

cutting edge. Scales in mm.

117

118 S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

Fig. 8. Crassarietellus sp., male. A. Praecoxal arthrite and coxal endite of maxillule; B, Maxillulary endopod; C, First and second praecoxal

endites of maxilla; D, Basal spine of maxilla; E, Fourth to sixth endopod segments of maxilliped, inner seta on fourth and fifth segments

indicated by arrowhead; F, Leg 1, anterior surface; G, Leg 2, anterior surface; H, Outer distal process on second exopod segment of leg 3;

I, Outer process on second endopod segment of leg 3; J, Leg 5, anterior surface, ancestral second and third segments of right exopod

missing. Scales in mm.

PHYLOGENY OF ARIETELLID COPEPODS

segment not reduced, fifth segment with 2 developed setae.

Maxillule (Fig. 8A,B): praecoxal arthrite carrying 5 serrate

spines and 1| process, with numerous spinules of variable sizes

on both sides and patch of setules; coxal endite with long,

serrate seta; coxal epipodite bearing 6 setae.

Maxilla: first and second syncoxal endites (Fig. 8C) having

2 setae and vestigial element, and 2 spinulose setae, respec-

tively; basal spine (Fig. 8D) with 3 rows of spinules at

midlength.

Maxilliped (Fig. 8E): fourth and fifth endopod segments

each with non-reduced innermost seta, sixth segment with

setae a and b well developed.

Leg 1 (Fig. 8F): coxa with plumose seta at inner angle and

tuft of long setules near outer proximal margin; basis with

outer and inner plumose seta; endopod 3-segmented, all

segments with outer distal angle produced distally; exopod

3-segmented, first segment with outer setiform spine reaching

to distal end of second, third segment with 2 outer lateral

spines and 1 spiniform terminal seta. Legs 2 (Fig. 8G) and 3

with the same segmentation and setation; basal inner corner

rounded; outer process on second endopod segment (Fig. 81)

with minute spinules along inner margin; terminal outer

process on first and second exopod segments (Fig. 8H) also

carrying small projections midway along inner margin. Leg 4:

coxa unarmed; basis with outer seta on posterior surface;

endopod 3-segmented, setal formula 0—1;0—2;2,2,3; exopod

distal 2 segments missing, first segment with outer spine and

inner seta.

Leg 5 (Fig. 8J): coxae incompletely fused with intercoxal

sclerite; basis separate from coxa, bearing outer plumose seta

at midlength. Right leg lacking endopod; exopod missing

distal segment(s), at least, 2-segmented, first segment with

spinulose spine and pointed process at distal angle. Left leg

with indistinctly 2-segmented endopod, first segment large,

second hemispherical with minute prominence terminally;

exopod 3-segmented, first segment with spinulose spine and

pointed process on distal corner, second segment expanded,

carrying outer spinulose spine at midlength, third segment

small, tapering distally, with 1 minute basal element, 2 short

medial setae along outer margin and terminal spine as long as

second segment.

REMARKS. Since the third leg of Crassarietellus sp. has 3

outer spines on the third exopodal segment and 1 inner seta

on the first exopodal segment, as most other arietellids, the

third legs of the paratypes of C. huysi are here interpreted as

abnormal.

Genus Campaneria gen. nov.

DIAGNOsIs. Only male known. Cephalosome and first pedi-

gerous somite separate. Anal somite almost telescoped into

preceding somite; anal operculum not developed. Caudal

rami symmetrical, longer than wide, with vestigial seta I,

well-developed setae II-VI and minute seta VII.

Left antennule reaching almost to end of urosome, genicu-

late, 20-segmented; segments II to IV almost fused but

sutures clearly visible, segments II and III each bearing seta

and aesthetasc; segment XIII with seta, aesthetasc and pro-

| cess representing modified seta; segment XXI separate from

XXII; segment XXV incompletely fused with XXVI; seg-

ment XIII with seta and process; compound segment

XXVI-XXVIII with 8 setae and aesthetasc; segment II

119

(probably, originally from I) to XIII fringed with setules

posteriorly.

Antenna: first endopod segment having inner seta, second

segment bearing 3 inner setae subterminally and 5 setae

terminally; exopod indistinctly 8-segmented. Mandibular

gnathobase with tuft of setules. Mandibular palp: endopod

rudimentary, 1-segmented, with 2 setae; seta on first exopod

segment not reduced; outer seta on fifth exopod segment

relatively long.

Maxillule: praecoxal arthrite carrying 5 spines, 3 of which

weakly serrate medially, and process; coxal endite with long

seta; coxal epipodite with 6 setae; second basal endite repre-

sented by vestigial seta; endopod bulbous, 1-segmented,

having 2 setae.

Maxilla: first syncoxal endite with 2 setae and vestigial

element; second syncoxal endite with 2 setae; basal endite

bearing stout spine with 3 rows of spinules proximally.

Maxilliped: setal formula of endopod 1,4,4,3,3,4; fourth

endopod segment with non-reduced innermost seta, fifth

segment with shorter innermost seta than fourth, sixth seg-

ment with seta a vestigial and seta b relatively long.

Leg 1 with 2 outer spines on third exopod segment. Leg 4

lacking inner coxal seta. Leg 5 with coxae and intercoxal

sclerite fused to form a common plate; coxa separate from

basis. Right leg: endopod l-segmented, bulbous; exopod

indistinctly 3-segmented, distal 2 segments almost fused,

expanded medially, with rounded process medially and 2

setules and 1 prominence terminally. Left leg: endopod

indistinctly 2-segmented, unarmed; exopod 2-segmented, dis-

tal segment curved outwards near tip, with 3 setae terminally

and 1 seta medially.

TYPE SPECIES. Campaneria latipes gen. et sp. nov.

REMARKS. As already suggested by Bradford (1969), we

conclude that the single paratypic male of Scutogerulus

pelophilus belongs to a different species from the female.

Although sexual dimorphism in mouthparts is exhibited in

arietellids such as Arietellus (present study) and Paraugapti-

lus (Deevey, 1973; present study), the sexual differences are

restricted to the antennary rami and the first mandibular

exopod segment. However, the male differs from the holo-

type female of S. pelophilus in armature elements on the

mouthparts and leg 1 as follows: (1) the female has ‘shield-

shaped’ appendages (= ornamentation) (Bradford, 1969) on

terminal setae of the maxilla and maxilliped, while the male

lacks such ornamentation; (2) there is single inner seta on the

first antennary endopod segment in the male but none in the

female; (3) the praecoxal arthrite of maxillule has 6 elements

in the male (5 spines and 1 process) and 5 in female (4 spines

and 1 process); (4) the maxillulary endopod has 2 setae in the

male and 1 in the female; (5) the first and second praecoxal

endites of the maxilla bear 2 setae plus a vestigial element and

2 setae in the male, and 1 seta plus a vestigial element and 1

seta in the female, respectively; (6) seta b on the sixth

endopod segment of maxilliped is long in the male but short

in the female; (7) the third exopod segment of leg 1 has 2

outer spines in the male but only 1 in the female. As far as the

armature is concerned, the female shows more apomorphic

character states than the male. In particular, the magnitude

of the differences in the antenna, maxilla, maxilliped and leg

1 is greater than not only variation within a species but also

normal interspecific discrepancies between congeners. A new

genus is, therefore, established to accommodate the male.

120

The male of the new genus is similar to that of Crassarietel-

lus. However, the left antennule, the antennary exopod, the

maxillulary praecoxal arthrite, and the fifth and sixth endo-

pod segments of maxilliped are different: (1) left antennule

reaching almost to end of urosome in Campaneria, but,

possibly, at most to end of prosome in Crassarietellus; (2)

antennulary segments II to IV partly fused in Campaneria,

but almost completely so in Crassarietellus; (3) antennulary

segments II and III each bearing single seta and aesthetasc in

Campaneria, but 2 setae and aesthetasc in Crassarietellus; (4)

antennulary segments XXI and XXII completely separate in

Campaneria, but almost fully fused in Crassarietellus; (5) seta

on antennulary segment XV modified into process in Cras-

sarietellus, but not in Campaneria; (6) antennary exopod

indistinctly 8-segmented in Campaneria but 10-segmented in

Crassarietellus; (7) spines on maxillulary praecoxal arthrite

finely serrate in Campaneria, but strongly serrate in Crassari-

etellus; (8) innermost seta on the fifth endopod segment of

maxilliped relatively short in Campaneria, but long in Cras-

sarietellus; (9) seta a on the sixth endopod segment of

maxilliped relatively reduced in Campaneria, but not in

Crassarietellus.

The leg 5 of Campaneria is also similar to that of Crassari-

etellus sp., particularly in having a 2-segmented left endopod,

but can be distinguished by the presence of the right endopod

and by the 2-segmented left exopod.

ETYMOLOGY. The new genus Campaneria is named in

honour of the late Dr. A. Campaner who was the first to be

interested in the phylogenetic relationships between arietellid

genera (gender feminine). The specific name J/atipes (Latin

latus meaning broad; Latin pes meaning leg) refers to the

broad compound exopod segments of the right leg 5 of the

male.

ECOLOGICAL NOTE. Campaneria was collected by a trawl

from the near-bottom samples taken at depths of 1234-1260

m off northeastern New Zealand (Bradford, 1969). Since the

genus has never been captured in plankton hauls, it is most

likely hyperbenthic.

Campaneria latipes gen. et sp. nov. (Figs 9-10)

MATERIAL EXAMINED. O', New Zealand Oceanographic

Institute Reg. No. 121, labelled as Scutogerulus pelophilus

(C).

BODY LENGTH. 3.9 mm (after Bradford, 1969).

DESCRIPTION. Anal somite (Fig. 9A) small, almost tele-

scoped into preceding somite; caudal rami (Fig. 9A) sym-

metrical, seta I vestigial, setae II-VI developed, seta VII

minute.

Left antennule (Fig. 9B-F): segment I damaged, but with 3

setae and aesthetasc (only this segment still remained on the

body); segments II and III fused with suture visible ant-

eriorly; segments III and IV, and XXIV-XXV_ and

XXVI-XXVIII incompletely fused. Fusion pattern and arma-

ture elements as follows: I-IV-4 + 3 aesthetascs, V—2 +

aesthetasc, VI-2 + aesthetasc, VII-2 + aesthetasc, VIII-2 +

aesthetasc, IX-2 + aesthetasc, X-1 + aesthetasc + process,

XI-2 + aesthetasc, XII-1 + aesthetasc + process, XIII-1 +

aesthetasc + process, XIV—1 + aesthetasc + process, XV—2

+ aesthetasc, XVI-2 + aesthetasc, XVII-2 + aesthetasc,

XVIII-2 + aesthetasc, XIX-1 + aesthetasc + 2 processes,

XX-1 + aesthetasc + process, XXI-aesthetasc + 2 processes,

S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

XXII-XXII-1 + process (XXlII-process, XXIII-1),

XXIV-XXVIII-12 + 2 aesthetascs.

Antenna: inner basal seta present; endopod (Fig. 9G)

2-segmented, first segment with short inner seta, second

segment with 3 inner setae of unequal lengths subterminally

and 5 setae terminally; exopod (Fig. 10A) indistinctly

8-segmented, second segment elongate, setal formula

0;1,1,1,151,0;3.

Mandibular palp (Fig. 10E): endopod rudimentary,

1-segmented, carrying 2 setae of unequal lengths; first exo-

pod segment bearing non-reduced seta, fifth segment with 1

long and 1 shorter seta.

Maxillule: praecoxal arthrite (Fig. 10B) bearing 5 spines

and 1 process, 3 of which serrate medially, with row of long

setules and patch of minute spinules proximally; coxal endite

(Fig. 10C) with long spinulose seta terminally; coxal epi-

podite with 6 setae; minute endite seta present on basis (Fig.

10D), endopod bulbous, 1-segmented, with 2 spinulose setae

of unequal lengths.

Maxilla: first praecoxal endite with 2 spinulose setae and

vestigial element, second endite with 2 bipinnate setae (Fig.

10F); basal spine (Fig. 10G) with 3 rows of spinules of

different sizes proximally.

Maxilliped: fourth and fifth endopod segments (Fig. 10H)

each having non-reduced, spinulose innermost seta, but seta

on fourth segment much longer than on fifth; sixth endopod

segment (Fig. 101) with medium-length seta b and vestigial

seta a.

Leg 1 with 2 outer spines on third exopod segment. Leg 4

having outer basal seta, but lacking inner coxal seta.

Leg 5 (Fig. 10J): coxae and intercoxal sclerite almost fused,

but suture visible on posterior surface; basis separate from

coxa. Right leg: basal seta missing; endopod 1-segmented,

with tuft of short setules terminally; exopod indistinctly

3-segmented, first triangular, carrying spine at outer angle,

distal 2 segments almost fused, but suture visible on both

surfaces, expanded medially, having outer seta proximally,

round inner process with 3 minute prominences at tip medi-

ally, and 2 setae and 1 prominence along outer terminal

margin. Left leg: basal seta missing; endopod indistinctly

2-segmented, unarmed; exopod 2-segmented, first segment

triangular, bearing spine on outer corner, second segment

expanded inwards, curved outwards at about three quarters

length, with fine medial seta and 3 terminal setae of unequal

lengths.

REMARKS. In her original description Bradford (1969) over-

looked the antennary basal seta, the inner seta on the first

antennary endopod segment, 3 short setae on the distal 2

endopod segments of the maxilliped, the outer basal seta of

leg 4, and the fine midlength seta on the second exopod

segment of left leg 5.

Genus Paraugaptiloides gen. nov.

DIAGNOSIS. Only male known. Body similar to that of

Paraugaptilus; cephalosome separate from first pedigerous

somite; prosome rounded anteriorly and produced posteri-

orly, with small dorsolateral prominence and bluntly pro-

duced lateral lobe on each side; lateral flap of cephalosome

developed to cover bases of mouthparts. Caudal rami sym-

metrical with setae II and III normally developed.

Male left antennule 19-segmented, fringed with setules

along posterior margin of first segment only; segments I and

PHYLOGENY OF ARIETELLID COPEPODS 121

Fig. 9. Campaneria latipes gen. et sp. nov., male (holotype). A, Anal somite and caudal rami, dorsal view; B, Left antennulary segments II

to XV; C, Left antennulary segments XVI to XIX; D, Left antennulary segments XX to XXVIII; E, Anterior processes on segments

XX-_XXIV of left antennule; F, Left antennulary segments XXIV to XXVIII; G, Antennary endopod. Scales in mm.

122 S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

Fig. 10. Campaneria latipes gen. et sp. nov., male (holotype). A. Antennary exopod; B, Praecoxal arthrite of maxillule; C, Coxal endite of

maxillule; D, Maxillulary endopod with basal seta indicated by arrowhead; E, Mandibular endopod and exopod; F, First and second

praecoxal endite of maxilla; G, Basal spine of maxilla; H, Fourth and fifth endopod segments of maxilliped, innermost seta indicated by

arrowhead; I, Sixth endopod segment of maxilliped; J. Leg 5, anterior surface. Scales in mm.

PHYLOGENY OF ARIETELLID COPEPODS

II each with 1 seta; segment XIII with seta and process;

segment XXI fused with XXII; compound segment

XXIV-XXV with large cuticular process; compound segment

XXVI-XXVIII with 8 setae and aesthetasc. Antenna: first

endopod segment without inner seta, second segment with 2

inner setae at midlength and 5 setae and 1 setule terminally;

exopod indistinctly 8-segmented, _setal formula

0,1,1,1,1,1,0,3. Mandibular palp: endopod rudimentary,

1-segmented, with 2 setae; seta on first exopod segment not

reduced; outer seta on fifth exopod segment relatively long.

Maxillule: praecoxal arthrite with 5 spines and 1 process;

coxal endite carrying long seta; coxal epipodite bearing 8

setae; no basal seta; endopod 1-segmented, bearing 2 setae.

Maxilla: first praecoxal endite with 2 setae and 1 vestigial

element; second praecoxal endite with 2 setae; basal spine

with 2 rows of spinules. Maxilliped: endopodal setal formula

1,4,4,3,3,4; innermost seta on fourth and fifth endopod

segments not vestigial; seta a on sixth endopod segment

reduced, seta b relatively long.

Leg 1 with 2 outer spines on third exopod segment. Leg 4

with vestigial element on inner distal corner of coxa. Leg 5:

coxae fused with intercoxal sclerite; basis and coxa separate

in left leg and incompletely fused in right. Right leg: endopod

l-segmented, rudimentary, unarmed; second exopod seg-

ment expanded inwards, almost completely separate from

third segment, third segment triangular, tapering distally,

with 1 minute outer and 1 terminal setules. Left leg: endopod

2-segmented, unarmed; exopod 3-segmented, distal 2 seg-

ments completely separate, second segment expanded

inwards, third segment with 2 long stout processes directed

laterally.

TYPE SPECIES. Paraugaptilus magnus Bradford, 1974 (mono-

typic).

REMARKS. Bradford (1974) assigned a male collected from a

depth of 1697 m off the north-east coast of North Island, New

Zealand, to the genus Paraugaptilus, although she mentioned

seven distinct characters of the species that would possibly

necessitate its removal to a new genus. Morphological discon-

tinuities can be found between P. magnus and other species

of Paraugaptilus as follows: (1) left antennulary compound

segment XXVI-XXVIII with 8 setae and aesthetasc; (2)

antennary exopod indistinctly 8-segmented, with setal for-

mula 0,1,1,1,1,1,0,3; (3) mandibular endopod almost fused

with basis, but represented by a rudimentary segment with 2

setae; (4) maxillule with long seta on coxal endite, 1 basal

seta and 2 setae on 1-segmented endopod; (5) maxilla with 2

setae and 1 vestigial element on first praecoxal endite and 2

setae on second; (6) setae on maxillary endopod ornamented

with row of simple spinules along inner margin but lacking

triangular-shaped ornamentation found in other species of

Paraugaptilus; (7) seta b on sixth endopod segment of maxil-

liped not reduced; (8) second and third exopod segments of

right leg 5 almost completely separate; (9) leg 5 with

2-segmented left endopod.

In genera accommodating several species, such as Parami-

sophria, Arietellus and Metacalanus, the praecoxal arthrite,

coxal endite and endopod of maxillule, first praecoxal endite

of maxilla, and leg 5 exhibit wide interspecific variation in

armature. However, the armature of the antennary exopod,

mandibular palp, second praecoxal endite of maxilla, endo-

pods of male leg 5 are relatively consistent within each genus.

In particular, the significant differences found in the anten-

123

nary exopod, the mandibular endopod and the second prae-

coxal endite of the maxilla support the proposal to assign P.

magnus to a new genus, Paraugaptiloides.

The new genus is similar to Arietellus and Paramisophria in

the segmentation and setation of appendages, but can be

distinguished from these genera by: (1) the presence of a

large cuticular process on left antennulary segments

XXIV-XXV (shared with Paraugaptilus); (2) the lack of a

seta on the first endopod segment of antenna, also absent in

Arietellus but present in Paramisophria; (3) the 2 inner

medial setae on the second endopodal segment of antenna in

Paraugaptiloides and Arietellus, compared to 3 in Parami-

sophria; (4) outer seta on fifth exopodal segment of mandible

relatively long in Paraugaptiloides and Paramisophria, but

vestigial in Arietellus; (5) mandibular endopod 1-segmented

with 2 setae in Paraugaptiloides and Paramisophria, but

absent in Arietellus; (6) maxillule with 1 basal and 2 endopo-

dal setae in Paraugaptiloides and Paramisophria, but no basal

and, at most, single endopodal seta in Arietellus; (7) maxil-

lary basal spine ornamented with spinules in Paraugaptiloides

and Arietellus, but no ornamentation in Paramisophria; (8)

innermost seta on fourth and fifth endopodal segments of

maxilliped vestigial in Arietellus, but not in Paraugaptiloides

and Paramisophria; (9) seta a on the sixth endopodal segment

of maxilliped reduced only in Paraugaptiloides and Arietellus;

(10) the presence of vestigial element on inner distal angle of

coxa of leg 4 (shared with Paraugaptilus); (11) left leg 5

endopod 2-segmented in Paraugaptiloides and Arietellus, but

1-segmented in Paramisophria; (12) right endopod of leg 5

present in Paraugaptiloides and Arietellus, but absent in

Paramisophria.

ETYMOLOGY. The name refers to the close relationship of

the new genus to Paraugaptilus.

ECOLOGICAL NOTE. The male of P. magnus was first col-

lected from 1697 m depth off New Zealand (Bradford, 1974),

and has been reported recently from the near-bottom

(1060-1070 m depths) in the southwestern Indian Ocean

(Heinrich, 1993). It is likely that P. magnus is widely distrib-

uted in deep waters of the Indo-Pacific region. Although the

species was collected from the near-bottom in the Indian

Ocean (Heinrich, 1993), the well-developed antennules sug-

gest a relatively loose association with the bottom (Cam-

paner, 1984).

Paraugaptiloides magnus, new combination (Figs

11-12)

MATERIAL EXAMINED. OC’, holotype, New Zealand Oceano-

graphic Institute H-199.

BODY LENGTH. 4.85 mm (after Bradford, 1974).

DESCRIPTION. Cephalosome separate from first pedigerous

somite. Caudal ramus with setae II-VI well developed.

Left antennule (Fig. 11A,B) 19-segmented, the fusion

pattern and armature elements almost same as in Paraugapti-

lus, except for those of segments XXIV to XXVIII: segment

XXIV-XXV with large anterior process reaching well beyond

antennulary tip (Fig. 11B). Right antennule: segments I to X

fringed with long setules along posterior margin; segments X

and XI, and XIV and XV only partly fused; segment XXIII

and XXIV almost separate; segments XXV and XXVI almost

fused with suture visible; fusion pattern and armature ele-

ments as follows: I-III-7 + 3 aesthetascs, IV—2 (element

124 S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

Fig. 11. Paraugaptiloides magnus gen. et sp. nov., male (holotype). A. Left antennulary segments XIX to XXVIII; B, Left antennulary

segments XXVI to XXVIII; C, Antenna; D, Mandibular endopod and exopod; E, First and second praecoxal endites of maxilla; F, Basal

spine of maxilla; G, Terminal seta on fourth endopod segment of maxilla. Scales in mm.

PHYLOGENY OF ARIETELLID COPEPODS

125

Fig. 12. Paraugaptiloides magnus gen. et sp. nov., male (holotype). A, Praecoxal arthrite, coxal endite, basal endite and endopod of

maxillule, basal seta indicated by arrowhead; B, Fourth to sixth endopod segments of maxilliped, innermost seta on fourth and fifth

segments indicated by arrowhead; C, Inner coxal seta of leg 4; D, Leg 5, posterior surface, scar of element on third exopod segment of left

leg indicated by arrowhead; E, Right endopod of leg 5; F, Left endopod of leg 5; G, Inner distal process on second exopod segment of right

leg 5. Scales in mm.

missing), V—1 + aesthetasc (element missing), VI-2 + aes-

thetasc, VII-2 + aesthetasc, VIII-2 + aesthetasc, IX-2 +

aesthetasc, X-1 + aesthetasc + process, XI-2 + aesthetasc,

XII-2 + aesthetasc, XIII-2 + aesthetasc, XIV-1 + aes-

thetasc + process, XV—2 + aesthetasc, XVI-2 + aesthetasc,

XVII-2 + aesthetasc, XVIII-2 + aesthetasc, XIX-2 +

aesthetasc, XX-1 + aesthetasc (element missing), XXI-2 +

aesthetasc, XXII-1, XXIII-1, XXIV-XXVIII-12 + 2 aes-

thetascs (XXIV-2, XXV-2 + aesthetasc, XXVI-XXVIII-8

+ aesthetasc).

Antenna (Fig. 11C): first endopod segment lacking inner

seta, second segment with 2 inner setae of unequal lengths

subterminally and 5 setae and 1 setule terminally; exopod

indistinctly 8-segmented, setal formula 0,1,1,1,1,1,0,3.

Mandible: gnathobase with 3 cusped teeth, dorsalmost of

which bifid at tip; the medial part of gnathobase is damaged

and it is not known whether or not a tuft of setules is present.

Mandibular endopod (Fig. 11D) rudimentary, 1-segmented,

almost fused with basis, carrying 2 setae of unequal lengths;

first exopod segment with well developed seta, fifth segment

126

with non-reduced outer seta (Fig. 11D).

Maxillule (Fig. 12A): praecoxal arthrite with 5 bare spines

and 1 shorter process; coxal epipodite with 8 setae; coxal

endite with long, spinulose seta; vestigial basal seta present

(indicated by arrowhead); endopod bulbous, 1-segmented,

bearing 2 relatively long, spinulose setae terminally.

Maxilla: first praecoxal endite with 2 setae and 1 vestigial

element, second with 2 spinulose setae; basal spine (Fig. 11F)

with 2 rows of spinules; setae on endopod well developed,

ornamented with row of long, simple spinules along inner

margin (Fig. 11G). Maxilliped (Fig. 12B): innermost seta on

fourth and fifth endopod segments (indicated by arrowhead)

not reduced; seta a on sixth endopod segment reduced; seta b

relatively long; setae c and d simply ornamented with spinules

along inner margin.

Leg 1 with 2 outer spines on third exopod segment. Leg 4

with vestigial element on inner distal angle of coxa (Fig.

12C). Leg 5 (Fig. 12D-G): coxae and intercoxal sclerite

completely fused to form common base; coxa and basis

incompletely fused in right leg and separate in left. Right leg:

endopod (Fig. 12E) 1-segmented, spatulate, with minute

sensillum on outer proximal margin and tubular prominences

terminally; first exopod segment produced on outer angle,

with minute spine, second segment almost completely sepa-

rate from third, with 2 tufts of fine setules at inner distal

angle, minute sensillum at midlength of inner distal triangular

process (Fig. 12G) and outer terminal spiniform seta, third

segment triangular, tapering distally, with minute sensillum

at outer middle margin and short vestigial element termi-

nally; third segment with well developed muscles proximally.

Left leg: endopod (Fig. 12F) distinctly 2-segmented, first

segment produced terminally, second separate from first,

spatulate, covered by numerous fine setules on outer surface,

with attachment of muscles proximally; first exopod segment

similar to that of right leg, second expanded inwards with

outer seta subterminally, third segment small, separate from

second, with 2 elongate, chitinized processes terminally and

minute setule and scar of outer element proximally.

REMARKS. The fifth leg of the new genus exhibits a more

primitive state than Paraugaptilus in: (1) 2-segmented left

endopod; (2) both exopods 3-segmented. The right third exopo-

dal segment of Paraugaptiloides is certainly movable with well-

developed muscles originating in the preceding segment, while

the counterpart of Paraugaptilus is almost fused with the preced-

ing segment and has reduced musculature (see Figs 30F,32H). It

is probably not movable. In addition, the second segment of the

left endopod in Paraugaptiloides is likely to be movable as

indicated by the presence of a muscle extending between first

and second segments.

Genus Arietellus Giesbrecht, 1892

DIAGNOSIS (emended). Female. Body relatively large, mea-

suring approximately 3 to 7 mm in total length. Prosome

pointed or rounded frontally; cephalosome separate from

first pedigerous somite; last prosomal somite with pair of

blunt dorsolateral processes and paired ventrolateral pro-

cesses, symmetrical or asymmetrical, strongly or weakly

produced backwards. Genital double-somite longer than

wide, with pair of gonopores ventrolaterally and copulatory

pore ventromedially; seminal receptacle relatively large, bul-

bous, located laterally. Anal somite large; anal operculum

not developed. Caudal rami symmetrical, longer than wide,

S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

divergent or not, with well developed setae II to VII.

Antennule symmetrical, distinctly 20-segmented; posterior

margin fringed with long setules from segment I to X;

segments I to IV and XXIII to XXVIII fused; segments IV,

VI and XII without aesthetasc; compound segment

XXVI-XXVIII with 7 setae and aesthetasc. Antenna: first

endopod segment unarmed; second segment with 2 inner

setae, reduced in some species, and 5 setae and setule

terminally; exopod indistinctly 7- or 8-segmented, segment

VIII unarmed. Mandibular gnathobase lacking tuft of setules

at midlength; 3 cusped teeth on cutting edge, dorsalmost of

which bifid at tip. Mandibular palp: endopod absent; first

exopod segment with reduced or normal seta, outer seta on

fifth segment vestigial. Maxillule: praecoxal arthrite with 6

elements (5 spines and 1 process); coxal endite bearing 1

relatively short, thick seta, fringed with long setules; coxal

epipodite with 8 setae; outer basal seta absent; endopod

rudimentary, almost fused to basis or 1-segmented, bulbous,

with 1 seta terminally. Maxilla: first and second praecoxal

endites carrying 1 and 2 setae, respectively; basal spine with 2

rows of spinules; endopod setae armed with stout spinules

fringed with lamellar structure basally. Maxilliped: sgtal

formula of endopod segments of maxilliped: 1,4,4,3 or 2,3 or

2,4 (innermost seta on fourth and fifth segments reduced or

completely lacking in some species); setae a and b on sixth

segment vestigial.

First and third exopod segments of leg 1 bearing 1 and 2

outer spines respectively. Leg 5 reduced; coxae and inter-

coxal sclerite fused to form common transverse plate; basis

and coxa separate or fused; right basal seta longer than left;

endopod fused with basis, represented by small knob bearing

1 to 3 setae terminally, vestigial in some species; exopod

l-segmented, bulbous, carrying 1 terminal spine or almost

fused to basis, unarmed.

Male. Body as in female, about 4 to 6 mm in total length.

Left antennule 19-segmented, geniculate; segment XXI

fused with XXII; segments II and III with 1 seta; segment

XIII with seta; segments I to IX fringed with row of long

setules along posterior margin.

Second endopod segment of antenna with 1 long and 1

short seta medially; first exopod segment of mandible with

normally developed seta.

Leg 5: coxae and intercoxal sclerite fused to form common

plate; right coxa and basis incompletely fused; right basal seta

remarkably or normally elongate. Right leg: endopod

1-segmented, unarmed; exopod indistinctly 3-segmented, dis-

tal 2 segments incompletely fused, second segment with stout

process on inner angle, third segment spatulate, with 0-2

vestigial elements. Left leg: endopod indistinctly 2-segmented

or 1-segmented, unarmed; exopod 3-segmented, second seg-

ment expanded medially, third segment incompletely fused

with preceding one, bearing 2 terminal spines, with or

without outer minute spinule.

TYPE SPECIES. Arietellus setosus Giesbrecht, 1892 (mono-

typic).

OTHER SPECIES. A. aculeatus (T. Scott, 1894); A. giesbrechti

Sars, 1905; A. pavoninus Sars, 1905; A. plumifer Sars, 1905;

A. simplex Sars, 1905 (= A. major Esterly, 1906); A. armatus

Wolfenden, 1911; A. minor Wolfenden, 1911; A. pacificus

Esterly, 1913; A. tripartitus C.B. Wilson, 1950; A. sp. Brad-

ford, 1974; A. mohri (BjOrnberg, 1975), new combination; A.

sp. briefly described here.

PHYLOGENY OF ARIETELLID COPEPODS 127

XXIII-XXVIII—

——_—_—

Fig. 13. Arietellus plumifer, female. A, Genital double-somite, ventral view; B, Internal structure of right genital system; C, Antennulary

segments XXII to XXVIII; D, Antennary exopod; E, Mandibular exopod; F, Fifth segment of mandibular exopod, note reduced seta

indicated by arrowhead; G, Praecoxal arthrite, coxal endite and endopod of maxillule, rudimentary endopod indicated by arrowhead; H,

First and second praecoxal endites of maxilla; I, Basal spine of maxilla. Scales in mm.

128

REMARKS. The present study revealed that Paraugaptilus

mohri Bjornberg, 1975 belongs to the genus Arietellus (see

below). Arietellus shows sexual dimorphism in the antenna

and mandibular palp, as described in Paraugaptilus by

Deevey (1973). However, no sexual dimorphism is exhibited

in the maxillule, the maxilla and the maxilliped.

ECOLOGICAL NOTE. Species of the genus are pelagic and

distributed in deep water throughout the world’s oceans

(Brodsky, 1950; Vervoort, 1965; Roe, 1972, unpublished

data; Campaner, 1984).

Arietellus plumifer Sars, 1905 (Figs 13-15,17A,18L)

MATERIAL EXAMINED. 2 9 and CO’.

BODY LENGTH. Q 5.88 mm (28 VI 1985), 6.24 mm (26 XI

1965); CO 5.46 mm.

DESCRIPTION. Female. Cephalosome separate from first pedi-

gerous somite. Genital double-somite (Figs 13A,B,14) as long

as wide, almost symmetrical, with pair of gonopores ventrolat-

erally and anterior to single ventromedial copulatory pore;

paired copulatory ducts chitinized, each running anteriorly to

connect with seminal receptacle near genital operculum; semi-

nal receptacle located lateromedially, half as long as double-

somite, produced posteriorly with rounded posterior tip,

tapering anteriorly; receptacle duct beneath copulatory duct,

opening near inner corner of genital operculum.

Antennule symmetrical, 20-segmented; seventh (X) to

ninth (XII) segments and 11th (XIV) and 12th (XV) seg-

ments only partly fused. Fusion pattern and armature ele-

S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

ments as follows: I-IV—9 + 2 aesthetascs, V—2 + aesthetasc,

VI-2, VII-2 + aesthetasc, VIII-2, IX—2 + (small) aesthetasc,

X-2, XI-2 + aesthetasc; XII-2, XIIIJ-2 + aesthetasc, XIV—2

+ aesthetasc, XV-—2 +aesthetasc, XVI-2 + aesthetasc,

XVII-2 + aesthetasc, XVIII-2 + aesthetasc, XIX—2 +

aesthetasc, XX—2 + aesthetasc, XXI-2 + aesthetasc,XXII-1,

XXIII-XXVIII-12 + 2 aesthetascs (Fig. 13C). First (I-IV) to

seventh segments fringed with long setules along posterior

margin.

Antenna: first endopod segment without inner seta, second

segment with 2 short inner setae of unequal lengths (Fig.

15D) and 5 terminal setae and reduced setule terminally;

exopod indistinctly 7-segmented; setal formula 0,1,1,1,1,0,3.

Mandibular palp (Fig. 13E,F): endopod absent; first exopod

segment having relatively reduced seta, fifth segment carry-

ing normal seta and vestigial element.

Maxillule: praecoxal arthrite (Fig. 13G) with 5 naked

spines, 1 short process and row of long setules; coxal endite

(Fig. 13G) carrying relatively long spinulose seta, fringed

with numerous long setules along distal margin; basal seta

lacking; endopod (Fig. 13G, indicated by arrowhead) rudi-

mentary, almost fused with basis, unarmed. Maxilla: first

praecoxal endite bearing thick naked seta and vestigial ele-

ment, second praecoxal endite having 2 spinulose setae (Fig.

13H); basal spine (Fig. 131) with 2 rows of minute spinules

along ventral margin.

Maxilliped: sixth endopod segment (Fig. 15SA,B) having

elongate seta d with row of stout spinules whose base

ornamented with lamellar projection (Fig. 15C), finely ser-

rated, medial-length seta c and reduced setae a and b.

Fig. 14. Arietellus plumifer, female. SEM micrographs of genital double-somite of female. A, Genital double-somite, ventral view showing

large copulatory pore (indicated by an arrow), scale bar = 100 pm; B, Right gonopore, scale bar = 30 4m; C, Copulatory pore, scale bar =

20 pm.

PHYLOGENY OF ARIETELLID COPEPODS 129

0.1

Fig. 15. Arietellus plumifer, female (A-D), male (E-G). A, Fourth and fifth endopod segments of maxilliped, innermost vestigial seta

indicated by arrowhead; B, Sixth endopod segment of maxilliped; C, Spinule on seta d of sixth endopod segment of maxilliped; D,

Mid-margin setae on second segment of antennary endopod; E, Left antennulary segments XIX to XXVIII; F, Second endopod segment of

antenna; G, Mandibular exopod. Scales in mm.

130

Leg 1: third exopod segment with 2 subterminal serrate

spines.

Leg 5 (Fig. 17A): coxae incompletely fused with intercoxal

sclerite; right basal seta extremely elongate; endopod repre-

sented by knob with 2 plumose setae; exopod incompletely

fused with basis, 1-segmented, carrying 1 terminal spine.

Male. Left antennule (Fig. 15E) distinctly 19-segmented;

8th to 11th segments only partly fused near posterior margin;

fusion pattern and armature elements as follows: I-ITV—7 + 2

aesthetascs (I-3 + aesthetasc, II-1 + aesthetasc, IIJ-1 +

aesthetasc, IV—2 + aesthetasc), V—2 + aesthetasc, VI-2 +

aesthetasc, VII—2 + aesthetasc, VIIJ-2 + aesthetasc, [IX—2 +

aesthetasc, X-1 + aesthetasc + process, XI—2 + aesthetasc,

XII-1 + aesthetasc + process, XIII-1 + aesthetasc + pro-

cess, XIV—1 + 2 aesthetascs + process, XV—1 + aesthetasc +

process, XVI-2 + aesthetasc, XVII-2 + aesthetasc, XVIII-2

+ aesthetasc, XIX—1 + aesthetasc + 2 processes, XX—1 +

aesthetasc + process, XXI-XXIII-2 + aesthetasc + 2 pro-

cesses (XXI-aesthetasc + 2 processes, XXII-1, XXIII-1),

XXIV-XXVIII-11 + 2 aesthetascs (XXIV-1 + 1, XXV-1 +

1 + aesthetasc, XXVI-XXVIII-7 + aesthetasc); no suture

visible between segments XXV and XXVI. First (I-IV) to

sixth (IX) segments fringed with long setules along posterior

margin.

Antenna: second endopod segment (Fig. 15F) with 1 short

and 1 long seta medially and 5 setae and 1 vestigial setule

terminally. Mandibular palp (Fig. 15G): first exopod segment

with well-developed seta.

Leg 5 (Fig. 18L): both coxae fused to intercoxal sclerite to

form common plate, right coxa almost fused with basis, left

coxa completely separate from basis. Right leg: basal seta

considerably elongate; endopod 1-segmented, spatulate; exo-

pod indistinctly 3-segmented, first segment with 1 spine on

outer corner, second incompletely fused with third, furnished

with triangular process and 2 tufts of fine setules on inner

corner and 1 spine on outer corner, third segment spatulate,

with subterminal outer setule and terminal vestigial element.

Left leg: endopod indistinctly 2-segmented, first and second

segments unarmed; exopod indistinctly 3-segmented, first

segment with 1 spine on outer corner, second segment

incompletely fused with third, expanded inwards, bearing 1

subterminal outer spine, third segment small, having minute

spinule and 2 spines almost fused basally with segment,

terminal one bifid at tip.

Arietellus mohri (BjOrnberg, 1975), new combination

(Figs 16A, 17C, 18A,B,F,H)

MATERIAL EXAMINED. Q, U.S. National Museum, reference

number USNM 150095.

BODY LENGTH. 6.40 mm (after Bj6rnberg, 1975)

DESCRIPTION. Female. Cephalosome separate from first

pedigerous somite. Genital double-somite (Fig. 16A) as long

as wide, with anterior pair of gonopores located ventrolater-

ally anterior to single ventromedial copulatory pore as in A.

plumifer; copulatory ducts much more chitinized and wider

than in A. plumifer, slightly asymmetrical, left duct divergent

into blind tubule near left genital operculum; seminal recep-

tacle the same shape as in A. plumifer.

Right antennule (left antennule missing distal segments)

with fusion pattern and armature as A. plumifer except for

missing elements. Antenna: first endopod segment unarmed,

second segment with 2 inner setae of unequal length medially

S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

and 5 setae and 1 vestigial setule terminally; exopod

7-segmented, setal formula: 0,1,1,1,1,0,3. Mandibular palp

(Fig. 18A,B): endopod absent; exopod 5-segmented, first to

fourth segments each with 1 seta, first segment with well-

developed seta, fifth segment with 1 long seta and vestigial

seta.

Maxillule: praecoxal arthrite with 5 naked spines and 1

bare process; coxal endite with 1 naked, thick seta terminally,

fringed with long setules along ventral margin; coxal epi-

podite with 8 setae; endopod absent. Maxilla: first praecoxal

endite with long, bare seta and 1 vestigial element; basal

spine (Fig. 18F) with 2 rows of spinules along ventromedial

margin. Maxilliped: fourth and fifth endopod segments each

with only 2 well developed setae and lacking innermost seta,

sixth segment (Fig. 18H) with vestigial seta a and short seta b.

Leg 1: basis with inner and, possibly, outer (scar present on

outer margin) setae; third exopod segment with 2 lateral

bipinnate spines. Leg 4, possibly, with 1 basal outer seta (scar

present). Leg 5 (Fig. 17C): coxa and intercoxal sclerite almost

fused, but suture line visible on left side; basis completely

fused with coxa. Right leg: outer basal seta more elongate

than left one; endopod represented by small knob with

vestigial element at tip; exopod almost fused with basis, but

suture visible only on anterior surface, unarmed, round. Left

leg: basis with concavity on inner margin; outer basal seta

thick, plumose; endopod reduced to low prominence with

spinulose seta terminally; exopod almost completely fused

with basis, unarmed, round.

REMARKS. Bjérnberg (1975) assigned one female of a new

species collected from the southeastern Pacific (depths:

1932-3142 m) to the genus Paraugaptilus, probably because

of the remarkably reduced fifth legs. The present

re-examination revealed that it belongs to Arietellus not to

Paraugaptilus, on the basis of the following characters: (1)

the genital double-somite with single copulatory pore ventro-

medially; (2) the first, sixth and 10th antennulary segments

carrying 2, 1 and 1 aesthetascs, respectively; (3) the antennu-

lary segment XX VI-XXVIII with 7 setae and 1 aesthetasc;

(4) the coxal endite of maxillule bearing 1 relatively well

developed seta and fringed with long setules along ventral

margin; (5) the second praecoxal endite of maxilla having 2

setae; (6) the endopodal setae of maxilla carrying sharp

spinules with lamellar structure basally; (7) the fourth and

fifth endopodal segments of maxilliped lacking innermost

seta; (8) leg 4 without inner coxal seta; (9) leg 5 with distinct

distal lobe derived from exopod; (10) the right basal seta of

leg 5 considerably elongate.

Although Bj6rnberg (1975) described the species in rela-

tively great detail, the present re-examination of the holotype

revealed that her description included several misinterpreta-

tions, particularly in the mouthparts and legs. These are

amended in the present description.

Arietellus aculeatus (T. Scott, 1894) (Figs

16F,G,18D,E,O)

MATERIAL EXAMINED. 9 and2 0'C’.

BODY LENGTH. 9 4.62 mm; CO 3.77, 3.79 mm.

DESCRIPTION. Female. Cephalosome separate from first

pedigerous somite. Left antennule similar to that of female

A. plumifer except for following points: segments VIII and X

each with minute aesthetasc; segment XIV carrying 2 setae

PHYLOGENY OF ARIETELLID COPEPODS

and 2 aesthetascs. Antenna: second endopod segment (Fig.

16G) with 2 short inner setae medially and 5 setae and 1

vestigial seta terminally. Mandibular palp (Fig. 18D): first

exopod segment with reduced, short seta.

Male. Cephalosome separate from first pedigerous somite.

Left antennule exhibiting same fusion pattern and armature

elements as A. plumifer except for first segment: I-IV-7 + 7

aesthetascs (I-3 + aesthetasc, II-1 + 2 aesthetascs, III-1 + 2

aesthetascs, [V—2 + 2 aesthetascs). Antenna: second endo-

pod segment (Fig. 16F) bearing 1 long and 1 short seta

medially. Mandibular palp (Fig. 18E): first exopod segment

with well-developed seta. Maxillule: endopod almost fused

with basis, represented by small knob. Maxilliped: fourth and

fifth endopod segments each having vestigial innermost seta,

as in A. plumifer. Leg 5: left endopod (Fig. 18O) indistinctly

2-segmented, with suture visible on posterior surface; com-

pound distal exopod segment of right leg with minute termi-

nal element.

REMARKS. A. aculeatus exhibits sexual dimorphism in the

antenna and mandibular palp, as does A. plumifer.

Arietellus setosus Giesbrecht, 1892 (Figs 16J,181,M)

MATERIAL EXAMINED. CO’.

BODY LENGTH. 4.28 mm.

DESCRIPTION. Male. Cephalosome separate from first pedi-

gerous somite. Left antennule with same fusion pattern and

armature as A. plumifer. Antenna: exopod indistinctly

7-segmented; setal formula 0,1,1,1,1,0,3. Mandible: first exo-

pod segment with normally developed seta. Maxillulary

endopod (Fig. 16J) represented by unarmed, small knob.

Maxilla and maxilliped (Fig. 181) as in A. plumifer. Leg 5: left

endopod (Fig. 18M) indistinctly 2-segmented as in A. plumi-

fer, first segment produced ventrally to rounded tip, second

segment rising from inner side of first segment; terminal spine

on third exopod segment of left leg almost completely fused

to segment, subterminal spine incompletely coalesced with

segment; distal compound exopod segment of right leg

unarmed.

Arietellus pavoninus Sars, 1905 (Figs 16B,H,17B,18J)

MATERIAL EXAMINED. Q.

BODY LENGTH. 5.00 mm.

DESCRIPTION. Female. Cephalosome separate from first

pedigerous somite. Genital double-somite (Fig. 16B) similar

to that of A. plumifer, but readily distinguishable since

seminal receptacle relatively much larger than in A. plumifer,

over half length of genital double-somite.

Antennule with same fusion pattern and armature as A.

plumifer except for absence of aesthetasc on segment IX (this

aesthetasc may have been detached). Mouthparts similar to

those of female A. plumifer except for maxillulary endopod.

Maxillule (Fig. 16H): endopod distinctly 1-segmented, bul-

bous with 1 bipinnate seta. Maxilliped (Fig. 18J): fourth and

fifth endopod segments each with reduced innermost seta,

sixth segment with reduced setae a and b. Leg 5 (Fig. 17B):

coxae incompletely fused with intercoxal sclerite, in particu-

lar, more fused in right leg; endopod represented by 2

plumose setae not so produced as in A. plumifer; exopods

131

1-segmented, separate from basis, carrying 1 unipinnate spine

terminally.

Arietellus simplex Sars, 1905 (Figs 16E,1,18N)

MATERIAL EXAMINED. CO’.

BODY LENGTH. 6.10 mm.

DESCRIPTION. Male. Cephalosome separate from first pedi-

gerous somite. Left antennule with same fusion pattern and

armature as A. plumifer. :

Antenna: exopod (Fig. 16E) indistinctly 8-segmented; setal

formula 0,1,1,1,1,0,0,3. Mandible: first exopod segment with

normally developed seta. Maxillule: endopod represented by

low knob, almost fused with basis (Fig. 161). Maxilliped as in

A. plumifer. Leg 5: left endopod (Fig. 18N) indistinctly

2-segmented, suture visible on both surfaces; terminal and

- subterminal spines on third exopod segment of left leg

incompletely fused to segment, terminal spine with 4 minute

spinules terminally; terminal spine of distal compound exo-

pod segment of right leg unarmed.

Arietellus sp. (Figs 16C,D,17D,18C,G,K)

MATERIAL EXAMINED. Q.

BODY LENGTH. 5.15 mm.

DESCRIPTION. Female. Cephalosome separate from first

pedigerous somite. Posterolateral angles of prosome asym-

metrically produced into sharp lateral processes as in A.

giesbrechti (see Sars, 1924, 1925), left process slightly longer

and more produced than right. Genital double-somite (Fig.

16C) similar to that of A. mohri in having pair of laterally

expanded copulatory ducts, but differing in presence of better

developed muscles to genital operculum.

Left antennule with same segmentation and armature as A.

plumifer. Antennary endopod: first segment unarmed, sec-

ond (Fig. 16D) with 1 long and 1 short seta medially, and 5

setae and 1 vestigial element terminally. Mandibular palp

(Fig. 18C) with relatively long seta on first exopod segment.

Maxillulary endopod completely fused with basis. Maxilla:

basal spine (Fig. 18G) with 2 rows of spinules along ventral

margin. Maxilliped (Fig. 18K): fourth and fifth endopod

segments lacking innermost seta; sixth endopod segments

with setae a and b reduced.

Leg 5 (Fig. 17D) similar to that of A. mohri with intercoxal

sclerite, coxa, basis and both rami almost completely fused,

but distinguishable by: seta on both endopods represented by

low knob much better developed than in A. mohri; unarmed,

lobate exopods more developed than in A. mohri; left basal

seta longer than in A. mohri.

REMARKS. Arietellus sp., an as yet undescribed species, is

most closely related to A. mohri in having synapomorphic

characters such as no innermost seta on the fourth and fifth

endopodal segments of maxilliped and the reduced leg 5.

Genus Rhapidophorus Edwards, 1891

TYPE SPECIES. Rhapidophorus wilsoni

(monotypic).

Edwards, 1891

REMARKS. Fosshagen (1968) first pointed out the affinity of

this genus with Paramisophria. Campaner (1977) later

assigned the genus to the family Arietellidae. The genus,

132 S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

\ (SO) y

Fig. 16. Arietellus mohri, female (A); A. pavoninus, female (B,H); A. sp., female (C,D); A. simplex, male (E,I); A. aculeatus, female (G),

male (F); A. setosus, male (J). A-C, Genital double-somite, ventral view; D,F,G, Second endopod segment of antenna; E, Antennary

exopod; H, Praecoxal arthrite, coxal endite and endopod of maxillule, endopod indicated by arrowhead; I,J, Maxillulary endopod. Scales

in mm.

133

PHYLOGENY OF ARIETELLID COPEPODS

————— yy

yee

iy ——

Fig. 17. Fifth legs of females of Arietellus. A, A. plumifer; B, A. pavoninus; C, A. mohri, vestigial element on right endopod represented by

low knob incorporated in C; D, A. sp. Scales in mm.

134 S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

Fig. 18. Arietellus mohri, female (A,B,F,H); A. sp., female (C,G,K); A. aculeatus, female (D), male (E,O); A. setosus, male (I); A.

pavoninus, female (J); A. plumifer, male (L); A. setosus, male (M); A. simplex, male (N). A,C, Mandibular exopod; B, Fifth exopod

segment of mandible; D,E, First exopod segment of mandible; F,G, Maxillary basal spine; H, Sixth endopod segment of maxilliped; I-K,

Fourth to sixth endopod segments of maxilliped; L, Leg 5, anterior surface; M-O, Left endopod of leg 5. Scales in mm.

PHYLOGENY OF ARIETELLID COPEPODS

however, has peculiar characters in the mandibular palp,

maxillule, maxilliped and leg 1 as indicated by Fosshagen

(1968). We were unable to re-examine the male type speci-

men; it is deposited neither in the Berlin Zoological Museum

(Dr. H.-E. Gruner, personal communication) nor at the

University of Leipzig (Prof. K. Dréssler, personal communi-

cation), and may no longer be extant. Since Edwards’ (1891)

description is not accurate enough to compare Rhapidopho-

rus with the other genera, the present study does not include

the genus in the cladistic analysis.

ECOLOGICAL NOTE. Rhapidophorus was found in the water-

lung of a holothurian collected from the Bahamas, but was

stated to be free-living (Edwards, 1891). The compact body,

short antennule and stout legs suggest that it may originally

have been hyperbenthic.

Genus Paramisophria T. Scott, 1897

DIAGNOsIS. The diagnostic characters of the genus have

already been given in detail by Ohtsuka et al. (1993a).

Supplemental diagnostic characters are given briefly here.

Body lengths of female and male approximately 0.6 to 3

mm and 0.6 to 2 mm, respectively. Female antennules:

segments I-III fused; segments III and IV separate; segment

IV without aesthetasc; segments XXIII and XXIV separate;

posterior margin fringed with long setules from I to X. Male

left antennule: segments II and III with 1 seta; segment XIII

with 1 seta; segments XXI and XXII fused. Antenna: first

endopod with inner medial seta, second segment with 3 inner

setae at midlength, and 5 setae and 1 minute seta terminally;

exopod indistinctly 8- or 9-segmented, segment VIII with

seta. Mandibular gnathobase lacking or having a small tuft of

setules medially, with 3 teeth on cutting edge, dorsalmost of

which bifid at tip. Mandibular palp: seta on first exopod

segment not reduced; outer seta on fifth exopod segment

relatively long. Maxillulary coxal epipodite with 8 setae.

Maxilla: first praecoxal endite with 1-2 setae and vestigial

element, second endite with 2 setae. Maxilliped: setal for-

mula of endopod 1,4,4,3,3,4; innermost seta on fourth and

fifth endopod segments not rudimentary, setae a and b on

sixth segment not reduced.

TYPE SPECIES. Paramisophria cluthae T. Scott, 1897 (mono-

typic).

OTHER SPECIES. P. spooneri Krishnaswamy, 1959; P. ammo-

phila Fosshagen, 1968; P. giselae (Campaner, 1977); P. itoi

Ohtsuka, 1985; P. variabilis McKinnon and Kimmerer, 1985;

P. platysoma Ohtsuka and Mitsuzumi, 1990; P. japonica

Ohtsuka, Fosshagen and Go, 1991; P. fosshageni Othman

and Greenwood, 1992; P. reducta Ohtsuka, Fosshagen and

Iliffe, 1993; P. galapagensis Ohtsuka, Fosshagen and Iliffe,

1993; P. cluthae sensu Tanaka (1966).

REMARKS. Parapseudocyclops Campaner, 1977 was synony-

mized with the genus Paramisophria (Ohtsuka et al., 1991).

ECOLOGICAL NOTE. Paramisophria is mainly distributed in

the near-bottom communities on the continental shelf (Oht-

suka et al., 1991), but also colonizes marine caves (Ohtsuka

et al., 1993a).

135

Paramisophria japonica Ohtsuka, Fosshagen and Go,

1991 (Figs 19,20F)

MATERIAL EXAMINED. Q.

BODY LENGTH. 1.85—2.08 mm (after Ohtsuka et al., 1991).

DESCRIPTION. Female. Genital double-somite (Fig. 19A)

wider than long, with pair of gonopores anteroventrally and

single copulatory pore ventromedially; seminal receptacle

located lateromedially; copulatory duct thin.

Antennule: segments X to XII, and XIV and XV only

partly fused near posterior margin; segments XX V and XXVI

incompletely fused; segments I to X fringed by long setules

along posterior margin; fusion pattern and armature as

follows: I-IIIJ—7 + 2 aesthetascs (I-3 + aesthetasc, II-2, III-2

+ aesthetasc), [V-2, V-2 + aesthetasc, VI-2 + aesthetasc,

VII-2 + aesthetasc, VIII-2 + aesthetasc, [X-2 + aesthetasc,

X-2 + aesthetasc, XI—2 + aesthetasc, XII-2 + aesthetasc,

XIII-2 + aesthetasc, XIV-2 + aesthetasc, XV-2 + aes-

thetasc, XVI-2 + aesthetasc, XVIJ-2 + aesthetasc, XVIII-2

+ aesthetasc, XIX-2 + aesthetasc, XX-2 + aesthetasc,

XXI-2 + aesthetasc, XXII-1, XXIII-1, XXIV-XXVIII-12

+ 2 aesthetascs (XXIV-1 + 1, XXV-1 + 1 + aesthetasc,

XXVI-XXVIII-8 + aesthetasc).

Maxilla: first praecoxal endite with 1 seta and vestigial

element, second with 2 finely spinulose setae (Fig. 19C);

basal spine naked. Maxilliped: fourth and fifth segments (Fig.

19D) with relatively long innermost seta; sixth segment (Fig.

19E) with setae a and b not reduced.

Leg 5 (Fig. 20F): coxae and intercoxal sclerite almost

completely fused to form common base; endopod almost

completely fused to basis with fine suture visible on posterior

surface; first exopod segment clearly separate from second;

second and third exopod segments completely fused.

Paramisophria giselae (Campaner, 1977) (Fig. 20A-E)

MATERIAL EXAMINED. Q, holotype, Museu de Zoologia,

University of Sao Paulo, reference number 4004. Q,

paratype, Zoology Department, Instituto de Biociéncias,

University of Sao Paulo, number 173.

BODY LENGTH. 2.55, 2.60 mm (after Campaner, 1977).

DESCRIPTION. Posterior lateral.corners of second and third

pedigerous somites asymmetrically produced: corners more

sharply pointed on right side than on left. Genital double-

somite (Fig. 20A) longer than wide; genital system similar to

that of P. japonica, but differing in: copulatory pore located

on right side; seminal receptacle located near gonopore;

copulatory pore relatively thick.

Antennary exopod (Fig. 20B) indistinctly 9-segmented;

terminal segment with 2 long plumose setae and vestigial

seta. Mandibular gnathobase with small tuft of setules medi-

ally; 3 teeth on cutting edge, dorsalmost of which bifurcate at

tip. Mandibular palp similar to that of P. japonica: endopod

rudimentary, l-segmented, with 2 setae of unequal lengths;

seta on first exopod segment not reduced; outer seta on fifth

exopod segment relatively long. Maxillule similar to that of

P. japonica except for relatively long seta on coxal endite:

praecoxal arthrite with 5 naked spines and 1 process; coxal

epipodite with 8 setae; small basal seta present; endopod

bulbous, 1-segmented with 3 setae of unequal lengths. Max-

illa: first praecoxal endite (Fig. 20C) with 2 spinulose setae

and rudimentary element, second (Fig. 20C) bearing 2 spinu-

136

S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

Fig. 19. Paramisophria japonica, female. A, Genital double-somite, ventral view; B, Antennary exopod; C, First and second praecoxal

endites of maxilla; D, Fourth and fifth endopod segments of maxilliped; E, Sixth endopod segment of maxilliped. Scales in mm.

lose setae; basal spine (Fig. 20D) naked. Maxilliped with

same setal formula as P. japonica.

Leg 5 (Fig. 20E): coxae and intercoxal sclerite clearly

separate; setation and spinulation as in P. japonica; endopod

completely fused to basis; exopod almost completely fused to

basis with fine suture visible; first and second exopod seg-

ments fused with suture clearly visible on posterior surface;

second and third exopod segments completely fused.

REMARKS. Re-examination of the holotype and paratype

revealed the following: (1) since the antennules of both types

are missing (the proximal half remains on one side only), we

were unable to check the fusion and armature patterns; (2)

the terminal segment of the antennary exopod has only 2

developed setae plus 1 minute seta although 3 developed

setae were shown in the original description (Campaner,

1977); (3) the dorsalmost tooth on the mandibular gnatho-

base is bicuspid although it was originally drawn as monocus-

pid (Campaner, 1977); (4) the terminal segment of the

mandibular exopod has 2 relatively well developed setae (one

about 25% shorter than the other); (5) the setae on the

mandibular endopod are missing but there are 2 scars visible,

of different sizes, which suggests 2 unequal setae; (6) the

coxal epipodite of the maxillule of the holotype is damaged: 5

long setae are present, then a gap due to damage, then a short

seta; although the gap does not show clean scars where setae

were broken off, the gap is only big enough for 2 setae —

giving a total of 8 setae as in the paratype; (7) the first to sixth

endopodal segments of the maxilliped bearing 1, 4, 4, 3, 3 and

4 setae, respectively; (8) no seta originating from the poste-

rior surface of the first exopodal segment of leg 4.

Paramisophria reducta Ohtsuka, Fosshagen and Iliffe,

1993

Museum, BM (NH) Reg. No 1992. 1093.

BODY LENGTH. 1.60 mm (after Ohtsuka et al., 1993a).

DESCRIPTION. Male. Left antennule: segments XXI to

XXIII, XXIV and XXV, and XXVI to XXVIII completely

fused; segments XXIII and XXIV, and XXV and XXVI

incompletely fused; fusion pattern and armature elements as

follows: I-IV-7 + 4 aesthetascs (I-3 + aesthetasc, II-1 +

aesthetasc, IJI-1 + aesthetasc, IV-2 + aesthetasc), V—2 +

aesthetasc, VI-2 + aesthetasc, VII-2 + aesthetasc, VIII-2 +

aesthetasc, [IX—2 + aesthetasc, X—-1 + aesthetasc + process,

XI-2 + aesthetasc, XIJ-1 + aesthetasc + process, XIII-1 +

aesthetasc + process, XIV—1 + aesthetasc + process, XV-2

+ aesthetasc, XVI-2 + aesthetasc, XVII-2 + aesthetasc,

XVIII-2 + aesthetasc, XIX—-1 + aesthetasc + 2 processes,

XX-1 + aesthetasc + process, XXI-XXIII-2 + aesthetasc +

2 processes (XXI-aesthetasc + 2 processes, XXII-l,

XXIII-1), XXIV-XXV-4 + aesthetasc (XXIV-1 + 1,

XXV-1 + 1 + aesthetasc), XXVI-XXVIII-8 + 2 aes-

thetascs.

PHYLOGENY OF ARIETELLID COPEPODS 137

| Fig. 20. Paramisophria giselae, female (A-E); P. japonica, female (F). A, Genital double-somite, ventral view; B, Antennary exopod; C,

First and second praecoxal endites of maxilla; D, Basal spine of maxilla; E,F, Leg 5, anterior surface. Scales in mm.

138

REMARKS. The fusion pattern of the antennulary segments is

slightly different from the male of P. japonica in which

segments XXI and XXII are incompletely fused whereas

segments XXIII and XXIV are separate.

Genus Metacalanus Cleve, 1901

DIAGNOSIS (emended). Female. Body compact, small, mea-

suring approximately 1 mm in body length. Prosome oval in

dorsal view, not produced frontally; cephalosome and first

pedigerous somite separate or weakly fused; posterior cor-

ners of last prosomal somite produced to form ventrolateral

lobe, without dorsolateral processes; urosome short, less than

one-third length of prosome. Genital double-somite wider

than long, with ventrolateral pair of gonopores or only right

gonopore (left reduced) located posteriorly; paired copula-

tory pores small, located near inner corner of genital aperture

(in the case of reduction of left gonopore, only right copula-

tory pore present); anal operculum either developed, triangu-

lar or not. Caudal rami symmetrical, longer than wide, with

seta II reduced or completely lacking; seta III relatively

small.

Antennules asymmetrical, left longer than right and reach-

ing to end of prosome, different in fusion pattern and

armature; indistinctly 18- or 20-segmented in right antennule,

16- or 18-segmented in left; posterior proximal margin lack-

ing long setules; segments I-IV up to VI; segments IX and X

fused; segments XII to XIV fused in left; segments II, VII

and IX with 1 or 2 setae; segment XIII with 1 seta; segments

II, [V, VI, VIII and X lacking aesthetasc; segments V, XII

and XIII with or without aesthetasc; compound segment

XXVI-XXVIII with 8 setae and aesthetasc. Antenna: first

endopod segment with 1 inner seta, second with 2 setae

medially and 5 setae terminally; exopod indistinctly

7-segmented. Mandibular gnathobase lacking tuft of setules;

4 teeth on cutting edge, dorsalmost of which trifid at tip.

Mandibular palp: endopod almost fused to basis, represented

by small knob with 1 or 2 setae terminally; seta on first

exopod segment not reduced; outer seta on fifth exopod

segment relatively long. Maxillule: praecoxal arthrite with

0-2 spines; coxal endite with or without 1 short seta; coxal

epipodite with 5 setae; endopod absent or 1-segmented,

bulbous with 1 seta. Maxilla: first praecoxal endite with 1 seta

and 1 rudimentary element; basal spine with 2 rows of minute

spinules proximally; endopodal setae with row of spinules

along inner margin. Maxilliped: setal formula on first to sixth

endopod segments 1,4,4,3,3,4; innermost seta on fourth and

fifth endopod segments not reduced; only distalmost seta on

these segments well-chitinized and long; setae a and b on

sixth endopod segment not reduced.

Third exopod segment of leg 1 with single outer spine. Leg

5: coxae separate from intercoxal sclerite; endopod repre-

sented by 1 seta or completely absent; exopod and basis fused

or separate; exopod either 1-segmented, with 1-3 spines or

represented by small knob bearing 1| seta.

Male. Body as in female, measuring less than 1 mm in body

length. Left antennule 16-segmented; segments I-IV, IX-X

and XII-XIV fused; segment XIII without seta; segment XXI

separate from XXII. Leg 5: coxae and intercoxal sclerite

fused; basis separate from coxa; endopod absent; exopod

3-segmented, third segment with large seta almost fused with

segment.

TYPE SPECIES. Metacalanus aurivilli Cleve, 1901 (= Scottula

S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

ambariakae Binet and Dessier, 1968) (monotypic).

OTHER SPECIES. M. inaequicornis (Sars, 1902); M. acutioper-

culum Ohtsuka, 1984; M. curvirostris Ohtsuka, 1985; M.

species 1 and 2 from Okinawa.

REMARKS. Metacalanus was recognized as a senior synonym

of Scottula Sars, 1902 by Campaner (1984).

ECOLOGICAL NOTE. M. aurivilli seems to be epipelagic in

subtropical waters in the Indo-West Pacific (cf. Greenwood,

1978). Other species are hyperbenthic in shallow waters in

temperate and subtropical regions (cf. Sars, 1903; Ohtsuka,

1984, 1985), or are marine cave-dwellers (Ohtsuka et al.,

1993a).

Metacalanus species 1 (Figs 21B-I,23,25A,26A-G)

MATERIAL EXAMINED. 4 99 and O.

BODY LENGTH. Q 0.81, 0.83, 0.83, 0.84 mm; O' 0.77 mm.

DESCRIPTION. Female. Cephalosome only partly fused with

first pedigerous somite. Genital double-somite (Figs

21B,23B) wider than long, asymmetrical, left gonopore and

copulatory pore completely absent; right gonopore located

near posteroventral margin of double somite, anterior half

opening, covered by oval flap, possibly derived from leg 6;

outer half gonopore frilled with cuticular flap (Fig. 23A);

copulatory pore (Fig. 23C) small, oval in shape, approxi-

mately 4.0 1m in long axis and 1.0 wm in short axis, near

inner distal corner of gonopore (copulatory pore blocked by

spermatophore remnant); single seminal receptacle large,

about half width of somite, located ventromedially; copula-

tory duct short, curved. Anal operculum triangular as in M.

acutioperculum.

Antennules asymmetrical, left longer than right, different

in fusion pattern and armature (see Fig. 22). Right antennule:

segments X to XII and XIV and XV fused only partly near

posterior margin; fusion pattern and armature as follows:

I-IV-9 + 2 aesthetascs (I-3 + aesthetasc, II-2, III-2 +

aesthetasc, IV—2), V-2 + aesthetasc, VI-2, VII-2 + aes-

thetasc, VIII-2, [IXx-X-4 + aesthetasc (IX-2 + aesthetasc,

X-2), XI-2 + aesthetasc, XII-2 + aesthetasc, XIIJ-1 +

aesthetasc, XIV-—2 + aesthetasc, XV-—2 + aesthetasc, XVI-2

+ aesthetasc, XVII-2 + aesthetasc, XVIII-2 + aesthetasc,

XIX-2 + aesthetasc, XX—1 + aesthetasc, XXI-2 + aes-

thetasc, XXII-1, XXIII-1, XXIV—-XXVIII-12 + 2 aes-

thetascs (XXIV-1 + 1, XXV-1 + 1 + aesthetasc,

XXVI-XXVIII-8 + aesthetasc). Left antennule different

from right one in following: segments XII to XIV fused, with

5 setae and 2 aesthetascs (XII-2, XIII-1 + aesthetasc, XITV-—2

+ aesthetasc); segment XX with 2 setae and aesthetasc.

Antenna: exopod (Figs 21E,25A) indistinctly 7-segmented;

setal formula 0,1,1,1,1,1,3 (2 setae and vestigial element).

Mandibular palp (Fig. 21F): endopod 1-segmented, almost

fused with basis, with 2 setae of unequal lengths; first exopod

segment carrying long seta, fifth segment bearing 2 normal

setae of unequal lengths. Maxillule (Fig. 21G): praecoxal

arthrite with 2 slender setae; coxal endite having 1 short seta;

coxal epipodite having 5 setae; basal seta absent; endopod

(indicated by arrowhead) 1-segmented, bulbous, with short

seta terminally. Maxilla: first and second praecoxal endites

with 1 seta plus 1 vestigial element and 2 setae respectively

(Fig. 21H); basal spine (Fig. 211) with 2 rows of short spinules

proximally. Maxilliped: fourth and fifth endopod, segments

PHYLOGENY OF ARIETELLID COPEPODS 139

H 3

pees SIE

Fig. 21. Metacalanus sp. 1, female (B,E-I), male (C,D); Metacalanus sp. 2, female (A). A,B, Genital double-somite, ventral view; C, Left

antennulary segments I to XVII; D, Left antennulary segments XVIII to XXVIII; E, Antennary exopod; F, Mandibular palp; G, Praecoxal

arthrite, coxal endite and endopod of maxillule, endopod indicated by arrowhead; H, First and second praecoxal endites of maxilla; I,

Basal spine of maxilla. Scales in mm.

S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

Fig. 24. Metacalanus sp. 2, female. SEM micrographs of genital double-somite. A, Genital somite, copulatory pores indicated by arrows,

scale bar = 20 um; B, Left gonopore and copulatory pore (indicated by an arrow), scale bar = 10 4m; C, Right copulatory pore, scale bar

= 2 wm; D, Left copulatory pore, scale bar = 2 um.

PHYLOGENY OF ARIETELLID COPEPODS

143

Fig. 25. Metacalanus sp. 1, female (A); Metacalanus sp. 2, female (B). SEM micrographs of mouthparts. A, Detail of segments IV to VIII

of antennary exopod, scale bar = 10 »m; B, Mandibular endopod, indicated by arrow, scale bar = 5 pm.

Metacalanus species 2 (Figs 21A,24,25B,26H)

MATERIAL EXAMINED. 499.

BODY LENGTH. 0.84, 0.84, 0.86, 0.88 mm.

DESCRIPTION. Female. Cephalosome separate from first

pedigerous somite. Lateral lobe of last prosomal somite

produced backwards reaching halfway along second urosomal

somite (Fig. 24A). Genital double-somite (Figs 21A,24A)

wider than long, symmetrical, with paired gonopores and

copulatory pores located ventrolaterally near posterior end of

somite; each gonopore lacking outer cuticular lateral flap

found in M. species 1, anterior half opening, covered by oval

flap; copulatory pore (Fig. 24C,D) small, round, ca. 1.4 pm

_ in diameter, located near anterior inner corner of gonopore

(spermatophore remnant attached to opening). Internal geni-

tal system similar to that of M. species 1. Anal operculum

triangular as in M. species 1.

Antennule asymmetrical, left longer than right, different in

fusion pattern and armature (see Fig. 22). Right antennule:

segments X to XI, and XIV and XV only partly fused near

posterior margin; fusion pattern and armature as follows:

I-VI-12 + 2 aesthetascs (I-3 + aesthetasc, II-1, III-2 +

aesthetasc, IV-2, V—2, VI-2), VII-1 + aesthetasc, VIII-1,

IX-X-3 + aesthetasc (IX-1 + aesthetasc, X-2), XI-2 +

aesthetasc, XII-2 + aesthetasc, XIII—-1 + aesthetasc, XIV—2

+ aesthetasc, XV—2 + aesthetasc, XVI-2 + aesthetasc,

_XVII-2 + aesthetasc, XVIII-2 + aesthetasc, XIX—2 +

aesthetasc, XX-1 + aesthetasc, XXI-2 + aesthetasc,

XXII-1, XXIII-1, XXIV—XXVIII-12 + 2 aesthetascs

(XXIV-1 + 1, XXV-1 + 1 + aesthetasc, XX VI-XXVIII-8 +

aesthetasc). Left antennule: segments X and XI partly fused

near posterior margin; suture between segments XI and XII

visible on both surfaces, XII and XIII only on one surface,

XIII and XIV completely fused; fusion pattern and armature

as follows: I-V—10 + 2 aesthetascs (I-3 + aesthetasc, II-1,

III-2 + aesthetasc, IV-2, V—2), VI-2, VII-1 + aesthetasc,

VIII-1, [X—-X-3 + aesthetasc (IX-l1 + aesthetasc, X-—2),

XI-XIV-7 + 2 aesthetascs (XI-2 + aesthetasc, XII-2,

XIII-1, XI V—2 + aesthetasc), XV—2 + aesthetasc, XVI-2 +

aesthetasc, XVII—-2 + aesthetasc, XVIII-2 + aesthetasc,

XIX-2 + aesthetasc, XX-2 + aesthetasc, XXI-2 + aes-

thetasc, XXII-1, XXIII-1, XXIV-XXVIII-12 + 2 aes-

thetascs (XXIV-1 + 1, XXV-1 + 1 +. aesthetasc,

XXVI-XXVIII-8 + aesthetasc).

Antenna with same segmentation and setation as M. spe-

cies 1. Mandibular palp: endopod (Fig. 25B) rudimentary,

l-segmented, with 1 plumose seta; exopod with setation as in

M. species 1. Maxillule: praecoxal arthrite without elements;

coxal endite with short seta; coxal epipodite with 5 setae; no

basal seta; endopod represented by small, unarmed knob.

Maxilla and maxilliped as in M. species 1.

Legs 1 to 4 with same segmentation and setation as M. sp.

1. Leg 5 (Fig. 26H): coxae separate from intercoxal sclerite;

right basal seta thicker than left; endopod absent; right and

left exopods each 1-segmented, bulbous, with spiniform seta

terminally.

REMARKS. The fifth leg of this as yet undescribed species

resembles that of M. curvirostris but it can be distinguished

from the latter by the smaller body, the longer antennules,

and by differences in the mouthparts.

Genus Paraugaptilus Wolfenden, 1904

DIAGNOSIS (emended). Female. Body relatively large, mea-

suring about 3 mm in total length. Prosome: cephalosome

narrowed anteriorly, separate from or weakly fused with first

144

Fig. 26.

S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

Metacalanus sp. 1, female (A-F), male (G); Metacalanus sp. 2, female (H). A. Fourth endopod segment of maxilliped, innermost

seta indicated by arrowhead; B, Fifth endopod segment of maxilliped, innermost seta indicated by arrowhead; C, Sixth endopod segment of

maxilliped; D, Exopod of leg 1, anterior surface; E, Right leg 5, posterior surface; F, Left leg 5, anterior surface; G, Leg 5, anterior

surface; H, Leg 5, posterior surface. Scales in mm.

pedigerous somite; last pedigerous somite with short promi-

nence or curved process dorsally and weakly developed lobe

laterally on each side. Genital double-somite with pair of

gonopores located anteroventrally; copulatory pores asym-

metrically distributed posteroventrally, each copulatory duct

heavily chitinized; seminal receptacle relatively small. Caudal

rami symmetrical, longer than wide, with setae II and III

normally developed.

Antennule symmetrical or slightly asymmetrical in orna-

mentation of terminal segments (outer seta on segments

XXV and XXVI with thicker setules in one antennule than in

other) and in length, left slightly longer than right,

20-segmented; segments I to IV fused; segments XXIII-—

XXVIII fused; segments II, IV, VI, VIII-X, XII and XIII

lacking aesthetasc; segment XIII with 2 setae; compound

segment XX VI-XXVIII with 7 setae and 1 or 2 aesthetascs.

Antenna: first endopod segment without inner seta, second

segment bearing 1 seta medially, and 5 setae and vestigial seta

terminally; exopod indistinctly 6-segmented, sixth segment

rudimentary, unarmed. Mandibular gnathobase with tuft of

setules; 3 teeth on cutting edge, dorsalmost of which bifid at

tip. Mandibular palp: endopod absent; first exopod segment

bearing vestigial seta, outer seta on fifth segment vestigial.

Maxillule: praecoxal arthrite with 5 spines; coxal endite

bearing no seta; coxal epipodite with 8 setae; endopod

absent. Maxilla: first and second praecoxal endite bearing 1

seta and 1 rudimentary element, and 1 seta, respectively;

basal spine bipinnate; endopodal setae with triangular

PHYLOGENY OF ARIETELLID COPEPODS

spinules along inner margin. Maxilliped: setal formula of

endopod 1,4,4,3,3,4; setae a and b on sixth endopod segment

reduced; seta c heavily chitinized, terminal inner spinules

fused to seta to form serrate margin.

Third exopod segment of leg 1 with 2 outer spines. Leg 4

with minute inner coxal seta, in addition to basal seta. Leg 5

rudimentary, represented by a plate with proximal (basal)

seta and terminal or subterminal (endopod) seta.

Male. Body as in female, measuring around 3 mm in

length. Left antennule 19-segmented; only first segment

fringed with setules along posterior margin; segments IT and

III with seta; segment XIII with 2 setae; segment XXI and

XXII fused; compound segment XXIV—XXV with large

cuticular process; segment XX VI-XXVIII with 7 setae and

aesthetasc.

Antenna: second endopod segment relatively shorter than

in female, with 1-2 setae medially; exopod indistinctly 6- or

7-segmented, segment VIII with or without seta, terminal

compound segment (IX—X) completely or incompletely fused

with segment VIII, bulbous, unarmed. Mandibular palp: first

exopod segment with well-developed seta.

Leg 5: coxae fused with intercoxal sclerite; basis and coxa

separate in left leg and incompletely fused in right. Right leg:

endopod 1-segmented, rudimentary, unarmed; second exo-

pod segment expanded inwards, almost completely fused

with third to form compound segment, tapering distally,

carrying proximal seta and subterminal setule along outer

margin. Left leg: endopod 1-segmented, unarmed; exopod

3-segmented, last 2 segments almost fused, second exopod

segment swollen medially, third segment with 2 stout long,

outwardly-directed process terminally.

TYPE SPECIES. Paraugaptilus buchani Wolfenden, 1904

(monotypic).

OTHER SPECIES. P. similis A. Scott, 1909; P. meridionalis

Wolfenden, 1911; P. mozambicus Gaudy, 1965; P. archimedi

Gaudy, 1973; P. bermudensis Deevey, 1973; P. buchani sensu

Bradford, 1974.

REMARKS. In P. bermudensis sexual dimorphism is exhibited

in the mouthparts and leg 1 (Deevey, 1973): second endopo-

dal segment of antenna carrying | short seta in female and 1

long plus 1 short seta in male, at midlength of the segment;

relative lengths of endopod and exopod of antenna; anten-

nary exopodal segment VIII unarmed in female, but bearing

long seta in male; first exopodal segment of mandible

unarmed (vestigial seta overlooked by Deevey (1973)) in

female but with well-developed seta in male; endopod of leg 1

indistinctly 3-segmented in female but distinctly in male.

Except for leg 1 the sexual dimorphism in P. bermudensis is

also found in P. similis (present study).

Since the superfamily Arietelloidea Sars, 1902 generally

exhibits distinctly 3-segmented rami in legs 14 (Andronov,

1974; Park, 1986) and no other congeners show such fusion in

endopod of leg 1, the incomplete fusion of the endopodal

segments in the female seems to be autapomorphic in P.

bermudensis. P. buchani exhibits sexual dimorphism only in

the relative lengths of the antennary rami and in the setation

of the mandibular palp (Deevey, 1973; present study).

Brodsky (1950) mentioned, in his definition of Paraugapti-

lus, that the left antennules of females are possibly longer

than the right, but P. similis has antennules of nearly equal

length (Scott, 1909; present study).

145

ECOLOGICAL NOTE. Paraugaptilus is mainly distributed

within the upper 1000 m, in particular, between 500 and 1000

m depths (Deevey, 1973). The genus appears to be meso- and

bathypelagic.

Paraugaptilus similis A. Scott, 1909 (Figs 27-30)

MATERIAL EXAMINED. Q and GC’.

BODY LENGTH. 9 3.32 mm; 0 3.03 mm.

DESCRIPTION. Female. Cephalosome separate from first

pedigerous somite. Genital double-somite (Figs 27A-C,28A)

asymmetrical, wider than long, swollen anteriorly, widest at

level of paired gonopores; each gonopore (Fig. 28B) covered

by operculum as in Arietellus, anterior half opening; copula-

tory pores remarkably asymmetrical, right pore located medi-

ally on right ventral side, slit-like, approximately 43 wm in

length, left pore located ventromedially at about two-thirds

distance along double-somite, with round opening, about 27

jum in diameter; both right and left copulatory ducts heavily

chitinized; right duct shorter than left, widest near pore

opening, constricted medially; left duct thick, with small

subchamber medially (see Fig. 27B); seminal receptacles

relatively small, right round in shape, left smaller than right,

spindle-shaped.

Antennule (Fig. 27D) 20-segmented; seventh (X) to ninth

(XII) segments and 11th (XIV) and 12th (XV) segments only

partly fused near posterior margin; 20th (XXIII-XXV) and

21st (XXVI-XXVIII) incompletely fused with suture clearly

visible. Fusion pattern and armature as follows: I-IV—9 +

aesthetasc (I-3, II-2, III-2 + aesthetasc, IV-2), V—2 +

aesthetasc, VI-2, VII-2 + aesthetasc, VIII-2, IX-2, X-2,

XI-2 + aesthetasc, XII-2, XIII-2, XIV-2 + aesthetasc,

XV-2 + aesthetasc, XVI-2 + aesthetasc, XVII-2 + aes-

thetasc, X VIII-2 + aesthetasc, XIX—2 + aesthetasc, XX-2 +

aesthetasc, XXI-2 + aesthetasc, XXII-1, XXIII-XXVIII-12

+ 2 aesthetascs (right), 12 + 3 aesthetascs (left) (XXIIF-1,

XXIV-1 + 1, XXV-1 + 1 + aesthetasc, XXVI-XXVIII-7 +

1 (right) or 2 (left) aesthetascs). First (I-IV) to seventh

segments fringed with long setules along posterior margin.

Posterior setae on segments XXV and XXVI having thicker

setules in right antennule than in left.

Antenna: first endopod segment without inner mid-length

seta, second segment (Fig. 29B) about 1.8 times as long as

first segment, with 1 inner short seta, and 5 setae and vestigial

seta terminally; exopod (Fig. 29A) indistinctly 6-segmented,

sixth segment bulbous, unarmed; setal formula 0,1,1,1,1,0.

Mandibular palp (Fig. 29C): endopod absent; first exopod

segment carrying vestigial seta, fifth segment having 1 normal

and 1 reduced seta. Maxillule (Fig. 27E): praecoxal arthrite

with 5 spines, 2 of which serrate subterminally, ornamented

by minute spinules on both surfaces; coxal endite unarmed;

coxal epipodite with 8 setae; basal seta and endopod absent.

Maxilla: first praecoxal endite with 1 serrate seta and 1

vestigial element, second endite having single bipinnate seta

(Fig. 30A); basal spine (Fig. 29D) with 3 rows of spinules.

Maxilliped: fourth and fifth endopod segments (Fig. 27F)

each bearing unipinnate innermost seta, sixth segment (Fig.

27G) carrying reduced setae a and b, medium-length serrate

seta c whose tip chitinized, and elongate seta d with row of

sharp triangular spinules along inner margin.

Leg 1: third exopod segment with 2 outer spines; endopod

distinctly 3-segmented. Leg 4: vestigial coxal seta present at

inner angle. Leg 5 (Fig. 30B): coxae, intercoxal sclerite, basis

146

S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

‘kA

LSS

_ A AATATTTA TS

FSO) ae

0.1

Fig. 27. Paraugaptilus similis, female. A, Genital double-somite, right lateral view; B, Genital double-somite, left lateral view; C, Genital

double-somite, ventral view; D, Antennulary segments XXII-XXVIII; E, Praecoxal arthrite, coxal endite and inner margin of basis; F,

Fourth and fifth endopod segments of maxilliped, innermost seta indicated by arrowhead, mid-margin seta on fourth segment missing; G

Sixth endopod segment of maxilliped. Scales in mm.

PHYLOGENY OF ARIETELLID COPEPODS

Fig. 28. Paraugaptilus similis, female. SEM micrographs of genital double-somite. A, Genital double-somite, ventral view, copulatory pores

indicated by arrows, scale bar = 100 um; B, Left gonopore, scale bar = 20 um.

and endopod fused to form flattened plate; basal setae of

almost equal length; endopod represented by plumose seta;

exopod completely absent.

Male. Left antennule (Fig. 30C-E) 19-segmented; segments

IX to XV only partly fused near posterior margin; segments

XXI and XXII almost fused, but suture visible near anterior

margin; segments XXIV-XXV and XXVI-XXVIII incom-

pletely fused; fusion pattern and armature as follows: I-IV—7

+ 4 aesthetascs (I-3 + aesthetasc, II-1 + aesthetasc, III-1 +

aesthetasc, [V—2 + aesthetasc), V-2 + aesthetasc, VI-2 +

aesthetasc, VII-2 + aesthetasc, VIII—2 + aesthetasc, [IX—2 +

aesthetasc, X-1 + aesthetasc + process, XI-2 + aesthetasc,

XII-1 + aesthetasc + process, XIII-1 + aesthetasc + pro-

cess, XIV-—1 + aesthetasc + process, XV—1 + aesthetasc +

process, XVI-2 + aesthetasc, XVII-2 + aesthetasc, XVIII-2

+ aesthetasc, XIX—1 + aesthetasc + 2 processes, XX—1 +

aesthetasc + process, XXI-XXIII-1 + aesthetasc + 3 pro-

cesses (XXI-aesthetasc + 2 processes, XXII-process,

XXIII-1), XXIV-XXVIII-11 + 2 aesthetascs + process (1

seta missing in Fig. 30E) (XXIV-1 + 1 + process, XXV-1 +

1 + aesthetasc, XXVI-XXVIII-7 + aesthetasc). Only first

segment fringed by short setules along posterior margin.

Antenna: second endopod segment (Fig. 29G) approxi-

mately 1.3 times as long as first segment, with 1 long and 1

short seta medially; exopod (Fig. 29E,F) indistinctly

7-segmented, terminal compound segment bulbous (IX—X),

sixth (VIII) carrying long seta, seventh (IX—X) unarmed.

Mandibular palp (Fig. 29H): first exopod segment with long

seta.

Leg 5 (Fig. 30F): coxae and intercoxal sclerite almost

completely fused; coxa and basis incompletely fused in right

leg, but separate in left; right and left endopods consisting of

1 segment. Right exopod 2-segmented, ancestral second and

third segments almost completely fused, proximal segment

triangular, with short seta at outer angle, distal compound

segment lamellar, expanded proximally, tapering distally,

carrying short outer seta near base, triangular inner process

and 2 patches of setules medially. Left exopod indistinctly

3-segmented, first segment with short seta at outer angle,

second swollen inwards, bearing minute setule subterminally,

third segment incompletely fused with second segment, hav-

ing 2 processes, outer bifid at tip, and minute subterminal

outer setule.

REMARKS. The large process on segment XXIV of the left

antennule probably represents an extension of the cuticular

surface rather than a modified setation element. The anterior

subterminal process on the counterpart of the male left

antennule of Paraugaptiloides magnus is possibly homologous

to that of Paraugaptilus. The presence of 2 aesthetascs

located immediately adjacent to each other on the extreme

tip of the left antennule is interpreted here as an abnormality.

Paraugaptilus buchani Wolfenden, 1904 (Figs 31,32)

MATERIAL EXAMINED. 9 and CO.

BODY LENGTH. 9 3.14 mm; C 3.25 mm.

DESCRIPTION. Female. Cephalosome separate from first

pedigerous somite. Genital double-somite (Fig. 31A) similar

to that of P. similis, but relatively shorter, left copulatory

pore located near posterior margin. Female left antennule

(Fig. 32A) with same fusion pattern and armature as in

female P. similis except for following: segment XXIII incom-

148

S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

Fig. 29. Paraugaptilus similis, female (A-D), male (E-H). A, Antennary exopod; B, Second endopod segment of antenna; C, Mandibular

exopod; D, Basal spine of maxilla; E, Antennary exopod; F, Detail of antennary exopod segments IV to X; G, Second endopod segment of

antenna; H, Mandibular exopod. Scales in mm.

pletely fused with segments XXIV—XXV; segments XXV and

XXVI_ incompletely fused; left compound segment

XXVI-XXVIII with 7 setae and aesthetasc.

Antenna: second endopod segment about 1.9 times as long

as first, with 1 minute inner seta at mid-length and 5 setae and

1 vestigial seta terminally, as in P. similis; exopod similar in

segmentation and setation to that of female P. similis.

Mandibular palp: first exopod segment with vestigial seta

(Fig. 32B) as in female P. similis. Maxilliped: sixth endopod

segment (Fig. 32E) similar to that of P. similis, but seta c with

terminal spinules incompletely fused to seta.

Male. Left antennule (Fig. 32F) with same fusion pattern

and armature as in P. similis except for following: seta on

segment XXII not modified into process; process on segment

XXIV-XXV not so developed as in male P. similis, not

reaching beyond end of antennule, directed straight for-

wards. Antenna similar in segmentation and setation to that

of female; second endopod segment ca. 1.4 times as long as

first. Mandibular palp: first exopod segment with well-

developed seta (Fig. 32G). Maxillule: praecoxal arthrite (Fig.

32C) with 5 spines; tubular gland opening on inner surface.

Leg 5: both coxae and intercoxal sclerite completely fused

as in male P. similis; coxa and basis separate in left leg and

incompletely fused in right (almost completely fused on

PHYLOGENY OF ARIETELLID COPEPODS

Fig. 30. Paraugaptilus similis, female (A,B), male (C-F). A, First and second praecoxal endites of maxilla; B, Leg 5, anterior surface; C,

Left antennulary segments I to XVI; D, Left antennulary segments XVII to XXVIII; E, Left antennulary segments XXIV to XXVIII; F,

Leg 5, anterior surface, minute seta indicated by arrowhead. Scales in mm.

149

S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

Fig. 31. Paraugaptilus buchani, female. SEM micrographs of genital double-somite. A, Genital double-somite, copulatory pores arrowed,

scale bar = 100 .m; B, Copulatory pores, scale bar = 50 ym; C, Right gonopore, scale bar = 20 ym; D, Left gonopore, scale bar = 20

wm.

PHYLOGENY OF ARIETELLID COPEPODS 151

Fig. 32. Paraugaptilus buchani, female (A-E), male (F-J). A, Antennulary segments XXII to XXVIII; B, Mandibular exopod; C, Praecoxal

arthrite and coxal endite of maxillule; D, Fourth to sixth endopod segments of maxilliped, innermost seta on fourth and fifth segments

indicated by arrowheads; E, Sixth endopod segment of maxilliped; F, Antennulary segments XIX to XXVIII; G, Mandibular exopod; H,

Second exopod segment of right leg 5; I, Inner medial process on second exopod segment of right leg 5; J, Outer margin of second exopod

segment of right leg 5. Scales in mm.

152

posterior surface); both endopods 1-segmented, lobate. Right

exopod (Fig. 32H-J): second and third segments almost

completely fused to form lamelliform compound segment,

tapering distally; inner medial triangular process with 2

minute spinules (Fig. 321) at tip; 1 subterminal outer and 1

terminal setule present (Fig. 32J); muscles between second

and third segments present, but less developed than in

Paraugaptiloides. Left exopod similar to that of male P.

similis.

REMARKS. Deevey (1973) first discovered sexual dimor-

phism in the mandibular palp of this species, but overlooked

the vestigial seta on the first exopodal segment of the female.

P. buchani shows no sexual dimorphism in setation of the

antennary endopod and exopod, unlike P. bermudensis

(Deevey, 1973) and P. similis (A. Scott, 1909; present study).

Unfortunately the only female of P. buchani lacked the

terminal segments of the right antennule. The posterior setae

on segments XXV and XXVI of the left antennule are

ornamented with thick setules as in the right antennule of

female P. similis. In P. buchani the asymmetrical pattern in

antennulary armature elements may be different from that of

P. similis.

Genus Scutogerulus Bradford, 1969

DIAGNOSIS (emended). Only female known. Body relatively

large, more than 3 mm long. Cephalosome separate from first

pedigerous somite; urosome about one-third as long as

prosome. Genital double-somite as long as wide; gonopore

and copulatory pore sharing common slit-like aperture,

gonopore located anteriorly, copulatory pore at innermost

corner of the slit; copulatory duct swollen anteriorly; seminal

receptacle relatively small and simple in shape. Caudal rami

slightly asymmetrical, left caudal ramus longer than right,

longer than wide, with setae II and III relatively long.

Antennules symmetrical, reaching almost to end of

prosome, 22-segmented; posterior margin of proximal seg-

ments bearing long setules from segment I to XIII; segment

III separate from IV; segment IV without aesthetasc; seg-

ment XIII with 2 setae; segment XXIII separate from XXIV.

Antenna: first endopod segment without inner seta; second

endopod segment with 3 medial and 5 terminal setae; exopod

indistinctly 8-segmented. Mandibular palp: endopod rudi-

mentary, 1l-segmented, with 2 setae; seta on first exopod

segment not reduced; outer seta on fifth segment relatively

long. Maxillule: praecoxal arthrite with 4 finely serrate spines

and 1 process; coxal epipodite with 6 setae; coxal endite

carrying 1 long seta; endopod having single seta. Maxilla: first

praecoxal endite with 1 relatively well developed seta and 1

vestigial element; second praecoxal endite with 1 seta; basal

spine with 3 rows of minute spinules; setae on endopod with

row of triangular spinules. Setal formula of endopod of

maxilliped: 1,4,4,3,3,4; setae a and b on sixth endopod

segment vestigial.

Third exopod segment of leg 1 with outer medial tuft of

short setules and subterminal outer spine. Leg 5 biramous,

carrying 1-segmented rudimentary endopod with 1 terminal

seta and 2-segmented exopod with 1 outer spine on first

segment and 2 terminal setae on second segment.

TYPE SPECIES. Scutogerulus pelophilus

(monotypic).

Bradford, 1969

S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

REMARKS. The new genus Campaneria is established for the

paratypic male of S. pelophilus.

ECOLOGICAL NOTE. Bradford (1969) suggested that S. pelo-

philus is a deep-sea hyperbenthic species. However, Cam-

paner (1984) considered that it was less associated with the

bottom than members of his second group, namely, Parami-

sophria, Rhapidophorus and some species of Metacalanus,

since S. pelophilus has well-developed antennules and anten-

nae for swimming. The presence of long caudal setae also

supports Campaner’s (1984) inference.

Scutogerulus pelophilus Bradford, 1969 (Figs 33,34)

MATERIAL EXAMINED. Q, Paratype, New Zealand Oceano-

graphic Institute, p-121.

BODY LENGTH. 3.6 mm (after Bradford, 1969).

DESCRIPTION. Female. Cephalosome separate from first

pedigerous somite. Urosome (Fig. 33A) slender. Genital

double-somite (Fig. 33B,C) as long as wide; paired gonop-

ores and copulatory pores symmetrically arranged; gonopore

sharing common slit-like aperture with copulatory pore;

gonopore located anteriorly in slit, genital operculum accom-

panied by muscles; copulatory pore small, located at inner-

most corner of slit; copulatory duct swollen anteriorly,

relatively short; seminal receptacle simple in shape, pea-like;

receptacle duct short, opening beneath gonopore. Left caudal

ramus slightly longer than right, with seta V longer than

urosome (Fig. 33A).

Antennule (Fig. 33D-F): eighth (X) to 10th (XII) segments

separate; 12th (XIV) and 13th (XV) segments partly fused

(Fig. 33D). Fusion pattern and armature elements as follows:

I-III-7 + 2 aesthetascs, IV-2, V—2 + aesthetasc, VI-2 +

(small) aesthetasc, VII-2 + aesthetasc, VIII-2 + (small)

aesthetasc, IX-2 + aesthetasc, X-2 + (small) aesthetasc,

XI-2 + aesthetasc, XII-2 + (small) aesthetasc, XIII-2 +

aesthetasc, XIV-—2 + aesthetasc, XV-XVI4 + 2 aesthetasc,

XVII-2 + aesthetasc, XVIII-2 + aesthetasc, XIX—2 +

aesthetasc, XX-2 + aesthetasc, XXI-2 + aesthetasc,

XXII-1, XXIII-1, XXIV-XXV-4 + aesthetasc, XXVI-

XXVIII-8 + aesthetasc. First to 11th (XIII) segments fringed

with long setules along posterior margin.

Antenna: first endopod segment without inner seta, second

segment carrying 3 inner setae and 5 terminal setae; exopod

(Fig. 33G) indistinctly 8-segmented, setal formula

0,1,1,1,1,1,0,3. Mandibular gnathobase missing, probably

lost during dissection. Mandibular palp (Fig. 33H): endopod

rudimentary, 1-segmented, bearing 2 setae of unequal

lengths; seta on first exopod segment not reduced, fifth

segment with 2 setae, one of which shorter but not reduced.

Maxillule: praecoxal arthrite (Fig. 34A) with 4 spinulose

spines and 1 process along inner margin and row of long

setules on surface; coxal endite with well-developed spinulose

seta; coxal epipodite with 6 setae (only 4 setae and 2 scars

remaining on slide); basal seta short, endopod rudimentary,

1-segmented, with 1 short seta terminally (Fig. 34B). Maxilla

(Fig. 34C): first praecoxal endite with spinulose seta and 1

vestigial element, second endite with bilaterally spinulose

seta. Maxilliped: innermost seta on fourth and fifth endopod

segments (Fig. 34E, indicated by arrowhead) not reduced;

sixth endopod segment (Fig. 34F) bearing stout, elongate

setae c and d with row of triangular spinules and reduced

setae a and b.

PHYLOGENY OF ARIETELLID COPEPODS 153

Fig. 33. Scutogerulus pelophilus, female (paratype). A, Urosome, ventral view; B, Genital double-somite, ventral view; C, Genital

double-somite, lateral view; D, Antennulary segments IX to XIV, armature omitted; E, Antennulary segments VI and VII, note that

aesthetasc on each segment differs in size; F, Antennulary segments XXI to XXVIII; G, Antennary exopod; H, Mandibular endopod and

exopod. Scales in mm.

154 S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

Fig. 34. Scutogerulus pelophilus, female (paratype). A, Praecoxal arthrite and coxal endite of maxillule; B, Maxillulary endopod, basal seta

indicated by arrowhead; C, First and second praecoxal endites of maxilla; D, Basal spine of maxilla; E, Fourth and fifth endopod segments

of maxilliped, innermost setae indicated by arrowheads; F, Sixth endopod segment of maxilliped; G, Exopod of leg 1, posterior surface.

Scales in mm.

PHYLOGENY OF ARIETELLID COPEPODS

Leg 1 (Fig. 34G): first exopod segment missing element on

outer corner, third segment with tuft of minute setules

medially and spinulose spine subterminally along outer mar-

gin. Leg 5 of paratype missing.

Genus Sarsarietellus Campaner, 1984

DIAGNOsIs (emended). Only female known. Body relatively

large, 3 to 5 mm in length. Prosome oblong in dorsal view;

cephalosome separate from first pedigerous somite; ventro-

lateral corner of last prosomal somite slightly produced;

urosome about one-third as long as prosome. Genital double-

somite longer than wide, produced ventrally; pair of gonop-

ores located anteroventrally, single copulatory pore

posteromedially; paired copulatory ducts medially fused to

form common duct, heavily chitinized; seminal receptacle

elongate, slender, with terminal part bulbous. Caudal rami

symmetrical, longer than wide, with setae II and III well

developed.

Antennules symmetrical, reaching to end of prosome,

22-segmented; posterior margin of ancestral segments I to X

fringed with long setules; segment III separate from IV;

segment IV without aesthetasc; segment XIII with 2 setae;

segment XXIII separate from XXIV. Antenna: second endo-

pod segment with 5 setae and 1 vestigial seta terminally;

exopod indistinctly 8-segmented. Mandibular gnathobase

lacking tuft of setules; 3 teeth on cutting edge, dorsalmost of

which bifid at tip. Mandibular palp: endopod rudimentary, 1-

segmented endopod with 2 setae; seta on first exopod seg-

ment not reduced; outer seta on fifth exopod segment

relatively long. Maxillule: praecoxal arthrite with 6 elements

(5 spines and 1 process); coxal epipodite with 8 setae; coxal

endite with 1 long seta; endopod bearing 2 setae and 1

vestigial seta. Maxilla: first praecoxal endite with 2 well-

developed setae; basal spine with 2 rows of long spinules.

Setal formula of endopod segments of maxilliped: 1,4,4,3,3,4;

seta a on sixth endopod segment vestigial, seta b relatively

long.

Third exopod segment of leg 1 with 2 outer spines. Leg 5:

coxa and intercoxal sclerite separate; basis fused to endopod.

Endopod represented by process with 2 terminal and 2 inner

setae. Exopod composed of 3 almost fused segments, bearing

3 outer spines and 2 terminal spines of unequal lengths.

TYPE SPECIES. Scottula abyssalis Sars, 1905.

OTHER SPECIES. Sarsarietellus natalis Heinrich, 1993.

REMARKS. Sars (1905) assigned this species to the genus

Scottula Sars, 1902. Scottula was synonymized with the genus

Metacalanus Cleve, 1901 by Campaner (1984), but he pointed

out that Scottula abyssalis was not congeneric with Metacala-

nus, and established Sarsarietellus to accommodate it. A

second species of Sarsarietellus, S. natalis, has been recently

described from the near-bottom (1083-1090 m depth) in the

southwestern Indian Ocean (Heinrich, 1993). S. natalis

exhibits a few more apomorphic characters than S. abyssalis:

(1) asymmetry in the genital double-somite; (2) reduction of

the elements on the exopod of the fifth leg.

ECOLOGICAL NOTE. Campaner (1984) suggested that the

genus is only loosely associated with the deep-sea near-

bottom as is Scutogerulus. The recent discovery of a second

congener from the near-bottom supports his opinion.

155

Sarsarietellus abyssalis (Sars, 1905) (Figs 35,36)

MATERIAL EXAMINED. 9, Holotype, Zoological Museum,

University of Oslo, Catalog No. F5447-5448.

BODY LENGTH. 3 mm (after Sars, 1925).

DESCRIPTION. Female. Genital double-somite (Fig. 35A,B)

longer than wide; its posterior end damaged, but single

copulatory pore possibly present posteroventrally (fragment

of copulatory pore still remained on slide); anterior paired

gonopores located ventro-laterally (since the specimen was

dried up, the urosome was so depressed t!.at the internal

structures have become artificially asymmetrical); copulatory

duct heavily chitinized, divergent anteriorly, each connecting

with elongate seminal receptacle (Fig. 35B) which curved

anteriorly and reaching to half length of somite with

expanded bulbous part terminally.

Antennule (Fig. 36A) 22-segmented; suture between seg-

ments XXIV—XXVI visible. Fusion pattern and armature as

follows: I-III-7 + aesthetasc, [V-2, V-2 + aesthetasc, VI-2

+ aesthetasc, VII—2 + aesthetasc, VIII-2 + aesthetasc, [X-2

+ aesthetasc, X-2 + aesthetasc, XI-2 + aesthetasc, XII-2 +

aesthetasc, XIII-2 + aesthetasc, XIV-2 + aesthetasc, XV-2

+ aesthetasc, XVI-2 + aesthetasc, XVII-2 + aesthetasc,

XVIII-2 + aesthetasc, XIX-2 + aesthetasc, XX-2 + aes-

thetasc, XXI-2 + aesthetasc, XXII-1, XXIII-1,

XXIV-XXV-4 + aesthetasc, XX VI-XXVIII-8 + aesthetasc.

First to eighth (X) segments fringed with row of setules

posteriorly.

Antennary endopod: first segment without inner seta;

second segment (Fig. 36B) with 3 setae of unequal lengths

medially, and 5 setae and 1 vestigial seta terminally. Anten-

nary exopod (Fig. 36C) indistinctly 8-segmented, first to fifth

segments almost fused or incompletely fused, setal formula as

follows: 0,1,1,1,1,1,0,3. Mandibular gnathobase with 3 stout

teeth, dorsalmost of which bifid at tip, lacking medial tuft of

setules as found in Crassarietellus sp.; basis fringed by row of

long setules along inner margin, and not furnished with

minute spinules as in male of Crassarietellus sp. Mandibular

palp (Fig. 36D): endopod rudimentary, 1-segmented, with 2

setae of unequal lengths; exopod indistinctly 5-segmented,

seta on first segment not reduced, outer seta on fifth segment

relatively long.

Maxillule (Fig. 36E) praecoxal arthrite with 5 naked spines

and 1 process; coxal endite carrying long serrate seta; coxal

epipodite with 8 plumose setae; second basal endite bearing 1

vestigial seta; endopod bulbous, l-segmented, bearing 3

setae, one of which rudimentary. Maxilla: first praecoxal

endite (Fig. 36F) with 2 spinulose setae and vestigial element;

basal spine (Fig. 36G) stout, bearing 2 rows of long spinules.

Maxilliped: fourth endopod segment (Fig. 35C) with rela-

tively developed spinulose innermost seta, fifth segment (Fig.

36D) also having spinulose innermost seta, but much shorter

and thinner than on fourth segment; sixth endopod segment

(Fig. 36E) with seta a reduced, seta b over half length of

medial-length seta c, medium-length spinulose seta c, spinu-

lose seta d elongate.

Leg 4 without inner coxal seta. Leg 5 (Fig. 36H): intercoxal

sclerite more or less fused; endopod almost fused with basis,

medial suture visible; exopod separate from basis, indistinctly

3-segmented, sutures between segments visible, terminal

outer spine almost fused with segment.

REMARKS. Sars (1924, 1925) overlooked the vestigial seta on

156

S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

Fig. 35. Sarsarietellus abyssalis, female (holotype). A, Genital double-somite, ventral view, part around copulatory pore missing; B, Internal

structure of right genital system; C, Fourth endopod segment of maxilliped, innermost seta indicated by arrowhead; D, Fifth endopod

segment of maxilliped, innermost seta indicated by arrowhead; E, Sixth endopod segment of maxilliped. Scales in mm.

the second endopodal segment of the antenna, the rudimen-

tary 1-segmented mandibular endopod with 2 setae, and the

rudimentary seta on the second basal endite of the maxillule.

The terminal segments of the female antennule were

re-examined in detail, revealing that there were several

misinterpretations of the segmental fusion pattern and of the

setation pattern in Sars’ (1924, 1925) descriptions.

Genus Pilarella Alvarez, 1985

DIAGNOSIS (emended). Only female known. Body relatively

small, 1.5 to 1.7 mm in length. Prosome oblong in dorsal

view; cephalosome separate from first pedigerous somite;

ventrolateral corner of last prosome somite pointed; urosome

nearly half as long as prosome. Genital double-somite slightly

PHYLOGENY OF ARIETELLID COPEPODS

157

Fig. 36. Sarsarietellus abyssalis, female (holotype). A, Antennulary segments XXI to XXVIII; B, Terminal part of second endopod segment

of antenna, vestigial innermost seta indicated by arrowhead; C, Antennary exopod; D, Mandibular endopod and exopod; E, Praecoxal

arthrite, coxal endite, basis and endopod of maxillule, vestigial basal seta indicated by arrowhead; F, First praecoxal endite of maxilla; G,

Basal spine of maxilla; H, Leg 5, posterior surface. Scales in mm.

wider than long; entire reproductive system paired, sym-

metrical; large circular gonopore and small copulatory pore

located at outer and inner ends of slit-like aperture, respec-

tively; copulatory duct short, simple; seminal receptacle

relatively small, located medial to gonopore. Caudal rami

slightly asymmetrical, with right ramus narrower and just

shorter than left, with setae II and III relatively long.

Antennules asymmetrical, left longer than right and reach-

ing to end of caudal rami; antennules 21-segmented on both

sides; posterior proximal margin lacking long setules; seg-

ments I to IV fused, segments IX to XII partially fused;

segments XXIV to XXVIII fused into compound apical

segment. Antenna: first endopod segment with 1 mid-margin

inner seta, second with 3 setae at midlength and 5 setae

terminally; exopod indistinctly 7-segmented. Mandibular

gnathobase lacking tuft of setules; 4 teeth on cutting edge,

158

dorsalmost of which tricuspid; endopod rudimentary,

l-segmented with 2 setae; seta on first exopod segment not

reduced; outer seta on fifth segment relatively long. Maxil-

lule; praecoxal arthrite with 6 elements (5 setae and 1

process); coxal epipodite with 5 setae; coxal endite with 1

long seta; basal seta absent; endopod bearing 2 setae. Max-

illa: first praecoxal endite with 2 setae and vestigial element,

second praecoxal endite with 2 setae; basal spine with 2 rows

of spinules. Setal formula of endopod segment of maxilliped

1,4,4,4,3,3,4; setae a and b on sixth endopod segment rela-

tively well developed.

Leg 1 with 1 outer spine on third exopod segment. Leg 4

with inner seta on coxa. Leg 5: coxae separate from reduced

intercoxal sclerite; endopod represented by 1 seta; exopod

and basis separate. Exopod 1-segmented bearing 1 short

spine on outer margin and 1 short and 1 long spine terminally.

TYPE SPECIES. Pilarella longicornis Alvarez, 1985 (mono-

typic).

REMARKS. As Alvarez (1985) has already pointed out, the

genus Pilarella is very similar to Metacalanus, but can be

distinguished from the latter in the structures of antennules,

maxillule and caudal rami. The present study revealed that

the genital double-somite of Pilarella resembles that of Scu-

togerulus. A short supplementary description follows, provid-

ing details of setation and genital structure that were not

apparent in the original description (Alvarez, 1985).

ECOLOGICAL NOTES. The species was collected from near-

bottom at a depth of 135 m (Alvarez, 1985), and is, presum-

ably, a shallow-water hyperbenthic species.

Pilarella longicornis Alvarez, 1985 (Fig. 37)

MATERIAL EXAMINED. 39 9, paratypes, Copepod collection

of Departmenta de Zoologia, Instituto de Biociéncias, Uni-

versidade de Sao Paulo, Brasil, No. 186.

BODY LENGTH. 1.53 to 1.73 mm (after Alvarez, 1985).

DESCRIPTION. Genital double-somite (Fig. 37A) wider than

long; genital system symmetrical; genital aperture slit-like,

located just posterior to mid-length; large circular gonopores

present at outermost extremity of genital aperture and small

copulatory pore at innermost extremity; copulatory and

receptacle ducts short; seminal receptacle relatively small,

located medial to gonopore. Caudal rami slightly asymmetri-

cal, with right ramus narrower and just shorter than left, with

setae II and III relatively long.

Antennules (see Fig. 39 ) asymmetrical, left longer than

right and reaching to end of caudal rami; both antennules

21-segmented; posterior proximal margin lacking long set-

ules. Fusion pattern and armature as follows: I-IV-9 + 2

aesthetascs, V—2 + aesthetasc, VI-2, VII-2 + aesthetasc,

VIII-2 + aesthetasc, [IX—2 + aesthetasc, X—2 + aesthetasc,

XI-2 + aesthetasc, XII-2 + aesthetasc, XIII-2 + aesthetasc,

XIV-2 + aesthetasc, XV-—2 + aesthetasc, XVI-2 + aes-

thetasc, XVII-2 + aesthetasc, XVIII-2 + aesthetasc, XIX—2

+ aesthetasc, XX-2 + aesthetasc, XXI-2 + aesthetasc,

XXII-1, XXIII-1, XXIV—XXVIII-12 + 2 aesthetascs.

Antenna: second endopod segment (Fig. 37B) with 3 setae

of unequal lengths at midlength and 5 setae terminally;

exopod indistinctly 7-segmented. Maxillule: praecoxal arth-

rite (Fig. 37C) with 6 elements (5 setae and 1 process); coxal

epipodite with 5 setae; endopod bearing 2 setae of unequal

S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

lengths (Fig. 37D). Maxilla: first praecoxal endite with 2

setae and vestigial element (Fig. 37E), second praecoxal

endite with 2 spinulose setae; basal spine with 2 rows of

spinules. Maxilliped: setae a and b on sixth endopod segment

(Fig. 37F) relatively well developed.

Leg 1 with 1 outer spine on third exopod segment. Leg 4

with short inner seta on coxa. Leg 5: coxae separate from

small intercoxal sclerite; endopod represented by 1 relatively

long seta; exopod and basis separate; exopod 1-segmented

bearing 1 short spine on outer middle margin and 1 short

outer and 1 long inner spine terminally.

DISCUSSION

Ancestral states and character transformation

All genera of the family Arietellidae except Rhapidophorus

are described in detail and their characters are discussed prior

to analysis of the phylogenetic relationships between the

genera. Within a single genus various states can be observed

in appendage segmentation and setation patterns. For

example, Metacalanus species show a variety of character

states in the antennules (Fig. 22) and fifth legs (Fig.

26E,F,H). In such a case, the most plesiomorphic state is

selected as the ancestral state for the genus, using the

principle of deduction of ancestral states proposed by Huys &

Boxshall (1991). Fig. 22 schematically depicts the segmenta-

tion and setation of right and left female antennules of 2 new

species of Metacalanus collected from Okinawa, South Japan.

Asymmetry in segmentation and setation is exhibited in both

species. The fewest segmental fusions and the greatest num-

ber of armature elements on each segment are combined

from both antennules of these two species in order to arrive at

a hypothetical ancestral condition. The hypothetical anten-

nule of ancestral Metacalanus so constructed is used for

comparison with antennules of other arietellid genera.

In the antenna and mandibular palp of Arietellus and

Paraugaptilus, which show sexual dimorphism, the more

plesiomorphic state from either sex is selected as the generic

character state. By reference to the ancestral character states

for Calanoida (Huys & Boxshall, 1991) the evolutionary

trends within the family are traced.

1. Body plan. The most primitive condition in the family

can be seen in Crassarietellus and Sarsarietellus. The body is

symmetrical with complete separation between the cephalo-

some and the first pedigerous somite; there is no projection at

the tip of the cephalosome, no strong dorso- and ventrolat-

eral processes on the last prosomal somite, and no specializa-

tion of the caudal ramus.

Asymmetry in the body, except for female genital double-

somites, can be seen in the ventrolateral processes on the last

prosomal somite in Arietellus giesbrechti (Sars, 1924, 1925),

A. mohri (Bjornberg, 1975), and A. sp.; in the ventrolateral

corners of the second and third pedigerous somites in Parami-

sophria giselae (Campaner, 1977); and in the prosome of

Paramisophria platysoma (Ohtsuka & Mitsuzumi, 1990).

These are more apomorphic states compared with congeners

which have symmetrical counterparts. The asymmetrical

prosome of P. platysoma appears to result from its specialized

adaptation to the hyperbenthic zone (Ohtsuka & Mitsuzumi,

1990).

The cephalosome is separate from the first pedigerous

PHYLOGENY OF ARIETELLID COPEPODS 159

Fig. 37. Pilarella longicornis, female (paratype). A, Genital double-somite, ventral view; B, Apical endopod segment of antenna; C,

Praecoxal arthrite of maxillule; D, Maxillulary endopod; E, Praecoxal endites of maxilla; F, Tip of endopod of maxilliped showing setae a

and b. Scales in mm.

somite in almost all arietellids. Re-examination of those taxa arietellids the fourth and fifth pedigerous somites are invari-

in which the cephalosome and the first pedigerous somite ably fused, with or without a suture.

were previously reported to be fused (for example, Paraugap- Within the genus Arietellus, A. setosus has a well-

tilus magnus), has revealed that these somites are clearly developed cephalic projection, a pair of strong ventrolateral

separate. In Metacalanus species 1 the cephalosome is weakly processes on the last prosomal somite and a posteriorly

fused with the first pedigerous somite ventrolaterally. In all swollen caudal ramus with remarkably elongate setae. In

160

contrast A. simplex lacks all these characteristics (see Sars,

1924, plates 118, 120). Paramisophria species typically have a

pair of pointed dorsolateral and rounded or prominent vent-

rolateral processes on the last prosomal somite (e.g., Sars,

1903; Fosshagen, 1968; Campaner, 1977; McKinnon & Kim-

merer, 1985; Ohtsuka, 1985; Ohtuska & Mitsuzumi, 1990).

Although some cave-living species of Paramisophria lack

such processes (Ohtsuka et al., 1993a), there is a cave-living

Paramisophria with processes in Bermuda (Fosshagen, per-

sonal communication). The genera Paraugaptilus and

Paraugaptiloides consistently exhibit a pair of dorsolateral

processes on the last prosomal somite and no cephalic projec-

tion (Sars, 1924; Gaudy, 1965; Deevey, 1973; Bradford,

1974). Sarsarietellus has weakly developed dorsolateral

and/or ventrolateral processes on the last prosomal somite

(Sars, 1924, 1925; Heinrich, 1993). Crassarietellus, Metacala-

nus, Scutogerulus, Pilarella and, possibly, Campaneria lack

dorsolateral processes on the last prosomal somite and a

cephalic projection (Bradford, 1969; Alvarez, 1985; present

study).

2. Genital double-somite. The present study has revealed

an amazing variety of genital systems of arietellid females.

The hypothetical ancestral calanoid proposed by Huys &

Eoxshall (1991) was characterized by paired genital apertures

located about in the middle of the genital double-somite. This

basic condition is displayed by the genera Crassarietellus (Figs

1D,E,2A), Scutogerulus (Fig. 33B,C) and Pilarella (Fig.

37A). The paired gonopores are ventrolaterally located at

about the midlength of the genital double-somite, and the

paired copulatory pores are situated either posterior to the

gonopores or at the midlength of the somite. Scutogerulus

exhibits the most plesiomorphic state, similar to that of the

primitive family Pseudocyclopidae (see Huys & Boxshall,

1991, Fig. 2.2.32): the gonopore and the copulatory pore

share a common opening, with the copulatory pore located

on the innermost part of the common opening; the gonopore

is located in the outer part of the common opening. Although

Huys & Boxshall (1991) did not mention the location of

paired seminal receptacles of the ancestor, it is likely that

they lie ventrally just beneath the gonopores as proposed for

the ancestor of the Cyclopoida (see Huys & Boxshall, 1991,

Lene, DaSeSi7)))-

Fig. 38 schematically depicts possible evolutionary trends

in structure of the female genital system in the Arietellidae,

based on the relative positions of gonopores and copulatory

pores. Five major trends are recognizable: (A) fusion of

copulatory pores to form a single common pore and antero-

lateral migration of gonopores; (B) posterior migration of

both gonopores and copulatory pores; (C) anterolateral

migration of gonopores, and asymmetrical arrangement and

enlargement of copulatory pores; (D) lateral migration of

both gonopores and copulatory pores, and copulatory pore

covered by ventral flap; (E) lateral migration of both gono-

pores and copulatory pores, copulatory pore uncovered. The

first three trends (A-C) are accompanied by the formation of

a pair of genital opercula, each of which closes off a gonopore

and opens anteriorly with a posterior hinge. The gonopore is

separate from the copulatory pore in all except the last trend

(E). The first evolutionary trend (A) is exhibited in Parami-

sophria, Arietellus and Sarsarietellus. The copulatory ducts

are heavily chitinized in Arietellus and Sarsarietellus (see Figs

13B,16A-C) but not so in Paramisophria (Figs 19A,20A). In

addition, each copulatory duct is connected to a medial part

of the seminal receptacle, but not so anteriorly as in Arietellus

S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

and Sarsarietellus. Even within the genus Paramisophria, a

remarkable trend is exhibited. In P. japonica and P. reducta,

the copulatory pore is located ventro-medially, whereas in P.

platysoma, P. itoi and P. cluthae the pore is present on the

left side of the genital double-somite (Ohtsuka & Mitsuzumi,

1990; Huys & Boxshall, 199i; Ohtsuka et al., 1991, 1993b).

Alternatively, the copulatory pore can be located on the right

side as in P. giselae. These asymmetrical species are thought

to be more derived than P. japonica.

In Arietellus the genital system is essentially the same as in

Paramisophria, but may be relatively more apomorphic in

having: (1) copulatory ducts much more heavily chitinized;

and (2) enlargement of the copulatory pore. In Sarsarietellus

the systems are basically similar to those of both Parami-

sophria and Arietellus, but are more closely related to Arietel-

lus in having the two previously mentioned apomorphic

states.

The genus Metacalanus exhibits the second trend (B).

Primitively, M. species 2, M. inaequicornis (Campaner,

1984), M. acutioperculum (Ohtsuka, 1984), M. curvirostris

(Ohtsuka, 1985) and, possibly, M. aurivilli display paired

gonopores and copulatory pores which are located along the

posterior margin of the genital double-somite. The gonopores

are relatively large. The copulatory pore is clearly separate

from the gonopore (see Figs 23A,24A), and is located near

the anterior inner corner of the gonopore. The seminal

receptacles are located ventrally at almost the same level as

the gonopore, and each is connected via a short, chitinized

copulatory duct. M. species 1 shows a further derived state

since it completely lacks the genital system on the left side.

The right genital structure of this species is quite similar to

that of the right side of other Metacalanus species, but is

bounded by a chitinized flap along the outer lateral margin

and the copulatory pore is slightly oblong in shape compared

with the rounded pore of the other congeners (see Figs

23,24).

The third trend (C) is exhibited by the genus Paraugaptilus.

The gonopores are almost symmetrically sited anteriorly (see

Figs 28A, 31A) while the copulatory pores are extremely

asymmetrical (see Fig. 31B). The right copulatory pore is

slit-like and situated in a large circular ventral depression; the

left pore is a large pore located posterior to the right. The left

copulatory duct is much longer than the right, although both

ducts are heavily chitinized. The seminal receptacles are

relatively small, bulbous, and located just posterior to the

gonopore; the right is better developed than the left (Fig.

27A,B). However, both genital systems are probably func-

tional because of the presence of well-developed muscles

which provide an opening-closing mechanism for the genital

operculum on both sides. In the recently established calanoid

family Hyperbionycidae (Ohtsuka et al., 1993b), only the left

genital system is functional; the right side lacks musculature

around the gonopore and is probably no longer functional.

Only two species of Paraugaptilus were available for the

present study but Gaudy’s (1965) and Deevey’s (1973) illus-

trations of the ventral surfaces of the female genital double-

somites of P. mozambicus and P. bermudensis suggest that

these species exhibit the same genital systems.

The fourth (D) and fifth trends (E) are displayed by

Crassarietellus, and by Scutogerulus and Pilarella, respec-

tively. Both trends show primitive states of the female genital

system in the presence of paired and symmetrically arranged

gonopores, copulatory pores and seminal receptacles. How-

ever, both trends exhibit different variations of the plesio-

PHYLOGENY OF ARIETELLID COPEPODS

way Parrot,

j ~

Oe ang aes

Ancestor

161

“OTN

} seminal receptacle

====> copulatory duct

receptacle duct

Fig. 38. Evolutionary trends in the structures of the female genital systems of the arietellid genera. A, Fusion of copulatory pores to form

single pore, and anterolateral migration of both gonopores; B, Posterior migration of both gonopores and copulatory pores, and separation

of copulatory pore from gonopore; C, Anterolateral migration of gonopores, and separation of copulatory pore from gonopore and their

asymmetrical arrangement and enlargement; D, Lateral migration of both gonopores and copulatory pores, and separation of copulatory

pore from gonopore; E, Lateral migration of both gonopores and copulatory pores. Pg: Paramisophria giselae; Pj: Paramisophria japonica;

Pe: Paramisophria cluthae; Sa: Sarsarietellus abyssalis; Ap: Arietellus plumifer; M1: Metacalanus species 1; M2: Metacalanus sp. 2; Ps:

Paraugaptilus similis; Ch: Crassarietellus huysi; Sp: Scutogerulus pelophilus. g: gonopore; c: copulatory pore.

morphic genital system. In Scutogerulus and Pilarella each

copulatory pore shares a common opening with the gonop-

ore, whereas in Crassarietellus each copulatory pore is sepa-

rate from the gonopore and located beneath the ventral flap.

The latter is probably more derived since the copulatory

pores are separate from the gonopores. In both trends, the

copulatory duct is relatively short and the seminal receptacle

is a simple spherical shape.

In the specimens of Crassarietellus examined, a pair of

fertilization tubes from the spermatophore remnant (Figs

2A,3) was still connected to the copulatory pores. In this

genus each copulatory pore seems to be relatively large and

Opens onto the inner surface of the ventral flap. The end of

the fertilization tube terminates in a mass of brownish opaque

material (see Fig. 1E, dotted) positioned where the copula-

tory pore opens. The gonopore is not covered by a genital

operculum, as in other arietellid genera (Fig. 2C,D). An

exposed gonopore, as in Crassarietellus, is also found in the

deep-sea hyperbenthic calanoid family Hyperbionycidae

(Ohtsuka et al., 1993b). Owing to the complete absence of

armature elements on leg 6 in the Calanoida, it is unknown

whether the absence of a genital operculum in Crassarietellus

represents a secondary loss or a more plesiomorphic state

than other arietellids. Radiation of the genital systems of

arietellids can be related to their different habitats. Gener-

ally, deep-sea hyperbenthic genera such as Crassarietellus and

162

Scutogerulus exhibit a more primitive state than genera found

in other habitats, with the exception of Sarsarietellus which,

however, may be a deep-water hyperbenthic species (Cam-

paner, 1984). In contrast, the shallow-water pelagic and

hyperbenthic genera Metacalanus, Paramisophria and

Pilarella independently exhibit relatively derived genital sys-

tems. The bathypelagic genera Arietellus and Paraugaptilus

have also independently developed a more apomorphic geni-

tal system than the deep-sea hyperbenthic genera.

3. Caudal ramus. The caudal rami of almost all arietellids

are symmetrical. However, asymmetry of caudal rami is

exhibited in Scutogerulus, in which the left ramus is slightly

longer than the right (Bradford, 1969; present study, Fig.

33A), and in Pilarella in which the left caudal ramus is slightly

larger than the right (present study).

Except in Metacalanus the armature elements on the

caudal ramus are all retained. In all genera seta I is minute

and setae III-VII are developed to varying degrees. Seta II is

relatively minute or completely absent in Metacalanus, but

always present in the other genera. Arietellus pavoninus has

highly specialized caudal rami with densely plumose seta II

that is directed anteriorly (Sars, 1924, 1925).

4. Rostrum. All arietellids have a well-developed rostrum

produced ventrally with a pair of filaments. Both sexes of

Metacalanus curvirostris have a rostrum that curves to the left

(Ohtsuka, 1985).

5. Female antennule. The antennulary segmentation and

setation patterns of female arietellids are summarized in Fig.

39. Some genera show variability in segmentation and/or

setation. In particular, Metacalanus exhibits asymmetry in

both segmentation and armature (Fig. 22). The segmentation

and setation of Crassarietellus represent the most plesiomor-

phic state within the family, displaying both the maximum

segmentation and the greatest number of armature elements

as follows (Fig. 39A): separation of ancestral segment III

from IV; segments IV to XXI each with 2 setae and aes-

thetasc; segments X—XII separate; segments XIV and XV

separate; segments XXIII and XXIV separate.

Ancestral segments I-III are fused in Crassarietellus (Fig.

39A), Scutogerulus (Fig. 39C), Sarsarietellus (Fig. 39B) and

Paramisophria (Fig. 39D), and segments I-IV in Arietellus

(Fig. 39E), Metacalanus (Fig. 39G), Paraugaptilus (Fig. 39F)

and Pilarella (Fig. 39H). Segments XXIII and XXIV are

separate in Crassarietellus, Paramisophria, Scutogerulus, Sar-

sarietellus, Metacalanus and Pilarella, and fused in Arietellus

and Paraugaptilus. The complete fusion of segments IX and

X is unique to Metacalanus.

The loss of an aesthetasc on segment IV is found in seven

genera; that on segment II in Pilarella; that on segment VI in

Arietellus, Paraugaptilus, Metacalanus and Pilarella; those on

segments VIII and X in Paraugaptilus and Metacalanus; that

on segment XII in Arietellus and Paraugaptilus; those on

segments XXII and XXIII in Pilarella. One element on

segment XIII is reduced in Paraugaptilus and Metacalanus.

One seta on compound segment XX VI-XXVIII is reduced in

Arietellus and in Paraugaptilus similis.

The presence of a duplicated aesthetasc at the extreme tip

of antennule of Paraugaptilus similis is interpreted here as an

individual abnormality.

The right and left antennules are markedly asymmetrical in

length in the genera Paramisophria, Metacalanus and

Pilarella, which are mainly distributed near the sea bed. This

asymmetry has been related to the peculiar swimming behav-

iour of these genera at the sediment-water interface (see

S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

Ohtsuka & Mitsuzumi, 1990). The ornamentation of the right

and left antennules is slightly asymmetrical on the terminal

segments in the bathypelagic genus Paraugaptilus.

6. Male left antennule. The antennulary segmentation and

setation patterns of male arietellids are summarized in Fig.

40. Ancestral segments II to IV are incompletely fused in

Campaneria (Fig. 40A) and completely fused in the other six

genera. Fusion of segments [X—X is unique to Metacalanus

(Fig. 40G), whereas complete separation of segment XXI

from XXII is found only in Campaneria. Each of ancestral

segments II and III carries 2 setae and an aesthetasc in

Crassarietellus (Fig. 40B), and 1 seta and an aesthetasc in the

other genera. In Arietellus aculeatus segments I to IV bear 1,

2, 2 and 2 aesthetascs, respectively. The presence of one

additional aesthetasc on each segment from II to IV seem to

be a secondary addition found in the males of many pelagic

calanoids (see Huys & Boxshall, 1991). Huys & Boxshall

(1991) speculated that duplication of aesthetascs in males is

an adaptation for the open pelagic environment. The oceanic

pelagic species A. aculeatus shows duplication of aesthetascs,

and neither shallow- nor deep-water hyperbenthic arietellids

have such duplication. However, no other pelagic species of

either Arietellus or Paraugaptilus has such duplication, and its

occurrence within a single species of a relatively derived

genus may indicate that the duplication of aesthetascs in A.

aculeatus arose independently.

A seta on segment XV is modified, by loss of its proximal

articulation with the segment, into a process in Arietellus,

Paraugaptilus and Paraugaptiloides; a seta on segment XXII

is also modified into a process in Crassarietellus, Campaneria

and Metacalanus. Only in Paraugaptilus and Paraugaptiloides

does the compound segment XXIV-—XXV carry a large

distally directed process (Figs 11B, 30E, 32F). From its

position, this process may be derived from a setation element

of segment XXIV, but we consider it more likely that it

represents an outgrowth of the segment. The loss of a seta on

the compound segment XXVI-XXVIII is found in Arietellus

and Paraugaptilus. The lack of a seta on segment XIII is

unique to Metacalanus.

7. Antenna. The ancestral condition of the antennary

exopod of Copepoda is shown by Huys & Boxshall (1991):

the exopod consists of 10 separate segments; first to ninth

segments each bearing a single seta, the 10th segment with 3

setae (Fig. 41A). The segmentation and setation patterns of

the arietellid genera are schematically depicted in Fig.

41B-H. In all genera, ancestral exopodal segments I and II, V

and VI, VI and VII, and VII and VIII are either completely

separate or incompletely fused with a suture still visible. In all

genera, ancestral segments IV, V, VI and VII each carry 1

seta while segments I, II, III and IX are unarmed. Segment X

carries 3 setae except in Paraugaptilus (Fig. 41G,H). A seta is

present on segment VIII in Crassarietellus (Fig. 41B), Cam-

paneria (Fig. 41D), Paraugaptiloides (Fig. 41D), Parami-

sophria (Fig. 41D), Metacalanus (Fig. 41E), Sarsarietellus

(Fig. 41D), Scutogerulus (Fig. 41D) and Pilarella (Fig. 41D),

but absent in Arietellus (Fig. 41F) and female Paraugaptilus

(Fig. 41H). Complete fusion of ancestral segments I-IV

occurs in Campaneria, Paraugaptiloides, Arietellus, Parami-

sophria, Metacalanus, Paraugaptilus, Sarsarietellus, Scu-

togerulus and Pilarella. Complete fusion of segments VIII-IX

occurs in Arietellus, Metacalanus and female Paraugaptilus.

The most advanced state is found in female Paraugaptilus

(Fig. 41H): ancestral segments VIII to X are completely

fused to form an unarmed, bulbous compound segment in P.

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PHYLOGENY OF ARIETELLID COPEPODS

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PHYLOGENY OF ARIETELLID COPEPODS

similis, P. buchani, P. bermudensis (Deevey, 1973) and P.

meridionalis (= P. buchani sensu Sars, 1924, 1925). In

contrast, males of P. similis and P. bermudensis are relatively

plesiomorphic in that compound segment VIII-X retains a

seta which is derived from ancestral segment VIII.

In contrast to the exopodal segmentation, the endopods of

arietellids are constantly 2-segmented with the second to

fourth ancestral segments almost completely fused. The first

segment bears a single minute seta in Crassarietellus, Cam-

paneria, Paramisophria, Metacalanus and Pilarella, and is

unarmed in Paraugaptiloides, Arietellus, Paraugaptilus, Sar-

sarietellus and Scutogerulus. The number of inner setae on

the second compound segment is variable: 3 in Crassarietel-

lus, Campaneria, Paramisophria, Sarsarietellus, Scutogerulus

and Pilarella; 2 in Paraugaptiloides, Arietellus, Paraugaptilus

and Metacalanus (Paraugaptilus has 1 or 2 setae on it). The

number of terminal setae on the compound segment is 6 in

Paraugaptiloides, Arietellus, Paramisophria, Paraugaptilus

and Sarsarietellus, and 5S in Crassarietellus, Campaneria,

Metacalanus, Scutogerulus and Pilarella.

Sexual dimorphism is found in the antennary rami of

Arietellus and Paraugaptilus. The reduction of one of the 2

medial setae on the second endopodal segment of Arietellus

and some species of Paraugaptilus is retained only in the

female. In Paraugaptilus the relative length of the first and

second endopodal segments is different in the sexes. In

addition, some species of Paraugaptilus (Deevey, 1973;

present study) exhibit sexual differences in the exopod in that

the ancestral segment VIII is completely fused with segment

IX-X in the female and is unarmed, but incompletely fused

165

with the compound segment and carrying 1 seta in the male.

The male shows a more plesiomorphic state in antennary

rami than the female.

8. Mandible. Arietellids are typically carnivorous, feeding

on copepods and other small organisms (e.g., Ohtsuka &

Mitsuzumi, 1990; Ohtsuka et al., 1991). Their mandibular

gnathobases are well developed and heavily chitinized, with

three or four sharp teeth.

The endopod is either reduced to 1 segment with 1 or 2

setae, or is unarmed and completely fused with the basis. The

more plesiomorphic state is retained in Crassarietellus, Cam-

paneria, Paraugaptiloides, Paramisophria, Metacalanus, Sar-

sarietellus, Scutogerulus and Pilarella, and the derived state

found in Arietellus and Paraugaptilus.

The first exopodal segment has a normally developed seta

in all genera, except for some species of Arietellus and

Paraugaptilus. In these two genera this seta is sexually

dimorphic. The males are furnished with a normally devel-

oped seta, whereas the females bear a vestigial seta (Sars,

1924; Deevey, 1973; present study). On the fifth exopodal

segment, the remarkable reduction of the outer terminal seta

is exhibited only by Arietellus (Figs 13D,18B) and Paraugap-

tilus (Fig. 32B).

9. Maxillule. Arietellids exhibit a wide variety of trans-

formed states in the praecoxal arthrite, the coxal endite and

epipodite, the basal endite and the endopod. These charac-

ters were used to define some arietellid genera by previous

authors such as Sars (1903), Rose (1933), Brodsky (1950),

Campaner (1977) and Ohtsuka et al. (1993a).

The maximum number of elements on the praecoxal arth-

Fig. 41.

Schematic illustration of fusion patterns and armature of antennary exopods of the arietellid genera. A, Hypothetical calanoid

ancestor; B, Crassarietellus; C, Paramisophria giselae; D, Campaneria, Paraugaptiloides, Paramisophria japonica, Sarsarietellus ,

Scutogerulus; E, Metacalanus; F, Arietellus; G, Paraugaptilus similis, male; H, P. similis, female. Solid and dotted lines indicating complete

separation between segments, and incomplete fusion or suture between segments, respectively.

166

rite (5 spines and process) occurs in Crassarietellus, Campan-

eria, Paraugaptiloides, Arietellus, Paramisophria,

Sarsarietellus and Pilarella. In Sarsarietellus the outer proxi-

mal spine is incompletely fused to the arthrite, while in the

other six genera the fusion is complete enough to form a

process. Both Paraugaptilus (5 spines) and Scutogerulus (4

spines and process) show more advanced states, and the

reduced element may be the inner proximal spine in both

genera. Metacalanus exhibits the most apomorphic state, in

the number of elements (0-2 setiform spines), and the

elements are not so strongly chitinized as in other genera.

On the coxal endite a single seta is present in all the genera

except for Paraugaptilus. The relative length and the orna-

mentation of the seta are variable within polytypic genera.

The number of setae on the coxal epipodite varies in ari-

etellids. The maximum number (8 setae) is retained in

Paraugaptiloides, Arietellus, Paraugaptilus and Sarsarietellus ,

whereas there are 6 in Crassarietellus and Campaneria, 5 in

Metacalanus, Scutogerulus and Pilarella. A vestigial basal

seta is present in Crassarietellus, Campaneria, Paraugapti-

loides, Paramisophria and Sarsarietellus, but absent in Ari-

etellus, Metacalanus, Paraugaptilus and Pilarella. The

position of this seta indicates that it probably represents the

second basal endite.

The endopod is variously modified. The most plesiomor-

phic state, 1-segmented with 3 setae, is found in several

species of Paramisophria. A 1-segmented endopod with 2

setae is present in Crassarietellus, Campaneria, Paraugapti-

loides, Arietellus, Sarsarietellus and Pilarella; a 1-segmented

endopod with a single seta in Arietellus, Metacalanus and

Scutogerulus. Species of Arietellus and Metacalanus, espe-

cially the former, exhibit a variety of transformed states in the

endopod. The most apomorphic state in these 2 genera is

complete incorporation into the basis. Several species of

Arietellus display an intermediate state with the endopod

represented by a rudimentary, unarmed knob, almost fused

to the basis. In Paraugaptilus the endopod is completely

incorporated into the basis.

10. Maxilla. The armature elements on the first and second

praecoxal endite, and the ornamentation on the basal and

endopodal setae are unique to each genus. On the first

praecoxal endite the most primitive state (2 setae and a

vestigial element) is retained in Crassarietellus, Campaneria,

Paraugaptiloides, Sarsarietellus, Paramisophria (only P. gise-

lae) and Pilarella. Arietellus, Metacalanus, Paraugaptilus and

Scutogerulus share the more apomorphic state (1 seta and a

vestigial element). In all these genera it is the outer seta on

the endite of the more plesiomorphic genera that is absent

and the inner one that remains, based on the position of the

setae on the endite.

On the second praecoxal endite, 2 setae are present in

Crassarietellus, Campaneria, Paraugaptiloides, Arietellus,

Paramisophria, Metacalanus, Sarsarietellus and Pilarella, and

a single seta in Paraugaptilus and Scutogerulus. All genera

exhibit 2 setae on the first and second coxal endites. The

basal spine is variously ornamented in all genera except for

Paramisophria whose spine is bare. In Campaneria (Fig.

10G), Paraugaptiloides (Fig. 11F), Arietellus (Figs

131,18F,G) and Sarsarietellus (Fig. 36G), the basal spine is

relatively elongate, ornamented with 2 rows of fine, long

spinules densely distributed along the entire length except for

the bare terminal part. Crassarietellus (Figs 5B,8D) also

carries a long basal spine with 2 rows of relatively thick

spinules distributed about at midlength. In Paraugaptilus

S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

(Fig. 29D) and Scutogerulus (Fig. 34D), the spinules are

minute and sparsely distributed. Metacalanus (Fig. 211) bears

a basal spine unique within arietellids; the spine is relatively

short, with 2 rows of minute, rigid spinules at midlength. In

Pilarella the basal spine is elongate with a single row of

spinules at midlength.

The ornamentation on the endopodal setae is also charac-

teristic of each genus. In Crassarietellus, Campaneria,

Paraugaptiloides, Paramisophria, Metacalanus, Sarsarietellus

and Pilarella, the inner margin of these setae is furnished with

a row of slender, simple spinules (see Fig. 11G), whereas in

Arietellus, Paraugaptilus and Scutogerulus the ornamentation

is variable. Arietellus develops a lobate structure basally on

each spinule (Fig. 15B,C), while both Paraugaptilus (Fig.

27G) and Scutogerulus (Fig. 34F) carry a row of triangular

spinules along the inner margin of each seta. In arietellids

such setal ornamentation on the maxilla is also found on the

well-developed setae of the terminal endopod segments of

the maxilliped. Bradford (1969) referred to the setal orna-

mentation on the maxilla and maxilliped of Scutogerulus as

‘shield-shaped appendages’ in her definition of the genus.

11. Maxilliped. Variation in arietellids can be found in the

armature on the fourth to sixth endopodal segments. The

innermost seta on the fourth and fifth segments is relatively

well-developed in all the genera except for Arietellus, in

which it is reduced to a vestigial element or is completely

absent. In Crassarietellus (Figs 6B,C,8E), Metacalanus (Fig.

26A,B), Paramisophria (Fig. 19D), Paraugaptilus (Fig. 27F)

and Pilarella, the innermost setae on the fourth and fifth

endopodal segments are of almost equal length; in Campan-

eria (Fig. 10H), Paraugaptiloides (Fig. 12B), Sarsarietellus

(Fig. 35C,D) and Scutogerulus (Fig. 34E) the innermost seta

on the fourth endopodal segment is longer than that on the

fifth.

On the sixth endopodal segment the most plesiomorphic

state, with setae a and b developed, is retained in Crassari-

etellus (Fig. 5C), Paramisophria (Fig. 19E), Metacalanus

(Fig. 26C) and Pilarella (Fig. 37F); the most apomorphic

state, namely, reduced setae a and b is found in Arietellus

(Fig. 18H-K), Paraugaptilus (Fig. 27G) and Scutogerulus

(Fig. 34F). Campaneria (Fig. 101), Paraugaptiloides (Fig.

12B) and Sarsarietellus (Fig. 35E) show an intermediate

condition: only seta a is reduced and seta b is relatively long.

In Paraugaptilus only seta c is specialized, with its terminal

part heavily chitinized and serrated along the inner margin

(Figs 27G, 32E). Paraugaptiloides, however, shows no spe-

cialization of seta c (Fig.12B).

12. Leg 1. On the third exopodal segment two outer spines

are retained in Crassarietellus, Campaneria, Paraugaptiloides,

Arietellus, Paramisophria, Paraugaptilus and Sarsarietellus.

A single outer spine is found in Metacalanus, Scutogerulus

and Pilarella. Consideration of the relative position of the

spines suggests that it is the proximal spine that is lost in these

three genera.

13. Legs 2 and 3. All genera and species, except for the

cave-dwelling Paramisophria galapagensis, retained the maxi-

mum setation of the endopods of legs 2 and 3: seta and spine

formula 0-1;0-2;2,2,4. In P. galapagensis the seta and spine

formula of the endopod is 0—1;0—2;2,2,3 (Ohtsuka et al.,

1993a). This represents the most apomorphic state known in

arietellids.

14. Leg 4. An inner coxal seta or a vestigial element is

present only in Paraugaptiloides, Paraugaptilus and Pilarella.

It is absent in the other genera, although a fourth copepodid

PHYLOGENY OF ARIETELLID COPEPODS

stage of Paramisophria sp. collected from South Japan carries

a minute inner coxal seta (Ohtsuka et al., 1991, Fig. 6J,K).

The maximum setation on the third endopodal segment is

retained in all the genera and species except for P. galapagen-

sis: 2,2,2 in P. galapagensis and 2,2,3 in other taxa (Ohtsuka

et al., 1993a).

15. Female leg 5. The female fifth legs of arietellids are

variable, as in several other calanoid families and the

misophrioid family Misophriidae by Huys & Boxshall (1991).

Campaner (1984) compared the structure of leg 5 in both

sexes but drew no strict homologies of segmentation and

armature elements.

Fig. 42 schematically depicts apparent evolutionary trends

in the structure of female leg 5 within the genera Arietellus,

Paraugaptilus, Paramisophria, Metacalanus and Pilarella.

Within the genus Arietellus, three obvious evolutionary

trends in segmentation and setation can be recognized:

incorporation of the endopod into the basis, reduction of

endopodal setae, and fusion of coxa, basis and both rami. The

genus Paramisophria also exhibits two distinct evolutionary

trends: reduction in numbers of endopodal setae and of

exopodal spines. In the genus Metacalanus reduction of the

endopod, and fusion of both rami into the basis plus reduc-

tion in number of elements on the exopod occur. Based on

these evolutionary trends, the derivation of the Paraugaptilus

state from an Arietellus-like condition, the relationships

between Sarsarietellus and Paramisophria spp., and the deri-

vation of Metacalanus from a Paramisophria-like ancestor, as

already proposed by Campaner (1984), are supported. The

setation of Crassarietellus (Fig. 6K,L) suggests a close rela-

tionship with Paramisophria, especially in the endopod seta-

tion.

Consideration of the plesiomorphic states exhibited in leg 5

of all female arietellids indicates that the hypothetical ances-

tor may be characterized by having retained a) the coxa, the

basis and 3-segmented exopod and 2-segmented endopod as

separate segments; b) basal seta present; c) intercoxal sclerite

separate from coxae; d) setal formula of endopod segments

0-2;0,1,1; and e) setal formula of exopod I-0;I-0;I1,1,0.

In Crassarietellus and Scutogerulus the endopod is dis-

tinctly separate from the basis, is 1-segmented, and bears 2

and 1 setae respectively. In Arietellus, Paramisophria, Meta-

calanus, Paraugaptilus, Sarsarietellus and Pilarella the endo-

pod is completely or incompletely fused with the basis, and is

represented by 0-4 setae. In Paramisophria the number of

setae on the endopod ranges from 0 to 2; in Arietellus from 1

to 3 setae. In Metacalanus, Paraugaptilus and Pilarella the

endopod is represented by 0-1 seta, and is almost completely

incorporated into the basis.

In P. japonica (Ohtsuka et al., 1991, Fig. 3F,G) and

Scutogerulus (Bradford, 1969, Fig. 181) the exopod is com-

posed of 2 distinct segments. Particularly in P. japonica the

ancestral second and third exopodal segments are incom-

pletely fused with a suture visible on the anterior surface. In

Crassarietellus (Fig. 6K,L) and Sarsarietellus (Fig. 36H) the

first to third exopodal segments are almost fused with a

suture just visible. In Arietellus (except for A. mohri and A.

sp.), almost all species of Paramisophria (except for P.

giselae), Metacalanus (except for M. aurivilli and M. acutio-

perculum) and Pilarella, the exopod is distinctly 1-segmented,

but variably armed. Arietellus carries only a single terminal

spine; Paramisophria bearing 2 or 3 lateral and 2 terminal

spines; Metacalanus has 1 terminal spine or 2 terminal and 1

lateral spine. The unarmed exopods of A. mohri and A. sp.

167

are lobate and almost completely fused with the basis. In M.

aurivilli and M. acutioperculum the exopod is represented by

a small knob with a single terminal seta. In Paraugaptilus the

exopod is completely incorporated into the basis.

The intercoxal sclerite and coxa are completely separate in

Sarsarietellus, Metacalanus, Pilarella and P. giselae, and

incompletely in Crassarietellus and Arietellus (except for A.

mohri and A. sp.). In Paramisophria (except for P. giselae),

Paraugaptilus, A. mohri and A. sp. fusion is almost or

completely accomplished.

Little attention was paid to the variability within a genus by

Campaner (1984). Within genera such as Arietellus, Parami-

sophria and Metacalanus, the reduction in segmentation and

setation is more variable than expected. Reductions in seg-

mentation and setation appear to occur independently within

each genus. For instance, the fusion between coxa and

intercoxal sclerite probably evolved independently in Arietel-

lus (see Fig. 17) and Paramisophria (Fig. 20E,F). The num-

ber of elements on both rami vary widely in these genera,

whereas the outer basal seta is consistently present in all

genera and species. In Arietellus the right basal seta is slightly

or considerably longer than the left.

16. Male leg 5. Campaner (1984) showed a possible rela-

tionship between the male fifth legs of arietellids, based

mainly on the presence or absence of the endopod on either

side. However, the homologies of segmentation and setation

were not considered in detail. Compared with the female fifth

legs, the male legs are less variable in segmentation and

setation within a genus. A scheme indicating possible deriva-

tions of segmentation and setation is given in Fig. 43.

The hypothetical ancestral state is based on all taxa and

consists of a) intercoxal sclerite and coxa separate; b) coxa

completely separate from basis; c) basal seta present; d)

2-segmented, unarmed left endopod; e) 1-segmented,

unarmed right endopod; f) 3-segmented right and left exo-

pods; and g) setal formula I-0;I-1;J1,1,0. The presence of a

basal seta and the numbers of first and second exopodal

elements are constant in all genera.

Although the left endopod of Paramisophria japonica

(Ohtsuka et al., 1991, Fig. 4K) and the right endopod of

Paraugaptiloides (Fig. 12E) each bear a minute terminal

spinule, we are not certain whether it is homologous with a

true setation element.

In Campaneria, Paraugaptiloides, Arietellus and Paraugap-

tilus, both right and left endopods are present. In the first

three genera a distinctly or indistinctly 2-segmented left

endopod is present, while the right endopods of all four

genera comprise a single segment. In Paraugaptiloides the

first and second endopodal segments are completely separate

and are accompanied by musculature, indicating that the

articulation between these segments is functional. In Cras-

sarietellus and Paramisophria (except for P. cluthae) only the

left endopod is retained and the right endopod is absent; the

former has an indistinctly 2-segmented left endopod while in

the latter this ramus is 1-segmented. In Metacalanus both

right and left endopods are completely absent.

The most plesiomorphic state in segmentation and arma-

ture of the exopod is retained in Paramisophria: in both legs,

the third segment is separate from the second (cf. Fosshagen,

1968) and 4 elements are present on the third segment of both

legs (see Ohtsuka & Mitsuzumi, 1990, Fig. 4E,F). In Cras-

sarietellus and Paraugaptiloides a vestigial outer proximal

element is present on the left third exopod segment, which

carries 4 elements in total. The number of elements on the

168

S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

Am Ps

Pi P

Fig. 42. Schematic comparison of patterns of segmentation and setation of female fifth legs in some arietellids. The arrows indicate possible

derivations of setation and segmentation patterns and are not indicative of ancestor-descendant relationships between taxa. Ap: Arietellus

pavoninus; As: A. sp.; Am: A. mohri; Ps: Paraugaptilus similis; Sa: Sarsarietellus abyssalis; Pj: Paramisophria japonica; Pi: P. itoi; Pp: P.

platysoma; Pg: P. giselae; Pr: P. reducta; Ch: Crassarietellus huysi; M1: Metacalanus species 1; M2: M. species 2; Ma: M. acutioperculum;

Pl: Pilarella longicornis. C: Coxa; B (in Ch): Basis; Is: Intercoxal sclerite; Ex: Exopod; En: Endopod. A-D (in Sa): setae on endopod; a-f:

spines on exopod.

third exopod segment of the right leg is 3 in Campaneria, 2 in

Paraugaptiloides, Arietellus and Paraugaptilus, and 1 in Meta-

calanus; on the left leg it is 3 in Campaneria and Arietellus

and 1 in Metacalanus. The distal two exopodal segments are

separate in both legs in Paraugaptiloides, Paramisophria and

Metacalanus, and fused in both legs of Campaneria and

Paraugaptilus and in the right leg only in Arietellus. The distal

two segments of the right leg are missing in the only known

male of Crassarietellus sp. The terminal and subterminal

elements on the third exopodal segment of the left leg are

heavily chitinized and almost fused to the segment only in

Paraugaptiloides, Arietellus and Paraugaptilus.

PHYLOGENY OF ARIETELLID COPEPODS

Ancestor

169

Fig. 43. Schematic comparison of segmentation and setation of male fifth legs in the Arietellidae. The arrows indicate possible derivations of

setation and segmentation patterns and are not indicative of ancestor-descendant relationships between taxa. Ch: Crassarietellus huysi; Pp:

Paramisophria platysoma; Pm: Paraugaptiloides magnus; Cl: Campaneria latipes; M1: Metacalanus species 1; Ap: Arietellus plumifer; Pb:

Paraugaptilus buchani. C: Coxa; B: Basis; Is: Intercoxal sclerite; Ex: Exopod; En: Endopod. a-f,k: elements on exopod. Setae and spines

are not distinguished here.

The intercoxal sclerite and both coxae are almost fused,

with the suture clearly visible in Crassarietellus and Campan-

eria, while in the other genera fusion is complete. The basis

and coxa are completely separate in both legs in Crassarietel-

lus, Campaneria, Paramisophria and Metacalanus, almost

completely fused in the right leg but completely separate in

the left leg in Paraugaptiloides, Arietellus and Paraugaptilus.

Phylogenetic relationships between arietellid

genera

Phylogenetic relationships between the 10 genera studied in

this paper were analyzed using PAUP 3.0 on a matrix of 44

characters (Tables 2,3). The matrix contains a significant

proportion of missing data, shown in the matrix by a ‘9’

(Table 3). These missing data correspond to the unknown

males of the genera Scutogerulus, Sarsarietellus and Pilarella

and to the unknown females of Campaneria and Paraugapti-

loides. Since most of the characters used in the analysis are

sexually dimorphic (30 out of 44 characters), only a minority

of characters (14 of 44) can be scored for all taxa. The

phylogenetic scheme presented here is necessarily tentative,

subject to re-examination as the gaps in the data matrix are

filled by the discovery of unknown sexes.

Four trees were generated by the analysis, all with the same

statistics: tree length = 179; consistency index = 0.263;

homoplasy index = 0.737. These four trees differed only in

the relative positions of Campaneria, Paraugaptiloides and

Sarsarietellus. The relative positions of all other genera are

the same. All three of these genera are known from only one

sex. Tree 1 (Fig. 44) was selected as the best working

hypothesis of relationships because Campaneria was the first

offshoot of the Arietellus-group, as it was in three of the four

trees, and because it placed Sarsarietellus as an earlier

offshoot than Paraugaptiloides which we consider to be the

more apomorphic genus of the two.

The genera of the Arietellidae form two lineages, the

Arietellus-group comprising six genera, and the Metacalanus-

group consisting of four genera. The Arietellus-group is

diagnosed by the apomorphic reduction of seta a on the

terminal segment of the maxillipedal endopod (character 27).

The Metacalanus-group lacks a simple diagnostic character.

The apomorphic state of character 38 (absence of endopod of

male right fifth leg) is found only within the group, in

Crassarietellus, Paramisophria and Metacalanus (the male of

Pilarella is unknown), and the apomorphic state of character

3 (asymmetrical antennules in females) is found only in

Paramisophria, Metacalanus and Pilarella. Crassarietellus

retains the plesiomorphic state.

This analysis suggests that there may have been several

170

Fig. 44. Cladogram depicting relationships among arietellid genera.

shifts in habitat utilization during the evolutionary history of

the family. Substitution of habitat type (Fig. 45) onto the

cladogram shown in Fig. 44 indicates that the Arietellidae

originated in the hyperbenthic zone. The most plesiomorphic

representatives of both lineages still inhabit this zone. The

Metacalanus-group has largely remained in the ancestral

hyperbenthic habitat although it has successfully colonized

anchialine caves (Ohtsuka et al., 1993a) and at least one

species of Metacalanus is epipelagic. In contrast, the most

apomorphic representatives of the Arietellus-group, the gen-

era Arietellus and Paraugaptilus, have successfully colonized

the open pelagic realm.

A similar analysis of habitat utilization was performed on

the genera of the copepod family Misophrioidae by Boxshall

(1989). The 10 genera of this family were placed in two

lineages, both of which originated in the deep-water hyper-

benthic zone. The first offshoot of the Archimisophria-

lineage, represented by the genus Archimisophria Boxshall,

1983, has remained in the ancestral habitat but all the derived

representatives of this lineage are found in anchialine caves

and crevicular habitats. The most plesiomorphic representa-

tive of the Misophria-lineage, the genus Misophriopsis Box-

shall, 1983, also inhabits the hyperbenthic zone but other

members of the lineage have successfully colonized the

pelagic zone, the shallow-water hyperbenthic zone and, inde-

pendently, anchialine habitats.

There are interesting parallels between the Arietellidae

and Misophriidae. The ancestry of both families appears to

be closely associated with the deep-water hyperbenthic zone.

Plesiomorphic genera in both families have remained in the

ancestral habitat but more derived representatives now utilize

a broader spectrum of habitat types, including the shallow-

S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE

Crassarietellus

Paramisophria

Metacalanus

Pilarella

Arietellus

Paraugaptilus

Scutogerulus

Paraugaptiloides

Sarsarietellus

Campaneria

water hyperbenthic zone, the open pelagic realm and anchia-

line caves. Certain habitat shifts appear to have occurred at

least twice, independently, within these two families. The

colonization of anchialine habitats appears to have taken

place twice in the Arietellidae, once within Metacalanus and

once within Paramisophria, just as Boxshall (1989) found for

the Misophriidae. Arietellids appear to have invaded the

open pelagic zone three times (the Arietellus-Paraugaptilus

group, Paraugaptiloides, and within the genus Metacalanus).

Key to genera of the family Arietellidae

la_ Leg 1 with 1 outer spine on third exopod segment ............. 2

1b Leg 1 with 2 outer spines on third exopod segment ............ 3

2a Maxillule with 5 spines and 1 process on praecoxal arthrite;

maxilla with 1 seta on distal praecoxal endite; caudal seta II

developed; genital double-somite (2) with paired genital sys-

tem, each copulatory pore opening within slit-like genital slit,

shared with gonopore .............. Scutogerulus Bradford, 1969

2b Maxillule with 5 spines and 1 process on praecoxal arthrite;

maxilla with 2 seta on distal praecoxal endite; caudal seta II

developed; genital double-somite (2) with paired genital sys-

tem, each copulatory pore opening within common genital

aperture, shared with gonopore ........ Pilarella Alvarez, 1985

2c Maxillule with 0-2 elements on praecoxal arthrite; maxilla with

2 setae on distal praecoxal endite; caudal seta II reduced;

genital double-somite (2) with gonopore and copulatory pore

separate and located posteriorly ..... Metacalanus Cleve, 1901

3a Maxillule with 6 setae on coxal epipodite ......................45- 4

3b Maxillule with 8-9 setae on coxal epipodite .....................- 5

PHYLOGENY OF ARIETELLID COPEPODS

171

Hyperbenthic

Hyperbenthic/cave

Hyperbenthic/cave/

epipelagic

Hyperbenthic

Pelagic

Hyperbenthic

Pelagic

Hyperbenthic

Fig. 45. Habitat cladogram of arietellid genera. Substitution of habitat type of each genus onto cladogram shown in Fig. 44.

Antennary exopod indistinctly 10-segmented; maxillulary prae-

coxal arthrite with strongly serrate spines; long innermost seta

on fifth endopod segment of maxilliped; outermost seta on sixth

endopod segment of maxilliped not reduced; left antennule (0’)

with 2 setae on segments II and III, and segments XXI and

XXII fused; right endopod of leg 5 (C’) lacking

Pee essere dae iscndesseccsecesetssesetenees Crassarietellus gen. nov.

Antennary exopod indistinctly 8-segmented; maxillulary prae-

coxal arthrite with weakly serrate spines; short innermost seta

on fifth endopod segment of maxilliped; outermost seta on sixth

endopod segment of maxilliped reduced; left antennule with 1

seta on segments II and III, and segments XXI and XXII

separate; right endopod of leg 5 (C’) present

Nt Pees Saris soma clasiotix eS nest nmecutaeeee.ss Campaneria gen. nov.

Innermost seta on fourth and fifth endopod segments of maxil-

INEGIVESMOTAl cc secswacsccssseasaneee tess Arietellus Giesbrecht, 1892

Innermost seta on fourth and fifth endopod of maxilliped not

SESH ICININ a - SECs See ence acme rc sales aaa on wndee een saaeenetes ase 6

Antennary exopod segment X unarmed

PERS Gad a \isaie oaMaie asiaelssive santas Paraugaptilus Wolfenden, 1904

Antennary exopod segment X with 3 elements

Leg 4 with inner coxal seta; second antennary endopod segment

with 2 inner setae at midlength .... Paraugaptiloides gen. nov.

Leg 4 without inner coxal seta; second antennary endopod

segment with 3 inner setae at midlength

Antennulary segments XXV and XXVI separated; basal spine

of maxilla ornamented with spinules; outermost seta on sixth

endopod segment of maxilliped vestigial; genital double-somite

(2) with copulatory pore located midventrally on median line or

on left side; copulatory duct heavily chitinized; seminal recep-

tacle elongate, its distal end bulbous in shape; inner process

(derived from endopod) of leg 5 (2) with 4 setae

Saeed ssSapiae adeiteletes webetceis Shes oe veiw alae Sarsarietellus Campaner, 1984

8b Antennulary segments XXV and XXVI fused; basal spine of

maxilla bare; outermost seta on sixth endopod segment of

maxilliped not vestigial; genital double-somite (9) with copula-

tory pore located posteroventrally; seminal receptacle not elon-

gate, its distal end not bulbous; inner process of leg 5 (Q) with

WEDISETAC 2eiimececcs sooo meta case Paramisophria T. Scott, 1897

ACKNOWLEDGEMENTS. We express our sincere thanks to Drs K.

Hulsemann and A. Fosshagen for critical reading of the manuscript.

We are also grateful to Dr. F.D. Ferrari, Dr. J.M. Bradford-Grieve,

Mr. P. Anderson, Miss V. Wilhelmsen and Dr. C.E.F. da Rocha for

lending us material. Thanks are due to Mr. R. Huys and the staff of

the Electron Microscopy Unit of The Natural History Museum,

London for their kind assistance, and to the captain and crew of

T.R.V. Toyoshio-maru of Hiroshima University for their coopera-

tion at sea. The first author (SO) appreciates Prof. T. Onbé for his

encouragement during the present study. The present study was

supported in part by Narishige Zoological Award (1992) and the

Fujiwara Natural History Foundation (1994) awarded to the first

author (SO).

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