. 7 este tale setetens fe oe Feet tasty i ; By His TH Be Ph AEN Hy aosbe tg Cota te Hh sa Henin ait Wine SE ay i aisha HRCI I GREE HEE HEUTE EPL GE ane its Hit } Aenea ie 1 stl f He Bug A a ih Hi i i! witli tt ATHY at tat listiie iH Rit steidt! “ihe RTP EH SHE oe Ht HN eta THES at EA EOL aE i ji fit iia Y ei ite he hy ae are cha i! ta i Hi At i th; ha at Hh Hi ffs 1 7 Ban His iia lai We Tas OL Tne f HTC RH Nie te tdi ; at iy i HU rt ie Puta eae Cane A aE atl tf if HMO RES AUR ea a HO es TRAE AE i i in| ti bs wild : Han fish th Pa) ish it ey il { i j Te at are HIN ah Eile Hat dh i Hi ; H a Se ae reese toe on Sane eae mreeee” Se eg Hi aS wre Fr ett > or osemiceta: - = = pee xe 12 3 Ss Sas = oe. ms aus ti ane, Wericn 7's" me ean Por oa 4 ear bee, bil ah 65, 4 P ib gherg \ 2a we | Ligne fa. ey ae that f° es ; rf rib 1 oe » ? @@ 042Aa o B HISTORY MUSE “9 JUL 1993 ; MNiw ~ iN ra | ZOOLOGY LIBRAR Zoology Series Sz THE NATURAL HISTORY MUSEUM VOLUME 59 NUMBER 1 24 JUNE 1993 The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum (Natural History)), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology. The Zoology Series is edited in the Museum’s Department of Zoology Keeper of Zoology: Dr C.R. Curds Editor of Bulletin: Dr N.R. Merrett Assistant Editor: Dr B.T. Clarke Papers in the Bulletin are primarily the results of research carried out on the unique and ever- growing collections of the Museum, both by the scientific staff and by specialists from elsewhere who make use of the Museum’s resources. Many of the papers are works of reference that will remain indispensable for years to come. All papers submitted for publication are subjected to external peer review for acceptance. A volume contains about 160 pages, made up by two numbers, published in the Spring and Autumn. Subscriptions may be placed for one or more of the series on an annual basis. Individual numbers and back numbers can be purchased and a Bulletin catalogue, by series, is available. Orders and enquiries should be sent to: Intercept Ltd. P.O. Box 716 Andover Hampshire SP10 1YG Telephone: (0264) 334748 Fax: (0264) 334058 World List abbreviation: Bull. nat. Hist. Mus. Lond. (Zool.) © The Natural History Museum, 1993 Zoology Series ISSN 0968 — 0470 Vol. 59, No. 1, pp. 1-96 The Natural History Museum Cromwell Road London SW7 5BD Issued 24 June 1993 Typeset by Ann Buchan (Typesetters), Middlesex Printed in Great Britain at The Alden Press, Oxford 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! y {_ 3 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;5/142;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 2 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 3 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 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 6 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’. i 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. 12 G.J. HOWES DATTA TTT TTT TTT TTT TTT TTT TTT TL Pepe TT TAGTTATp TNT HenT HTT TATA TTTTATT ee 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) 13 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 14 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 15 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. 16 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 17 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 18 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 fX 19 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. 20 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 21 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, 22 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 bh 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 23 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. 24 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 | 25 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. 26 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 nr J, =™ ot nvo gé n¥W nau 27 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. 28 & 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 ki md 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 29 GADOIDE! (Bathygadidae Steindachneriidae Moridoidea Gadoidea 16 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 30 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. REFERENCES Cohen, D.M. 1984. Gadiformes: an overview. Jn: Moser, H.G. (Ed.-in-chief), Ontogeny and systematics of fishes. American Society of Ichthyology and Herpetology Special Publication 1: 259-265. Cohen, D.M.., Inada, T., Iwamoto, T. & Scialabba, N. 1990. Gadiform fishes of the world. Species catalogue. FAO Fisheries Synopsis (125)10:1—442. Dunn, J.R. 1989. A provisional phylogeny of gadid fishes based on adult and early life-history characters. Jn: Cohen, D.M. (Ed.), Papers on the systemat- ics of gadiform fishes. Science Series No. 32. Natural History Museum of Los Angeles County: 109-235. Fahay, M. P. 1989. The ontogeny of Steindachneria argentea Goode & Bean, with comments on its relationships. Jn: Cohen, D.M. (Ed.) Papers on the ANATOMY OF THE MELANONIDAE systematics of gadiforms fishes Science series No.32. Natural History Museum of Los Angeles County: 143-158. Fahay, M.P. & Markle, D.F. 1984. Gadiformes: Development and relation- ships. In: Moser, H.G. (Ed.-in-Chief) Ontogeny and systematics of fishes. American Society of Ichthyology and Herpetology Special Publication 1:265-283. Freihofer, W.C. 1970. Some nerve patterns and their systematic significance in paracanthopterygian, salmoniform, gobioid and apogonid fishes. Proceed- ings of the California Academy of Sciences (4th ser.) 38 (12):215—264. Gosline, W.A. 1968. The suborders of perciform fishes. Proceedings of the United States National Museum 124:1-78. 1971.Functional morphology and classification of teleostean fishes. Uni- versity of Hawaii Press, Hononulu: 208pp. Howes, G.J. 1987. The palatine bone and its associations in gadoid fishes. Journal of Fish Biology 31:625-637. 1988. The cranial muscles and ligaments of macrouroid fishes (Teleostei: Gadiformes); functional, ecological and phylogenetic inferences. Bulletin of the British Museum (Natural History), Zoology series 54(1):1-62. 1989. Phylogenetic relationships of macrouroid and gadoid fishes based on cranial myology and arthrology. Jn: Cohen, D.M. (Ed.) Papers on the systematics of gadiform fishes. Science series No. 32. Natural History Museum of Los Angeles County: 114-128. 1990. The cranial osteology os the southern eel-cod family Muraenolepi- didae, with comments on its phylogenetic relationships and on the biogeog- raphy of sub-Antarctic gadoid fishes. Zoological Journal of the Linnean Society 100: 73-100. 199la. Biogeography of gadoid fishes. Journal of Biogeography 18: 595-622. 1991b. Anatomy, phylogeny and taxonomy of the gadoid fish genus Macruronus Gunther, 1873, with a revised hypothesis of gadoid phylogeny. Bulletin of the British Museum (Natural History), Zoology series 51 (1): 77-110. 1992. Notes on the anatomy and classification of ophidiiform fishes with particular reference to the abyssal genus Acanthonus Gunther, 1878. Bulletin of the British Museum (Natural History), Zoology series 58 (2): 95-111. & Crimmen, 1990. A review of the Bathygadidae (Teleostei: Gadiformes). Bulletin of the British Museum (Natural History) Zoology series 56 (2):155—203. Inada, T. 1989. Current status of the systematics of Merlucciidae. /n: Cohen, D.M. (Ed.) Papers on the systematics of gadiform fishes. Science series No. 32, Natural History Museum of Los Angeles County: 197-207. Iwamoto, T. 1989. Phylogeny of grenadiers (suborder Macrouroidei); another interpretation. Jn: Cohen, D.M. (Ed.) Papers on the systematics of gadiform fishes. Science series No. 32, Natural History Museum of Los Angeles County: 159-173. Lauder, G.V. 1983. Functional design and evolution of the pharyngeal jaw apparatus in euteleostean fishes. Zoological Journal of the Linnean Society ) 77 (1):1-38. Markle, D.F. 1989. Aspects of character homology and phylogeny of the | gadiforms. Jn: Cohen, D.M. (Ed.) Papers on the systematics of gadiform 31 fishes. Science series No. 32, Natural History Museum of Los Angeles County: 59-88. Markle, D.F. & Olney, J.E. 1990. Systematics of the pearlfishes (Pisces: Carapidae). Bulletin of Marine Science 47(2):269-410. Marshall, N.B. 1965. Systematics and biological studies of macrourid fishes (Anacanthini-Teleostii). Deep Sea Research 12: 299-322. 1966. The relationships of the anacanthine fishes Macruronus, Lyconus and Steindachneria. Copeia 1966: 275-280. & Cohen, D.M. 1973. Order Anacanthini (Gadiformes). Characters and synopses of families. Memoir, Sears Foundation for Marine Research 1. Fishes of the western North Atlantic (6): 479-495. Nolf, D. & Steurbaut, E. 1983. Révision des otolithes de téléostéens du Tortonien straotypique et de Montegibbio (Miocéne Supérieur d’Italie septentrionale). Mededelingen van de Werkgroep voor Tertiaire en Kwartaire Geologie 20 (4):143-197. 1989. Evidence from otoliths for establishing relationships within gadiforms. Jn: Cohen, D.M. (Ed.) Papers on the systematics of gadiform fishes Science series No.32, Natural History Museum of Los Angeles County: 89-111. Norman, J.R. 1930. Oceanic fishes and flatfishes collected in 1925-1927. Discovery Reports 2: 261-370. Okamura, O. 1970. Studies on the macrouroid fishes of Japan — morphology, ecology and phylogeny. Reports of the Usa Marine Biological Station, Kochi University 17 (1-2):1-179. 1989. Relationships of the suborder Macrouroidei and related groups, with comments on the Merlucciidae and Steindachneria. In: Cohen, D.M. (Ed.) Papers on the systematics of gadiform fishes. Science series No 32, Natural History Museum of Los Angeles County: 129-142. Patterson, C. & Rosen, D.E. 1989. The Paracanthopterygii revisted: order and disorder. In: Cohen, D.M. (Ed.) Papers on the systematics of gadiform fishes. Science series No. 32, Natural History Museum of Los Angeles County: 5-36. Paulin, C. D. 1983. A revision of the family Moridae (Pisces: Anacanthini) within the New Zealand region. Records, National Museum of New Zealand 2 (9): 81-126. Rosen, D.E. & Patterson, C. 1969. The structure and relationships of the paracanthopterygian fishes. Bulletin of the American Museum of Natural History 141 (3): 357-474. Schwarzhans, W. 1980. Die tertiare Teleosteer-Fauna Neuseelands, rekonstru- iert anhand von Otolithen. Berliner geowissenschaftliche Abhandlungen (A) 26: 1-211. 1984. Fish otoliths from the New Zealand Tertiary. Report New Zealand Geological Survey 113: 1-269. Svetovidoy, A.N. 1948. Treskoobraznye (Gadiformes). Fauna SSSR Zoologis- cheskii Institut Akademii Nauk SSSR (n.s.) (n.s.) 34, Ryby (Fishes) 9 (4):1-222. (English translation, 1962, Jerusalem: Israel Program for Scien- tific Translations, 304pp.) 1953. Additions to the knowledge of fish brain structure. 1. Structure of the brain in Gadidae.Trudy Zoologicheskoo Instituta Akademiya Nauk SSSR, Leningrad 13: 390-419. 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< 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 Heemstra for her excellent art-work. Balon, E.K. & Stewart, D.J. 1983. Fish assemblage in a river with unusual gradient (Luongo, Africa — Zaire system), reflections on river zonation, and description of another new species. Environmental Biology of Fishes 9 (3-4) 225-252. Bell-Cross, G. 1975. A revision of certain Haplochromis species (Pisces : 43 Cichlidae) of Central Africa. Occasional papers of the National Museums and Monuments of Rhodesia, Series B, Natural Sciences 5(7): 405-464. du Plessis, S.S. & Groenewald, A.A. 1953. The kurper of Transvaal. Fauna & Flora, Transvaal Provincial Administration 3: 35-43. Eccles, D.H. & Trewavas, E. 1989. Malawian cichlid Fishes. The classification of some haplochromine genera. 335pp. Lake Fish Movies, Hereten, Ger- many. Greenwood, P.H. 1965. Environmental effects on the pharyngeal mill of a cichlid fish, Astatoreochromis alluaudi, and their taxonomic implications. Proceedings of the Linnean Society of London 176: 1-10. 1979. A review of the pharyngeal apophysis and its significance in the classification of African cichlid fishes. Bulletin of the British Museum (Natural History), Zoology 33: 297-323. — 1979. Towards a phyletic classification of the ‘genus’ Haplochromis (Pisces, Cichlidae) and related taxa. Part I. Bulletin of the British Museum (Natural History), Zoology 35: 265-322. — 1980. Towards a phyletic classification of the ‘genus’ Haplochromis (Pisces, Cichlidae) and related taxa. Part II Bulletin of the British Museum (Natural History), Zoology 39: 1-101. — 1981. The Haplochromine Fishes of the East African Lakes. p. 839. British Museum (Natural History) London & Kraus International Publications, Munich. —— 1984. The haplochromine species (Teleostei, Cichlidae) of the Cunene and certain other Angolan rivers. Bulletin of the British Museum (Natural History), Zoology 47: 187-239. —— 1986. The pharyngeal apophysis on the base of the skull in cichlid fishes. A reply to Trewavas (1985). Netherlands Journal of Zoology 36 (4): 562-4. —— 1987. The genera of pelmatochromine fishes (Teleostei, Cichlidae). A phylogenetic review. Bulletin of the British Museum (Natural History), Zoology 53: 139-203. —— 1989. The taxonomic status and phylogenetic relationships of Pseudocre- nilabrus Fowler (Teleostei, Cichlidae). Ichthyological Bulletin of the J.L.B. Smith Institute of Ichthyology, Grahamstown No. 54: 1-16. — 1992. A revision and redescription of the monotypic cichlid genus Pharyngochromis (Teleostei, Labroidei). Bulletin of the British Museum (Natural History), Zoology 58: 37-52. Hert, E. 1989. The function of egg-spots in an African mouthbrooding cichlid fish. Animal Behaviour 37: 726-732. Hoogerhoud, R.J.C. 1986. Taxonomic and ecological aspects of morphological plasticity in molluscivorous haplochromines (Pisces, Cichlidae). Musee Royal de L’ Afrique Centrale, Tervuren; Annales Sciences Zoologique 251: 131-4. Jubb, R.A. 1967. The Freshwater Fishes of Southern Africa vii + 248 pp. Balkema, Cape Town. — 1968. A new Chetia (Pisces, Cichlidae) from the Incomati River system, Eastern Transvaal, South Africa. Annals of the Cape Provincial Museums (Natural History) 6 (7): 71-76. Konings, A. 1991. Cichlid and all other fishes of Lake Malawi 495 pp. T.F.H. Publications, New Jersey. Ladiges, W. 1964. Beitrage sur Zoogeographie und Oekologie der Stisser- wasserfische Angolas. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut 61: 221-272. Liem, K.F. 1991. Functional morphology. pp. 129-150. Jn: Keenleyside, M. (Ed.) Cichlid Fishes. Chapman & Hall, London. Lippitsch, E. 1991. Comparative investigation on scale characters in cichlids. Musee Royal de L’Afrique Centrale, Tervuren, Annales Sciences Zoologique 263: 97-102. Oliver, M.K. 1984. Systematics of African cichlid fishes: determination of the most primitive taxon, and studies of the haplochromines of Lake Malawi (Teleostei, Cichlidae). Ph.D. Thesis, Yale University. Poll, M. 1967. Contribution a la faune ichthyologique de l’Angola. Museu do Dundo, Publicacoes Culturais No. 75: 1-381. 1986. Classification des Cichlidae du lac Tanganika. Tribus, genres et especes. Académie Royale de Belgique; Mémoires de la Classe des Sciences, collection in 8°, 2e series, 45 (2): 1-163. Regan, C.T. 1920. The classification of the fishes of the family Cichlidae. I The Tanganyika genera. Annals and Magazine of Natural History (9)8: 632-639. —— 1922. The classification of the fishes of the family. Cichlidae. II. On African and Syrian genera not restricted to the Great Lakes. Annals and Magazine of Natural History (9)10: 249-264. Skelton, P.H. 1993. A complete Guide to the Freshwater Fishes of Southern Africa. Southern Book Publishers. Halfway House, South Africa. (In press). Stiassny, M.L.J. 1989. A taxonomic revision of the African genus Tylochromis (Labroidei, Cichlidae); with notes on the anatomy and relationships of the group. Musee Royal de l’Afrique Centrale, Tervuren; Annales Sciences Zoologiques 258: 1-161. — 1990. Tylochromis, relationships and the phylogenetic status of the African Cichlidae. American Museum Novitates no. 2993: 1-14. Trewavas, E. 1961. A new cichlid fish in the Limpopo basin. Annals of the South African Museum 46(5): 53-56. 44 —— 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 SLL IE tay EE cil LAM NG 9 i Doc yg | 6 ( anim f ne wey fe — 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 49 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 50 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 ie So Be, hy ) ‘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. NY Y i I uw v/a o. } = Y, < 7 \ \ ri eA 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. F R. HUYS AND J.M. GEE = yy Gif —— ~ AF y Lbs 5 Se ; i 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, i 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.] 66 labled VERN YER < > fag oo SS 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 LY | EST \ M@ \\ VXs Nf a . S tH 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: (s . | owed). tl dopo r d). ma 77 REVISION OF DANIELSSENIA AND PSAMMIS = > S—S 4 +} 4 i “5 > ee eee Ee = SS 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. 78 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 79 ( ¢ A 4A LNeone: a a f e d b Cc d 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: 80 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 REFERENCES Arlt, G. 1983. Taxonomy and ecology of some harpacticoids (Crustacea, Copepoda) in the Baltic Sea and Kattegat. Zoologische Jahrbiicher Systema- tik. Abteilung fiir Systematik 110: 45-85. Becker, K.-H. 1974. Eidonomie und Taxonomie abyssaler Harpacticoida (Crustacea, Copepoda). Teil I. Cerviniidae - Ameiridae. ’‘Meteor’- Forschungs-Ergebnisse Reihe D 18: 1-28. 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Novye massovye formy garpaktitsid (Copepoda, Harpac- ticoida) iz zaliva Pos’eta Yaponskogo morya [New common forms of harpacticids (Copepoda, Harpacticoida) from Possjet Bay of the Sea of Japan]. Issledovaniya Fauny More[ 8(14): 151-181. [In Russian. ] Coker, R.E. 1938. A copepod ‘Intersex’. Travaux de la Station zoologique de Wimereux 13: 97-100. Conover, R.J. 1965. An intersex in Calanus hyperboreus. Crustaceana 8:153-158. Francois, Y. 1949. Quelques cas d’intersexualité chez Eudiaptomus gracilis (Sars) (Copépode Calanoide). Etude morphologique. Bulletin de la Société zoologique de France 74: 232-239. Gee, J.M. 1988a. Some harpacticoid copepods (Crustacea) of the family Tachidiidae from sunlittoral soft sediments in Norway, the Celtic Sea and Gulf of Mexico. Zoologica Scripta 17: 181-194. —— 1988b. Taxonomic studies on Danielssenia (Crustacea, Copepoda, Har- pacticoida) with descriptions of two new species from Norway and Alaska. Zoologica Scripta 17: 39-53. — & Huys, R. 1990. 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(Copepoda, Harpacticoida) with a review of the taxonomic status of certain other deepwater harpacticoids. Hydrobiologia 185: 101-126. Ianora, A., Mazzocchi, M.G. & Scotto di Carlo, B. 1987. Impact of parasitism and intersexuality on Mediterranean populations of Paracalanus parvus Copepoda, Calanoida). Diseases of Aquatic Organisms 3: 29-36. Jespersen, P. 1939. Investigations on the copepod fauna in East Greenland waters. Meddelelser om Gronland 119 (9): 1-106. Klie, W. 1939. Diagnosen neuer Harpacticoiden aus den Gewassern um Island. Zoologischer Anzeiger 126: 223-226. — 1941. Marine Harpacticoiden von Island. Kieler Meeresforschungen 5: 1-44. — 1944. Ein gynandromorpher Amphiascus (Cop. Harp.) von Helgoland. Zoologischer Anzeiger 145: 77-79. Lang, K. 1936a. Undersokningar 6ver Oresund. Untersuchungen aus dem Oresund. XX. Harpacticiden aus dem Oresund. Acta Universitatis Lunden- sis.,n. ser., Avd. 2, 31: 1-52. — 1936b. Die wahrend der schwedischen Expedition nach Spitzbergen 1898 und nach Grénland 1899 eingesammelten Harpacticiden. Kungliga Svenska Vetenskapsakademiens Handlingar (3)15(4): 1-55. — 1944. Monographie der Harpacticiden (Vorldufige Mitteilung). 39 pp. Almgvist & Wiksell, Uppsala. REVISION OF DANIELSSENIA AND PSAMMIS — 1948. Monographie der Harpacticiden, volume I. 1-896, volume II. 897-1682. Hakan Ohlsson, Lund. Moore, C.G. & Stevenson, J.M. 1991. The occurrence of intersexuality in harpacticoid copepods and its relationship with pollution. Marine Pollution Bulletin 22: 72-74. Mrazek, A. 1913. Androgyne Erscheinungen bei Cyclops gigas Cls. Zoologi- scher Anzeiger 43: 245-250. Norman, A.M. & Scott, T. 1906. The Crustacea of Devon and Cornwall. i-xv + 232 pp. William Wesley and Son, London. Pirocchi, L. 1940. Eine Welle von Missbildungen in einer Bevolkerung von Arctodiaptomus bacillifer Koelb. (Crust. Copep.). Zoologischer Anzeiger 129: 269-271. Por, F. 1964. A study of the Levantine and Pontic Harpacticoida (Crustacea, Copepoda). Zoologische Verhandelingen. Leiden 64: 1-128. — 1965. Harpacticoida (Crustacea, Copepoda) from muddy bottoms near Bergen. Sarsia 21: 1-16. Sars, G.O. 1898. The Cladocera, Copepoda and Ostracoda of the Jana expedition. Ezhegodnik Zoologicheskago Muzeya Imperatorskoi Akademii Nauk 3: 324-359. — 1910. Copepoda Harpacticoida. Parts XXIX & XXX. Tachidiidae (con- cluded), Metidae, Balaenophilidae, supplement (part). An account of the Crustacea of Norway, with short descriptions and figures of all the species 5: 337-368. — 1921. Copepoda supplement. Parts IX & X. Harpacticoida (concluded), Cyclopoida. An account of the Crustacea of Norway, with short descriptions and figures of all the species 7: 93-121. Shen, C.-j. & Bai, S.-o. 1956. The marine Copepoda from the spawning ground of Pneumatophorus japonicus (Houttuyn) off Chefoo, China. Acta Zoologica Sinica 8: 177-234. [In Chinese with English summary. ] Smirnov, S.S. 1946. Novye vidy Copepoda Harpacticoida iz severnogo ledovi- 81 togo okeana. [New species of Copepoda-Harpacticoida from the Arctic Ocean). Trudy Dreifuyuschei Ekspeditsii Glavsevmorputi na Ledokol’nom Parokhode ‘G. Sedov’ 1937-1940 gg 3: 231-263. [In Russian with English summary.] Soyer, J. 1970. Contribution a l'étude des Copépodes Harpacticoides de Méditerranée Occidentale. 2. Tachidiidae Sars, Lang. Vie et Milieu 21 (2A): 261-277. Thompson, I.C. 1893. Revised report on the Copepoda of Liverpool Bay. Proceedings and Transactions of the Liverpool Biological Society 7: 175-230. Veldre, I. & Maemets, A. 1956. Eesti NSV_ vabaltevad aerjalalised (Eucopepoda) II. Soudikulised (Cyclopoida), Rullikulised (Harpacticoida). [Die freilebenden Ruderfiissler der estnischen SSR. II. Cycoploida, Harpac- ticoida]. Abiks Loodusevaat. 29: 1-128. [In Estonian. ] Wells, J.B.J. 1965. Copepoda (Crustacea) from the meiobenthos of some Scottish marine sub-littoral muds. Proceedings of the Royal Society of Edinburgh 69 (B): 1-33. 1967. The littoral Copepoda (Crustacea) of Inhaca Island, Mozambique. Transactions of the Royal Society of Edinburgh 67: 189-358. 1968. New and rare Copepoda Harpacticoida from the Isles of Scilly. Journal of Natural History 2: 397-424. Willey, A. 1920. Report on the marine Copepoda collected during the Canadian Arctic expedition. Report of the Canadian Arctic Expedition 1913-1918 7(K): 1-46. Wilson, M.S. 1966. North American harpacticoid copepods, 8. The Danielsse- nia sibirica group, with description of D. stefanssoni Willey from Alaska. Pacific Science 20: 435-444. Yashnov, V.A. 1935. Fauna solonovatovodnykh vodoemovy ostrova Vrangelya. {Fauna of brackish-water basins of the Wrangel Island]. /ssledovaniya More SSSR 22: 119-134. [In Russian with English summary. | 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 SELON CMO Fcc cn is oisieas ads hice dgode vu toon aclas cam MMM RTA asRces Sete Oaremsescedeenoes reece senses soccer sete ne Mer cnek etter cetera eT 83 MPATEIAISANGIMEtHOOS? 0.0.5 .ncsancandscosessceretentcmmieeemesacceetersncs se tnaeoceeee eee sce tasccen sect ar cmiee rasan case te rnanaecceett 83 PIES IE ON Strarac ae scaascuanesat iawanrs asincessdecacdaspamacan sera: Gc o0 ate ate ese esateee cnt ctaae dace Roce name = Aah e tam een e seen. 84 BAM © VMNGTOPSVINGAG SAS, VOUS” iio cces coseacas on snsesacctetaveonssdencmereaeteracaencstcectnaes date secon cuceetedcoveeete 84 Sublanilviveptopontiinae Wang, L948: .ccicencsscenls sucnnciatee eecadanan ote tate teers: ataaen cements dice oteerdaleeas tater as 84 Aurcolm Willies Ge. Clacy ss 1982 es. 2. ising BOM as SE «nnn an SERA aad Se etn a natin eave» « leaketeies Geena eealp icv weidaanls 85 SUZICOMPIMICTINEGIUS SP. DOV 50 o5x oss ios sacasassercsacsanccuossnsMemsene ae uas = eueeads an < calbw sige once wi tos toe aac s oeeei a mee 85 SyricOlanlanaricus Willetmistée'Clacys; L982 Bi IIIS, «0... /tteaewaass' dae eap oaddds cuews tents acne stan cases uaaeew ste uses 92 BREMIANKS ie ante seereatoesecs frank ota. tanta ae dice Ateuzewscuetebaacs onannnuge da dadusadaep aves ode otis sjant’slepinnastainametlyoaabaigsinea anes 92 PENIS EUISSIO LI me cae strcls tiscavin inti oss tnt ctonisc' slaiscinc esinal/otaie’naentlsino lls oine es oa a iaiNy abit ciara ole = = Se é nj 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 co NO 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 oO aoe OO mR ke L- t+ + & S = + K + + oO | (Ss) | ” 30 | O 28 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 ss 3 > ea = ies ls li Cc 5S 344 + se 4 KR ++ 2 o 32) as + 3 | ae ” | 30 4 <8 oO (lType of H. stewarti O 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. REFERENCES Ahmed, S. 1975. Sea-snakes of the Indian Ocean in the collections of the Zoological Survey of India together with remarks on the geographical distribution of all Indian Ocean species. Journal of Marine Biological Association of India 17:73-81. Anderson, J. 1872. On some Persian, Himalayan, and other reptiles. Proceed- ings of the Scientific Meetings of the Zoological Society of London, 1872:399. Boulenger, G. A. 1896. Catalogue of the British Museum (Natural History) Serpentes 3. British Museum, London. Bussarawit, S., Rasmussen, A. R. & Andersen, M. 1989. A preliminary study on sea snakes (Hydrophiidae) from Phuket harbour, Phuket Island, Thai- 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 4 72 Persian Gulf and Gulf of Oman 70 T T = T T T 1 162 166 170 174 178 182 186 190 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 + * + (+ © lS ae Ae ay 82 a5 Ap Se jae | Ox*+4+ 4+ 80 - + i 78 - 7668 170 172 174 176 178 180 182 184 186 188 Number of vertebrae Fig. 7 Relation between number of body vertebrae and position of heart tip in males (top) and females (bottom) of H. lapemoides, showing geographic variation. land. Natural History Bulletin of Siam Society 37:209-225. Cogger, H. G. 1975. Sea snakes of Australia and New Guinea pp. 59-139 In: W. A. Dunson (Ed.) Biology of Sea Snakes. University Park Press, Baltimore, London & Tokyo. De Silva P. H. D. H. 1980. Snake Fauna of Sri Lanka. National Museum of Sri Lanka, Colombo, Sri Lanka. Dowling, H. G., & Savage, J. M. 1960. A guide to the snake hemipenis: A survey of basic structure and systematic characteristics. Zoologica 45:17—-28. Dunson, W. A., & Minton, S. A. 1978. Diversity, distribution, and ecology of Philippine marine snakes (Reptilia, Serpentes). Journal of Herpetology 12:281-286. Gasperetti, J. 1988. Snakes of Arabia. /n: Buttiker, W. and F. Krupp (Eds.), Fauna of Saudi Arabia. 9:298-326. THE STATUS OF HYDROPHIS LAPEMOIDES Gray, J. E. 1849. Catalogue of the specimens of snakes in the collection of the British Museum, London. Giinther, A. 1872. New species of snakes in the collection of the British Museum. Annals and Magazine of Natural History, January 1872:33. Kharin, V. E. 1983. A new species of the genus Hydrophis sensu lato (Serpentes, Hydrophiidae) from the north Australian shelf. Zoologicheskii Zhurnal LXII:1751-1753. — 1984a. Sea snakes of the genus Hydrophis sensu lato (Serpentes, Hydrophiidae). On the taxonomic status of the New Guinea H. obscurus. Zoologicheskii Zhurnal LXIII:630-632. — 1984b. A review of sea snakes of the group Hydrophis sensu lato (Serpentes, Hydrophiidae). S. the genus Leioselasma. Zoologicheskii Zhur- nal LXIII:1535-1546. Lemen, C. A., & Voris, H. K. 1981. A comparison of reproductive strategies among marine snakes. Journal of Animal Ecology. 50:89-101. Leviton, A. E., Gibbs, R. H., Heal, E. & Dawson, C. E. 1985. Standards in Herpetology and Ichthyology: Part I. Standard Symbolic Codes for Institu- tional Resource Collections in Herpetology and Ichthyology. Copeia 1985:802-832. McCarthy, C., & Warrell, D. 1991. A collection of sea snakes from Thailand with new records of Hydrophis belcheri Gray. The Bulletin of the British Museum Natural History (Zoology Series) 57:161—166. McDowell, S. B. 1972. The genera of sea-snakes of the Hydrophis group (Serpentes: Elapidae). Transactions of the Zoological Society of London 32:189-247. Minton, S. A. 1966. A contribution to the herpetology of West Pakistan. Bulletin of the American Museum of Natural History 134:27-184. Murthy, T. S. N. 1985. Classification and distribution of the reptiles of India. The Snake 17:48-71. Rasmussen, A. R. 1987. Persian Gulf Sea Snake Hydrophis lapemoides (Gray): New record from Phuket Island, Andaman Sea, and the southern part of the Straits of Malacca. Natural History Bulletin of the Siam Society 35:57-S8. — 1989. An analysis of Hydrophis ornatus (Gray), H.lamberti Smith, and H.inornatus (Gray) (Hydrophiidae, Serpentes) based on samples from various localities, with remarks on feeding and breeding biology of H.orna- tus. Amphibia-Reptilia 10:397-417. — 1992. Rediscovery and redescription of Hydrophis bituberculatus Peters, 1872 (Serpentes, Hydrophiidae). Herpetologica 48(1):85—97. —— in press. A cladistic analysis of the Hydrophis subgenus Chitulia (Ser- pentes, Hydrophiidae). Zoological Journal of the Linnean Society. Smith, L. A. 1986. A new species of Hydrophis (Serpentes: Hydrophiidae) 105 from north-west Australian waters. Records of the Western Australian Museum 13:151-153. Smith, M. 1926. Monograph of the Sea-snakes (Hydrophiidae). British Museum (Natural History), London. — 1930. The Reptilia and Amphibia of the Malay Peninsula. Bulletin of the Raffles Museum Singapore, Straits Settlements. 3:74-83. — 1935. The sea snakes (Hydrophiidae). Dana-Report 8:1-6. — 1943. The fauna of British India Ceylon and Burma. Reptilia and Amphibia Vol. Ill — Serpentes. London. Tamiya, N., Maeda, N. & Cogger, H. G. 1983. Neurotoxins from the venoms of the sea snakes Hydrophis ornatus and Hydrophis lapemoides. Biochemical Journal 213:31-38. Taylor, E.H. 1965. The serpents of Thailand and adjacent waters. University of Kansas Science Bulletin 45:609-1096. Thomas, R. A. 1976. Dorsal scale row formulae in snakes. Copeia 1976:839-841. Toriba, M., & Sawai, Y. 1981. New record of Persian Gulf sea-snakes Hydrophis lapemoides (Gray) from Penang, Malaysia. The Snake 13:134-136. Tweedie, M. W. F. 1983. The Snakes of Malaya. 3rd ed. Singapore National Printers, Singapore. Volsge, H. 1939. The sea-snakes of the Iranian Gulf and the Gulf of Oman. Danish Scientific Investigations in Iran, 1:9-45, Voris, H. K. 1975. Dermal scale-vertebra relationship in sea snakes (Hydrophi- idae). Copeia 1975:746-757. —— 1977. A phylogeny of the sea snakes (Hydrophiidae). Fieldiana, Zoology 70:79-166. ., & Voris, H. H. 1983. Feeding strategies in marine snakes: An analysis of evolutionary, morphological, behavioral and ecological relationships. American Zoologist. 23:411-425. Wall, F. 1909. A monograph of the sea snakes. Memoirs of the Asiatic Society of Bengal 2:169-251. Williams, K. L., & Wallach, V. 1989. Snakes of the world Vol.1. Synopsis of Snake Generic Names. Krieger Publishing Company Malabar, Florida. Zann, L. P. 1975. Biology of a barnacle (Platylepas ophiophilus Lanchester) symbiotic with sea snakes pp.267—286 Jn: W. A. Dunson (Ed.) Biology of Sea Snakes. University Park Press, Baltimore, London & Tokyo. -, Cuffey, R. J., & Kropach, C. 1975. Fouling organisms and parasites associated with the skin of sea snakes pp. 251-265 Jn: W. A. Dunson (Ed.), Biology of Sea Snakes. University Park Press, Baltimore, London & Tokyo. ' * we eres ian 4 SOV) cape "| Daoww sad heigl a | Lee A Jipeecild t % ,wreda i ii ery ytd AL ered ote. ft ee ee A sw ,? sai 7 MELT? O22 Mew4(4 ; pai seer toe) - rd sindiel a Ww eit eR Sa oe ee | Pr ll Gla a) a A rw 1Gec> ma Pe cts A ' parry |) at.) by? nae ~ ~ > ‘ A >. & ie “soa y “ a , _—a ; —— & 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. © The Natural History Museum, 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 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. Z a O cal = o ‘e) O 5 = = = cs < ~% jaa =| & = < e4 eo) ee fh, eo) Zz fe) Z > ea] a4 = = Oo Z eo) s% < = 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. Zz 2 = O jaa 4 = fe) O z — = = & < m aa) = g = < a4 e) — Ky fo) Z fe) Zi > si % 2 = fe) Z fe) x < al 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. REFERENCES Acosta, J.T. 1940. Nuevos foraminiferos de la costa sur de Cuba. Memorias de la Sociedad Cubana de Historia Natural ’Felipe Poey’, Havana, 14: 269-276. Andersen, H.V.1951a. two new genera of foraminifera from Recent deposits of Louisiana. Journal of Paleontology, Tulsa, 25: 31-34. — 1951b. An addenda to Arenoparrella and Arenoparrella mexicana (Korn- feld). Contributions from the Cushman Foundation for Foraminiferal Research, Washington, 2: 96-97. — 1953. Two new species of Haplophragmoides from the Louisiana coast. 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Antiquariaat Junk, Lochem [Reprint]. Hoglund, H. 1947. Foraminifera in the Gullmar Fjord and the Skagerak. Zoologiska Bidrag fran Uppsala, 26: 1-327. Kornfeld, M.M. 1931. Recent littoral foraminifera from Texas and Louisiana. Contributions from the Department of Geology, Stanford University, Palo Alto, 2: 77-101. Lacroix, E. 1932. Discammina, nouveau genre Méditerranéen de foraminiféres arénacés. Bulletin de l'Institut Océanographique de Monaco, 600: 1-4. Loeblich, A.R. Jr. & Tappan, H. 1953. Studies of Arctic foraminifera. Smithsonian Miscellaneous Collections, Washington, 121 (7): 1-150. — & 1964. Treatise on Invertebrate Paleontology. Part C, Protista 2, Sarcodina, chiefly ‘Thecamoebians’ and Foraminiferida. 2 Volumes. xxxi+ 900pp. Geological Society of America and the University of Kansas Press. — & — 1987. Foraminiferal genera and their classification. 1: 1-970. 2: 1-212. Van Nostrand Reinhold Company, New York. Lutze, G.F. 1968. 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Zaninetti, L., Bronnimann, P., Beurlen, G. & Moura, J.A. 1977. La mangrove de Guaratiba et la Baie de Sepetiba, Etat de Rio de Janeiro, Brésil: Foraminiféres et écologie. Archives des Sciences, Genéve, 30: 161-178. 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 a Veale hed UL CNA fe te oe RR eels ocginile wee Vee onis's sis sOEVBRG Mela vin o¥iaichin sais alvanaisasiqeSie eas avis aiiecinssins seine civaaen ane 126 Sy es UTA AC COAL Vests te water a se Siete MERU se volte Sula odGs sis aio sie ani drtve ceWclapid paialele epilelasluSeinige de on sinjoasnaigae cup\einis smecie sdlevenceie 126 Functional morphology of the digestive system and feeding mechanisms in the Conoidea ......................00665 145 ea teRTCTESINN S(O 100 © TL OLCC ele ale eee a noc osslec ec ebadlo vin wn so cites cdeeanmacenth teideetharetecetwanest detsrerete servereditenent se 150 Glassiication OMGonoidearamd MiagnOSES OP NISMEL TAKA ....cccceccecescccnccaccccesceccscsccececcestrcenaechssescsenecaeeens 156 SGML CMC UES st MMMM TeaENCE REeMAG. dale ceieccassaiedensioess.necestestessececrastaccattrcetecceatdaecesccseceerercacceeceessnse 161 Appendix 1. Description of shell characters used in phylogenetic amalysis. ................0ccececnceececneneceeeenenenes 162 NONE U CIN © LASSINCAIOMOM NCCOLU MEIC PCHOTA ..cccsscsnenctecsededscocnstcscstevcsteerscescasecscasoctscnsctcatudvens ceeds 163 ESGRETENCES Mle Lente ren etaaema a atdce aoa eit PnAet a ecasi nieces dacnaiadieas Uivnlevadcscashbtavead ties talssanicodiuladvel srt cetotssine 168 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 © The Natural History Museum, 1993 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 7 ee SS \\ yo > ait, WPAN QP 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 Yf Wi 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 = wW e) 2) n > < Q Z < [og (S) i : > [a4 e) —] ral < lol a = 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 B 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 — present 0 — absent, 1 — one sphincter, 2 — two sphincters 0 — absent, 1 — present 0 — absent, 1 — present 0 — absent, 1 — present 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 NOGOOONFKF CORP CCOCCOCOFRPCOCOORPRFRP FPP NFP OOCOROCOrRRKROCOONOCOCOCOrFS VSS) SSS SS) OOS SSeS Sessa ooo eSs See oT oee a8 oae]) FPOoOrOrcOoOrRrPrRPrRP COCCCCCoCoCoCOoOCcCOCCCCocoocooooococoocoocoocoocoeoo orf orcoocoorFr S| OOo OOO CoOOO COR eo So ooo 0 oo So oo So oO ooo 'S : > WR VONMNOOO NV VON ON RP RPO NI VR REN RP RP RP EP N RP NR RFP ONKF PRE NOC CO 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 > > Sot RP VRP PP ya RP Se PRP RP RP Pree erererPocooocoooococoocoeocoeco VWV VV VV VV VV VV VV VV VV YY VY VV NV VY VV VNN VV VNNNR RV O NY ec ec ee eee ce a ee ee ee ee ee ee aan i a aa) 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) SS ROO EA IS StS Se OOS Oo OSS Soa ooo oF So ool eo SS Sis S Wor TO YNOOVVYVYNVYNYMR PrP OCOrcocooocoococooocoococoo'yon7sd ee oe on an an an an an an an an en en a> eo an eo ao em) SOR VV VV VV VV VV VV VV VV VV VRRP ROO VIO ORF RP ORCOrROSO 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 eoooqcoooqcooormrreooCorrerocoooceocooocoocoeooeoseoooooss NNR ROR PRO ORR RRP RP RP RP RRP OR RP NDR RP RB RN EP NNNE PNR RP RP RROD ooonor SOY HSE rH Oe HPrPRPrereoOooocoeoescoooeeocoooos © 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 eSeoeooooororrPrPooCooooOrFOOCOORrR OR OR OR OR COR RP Re ROR HEHE OWN SOOCONVNNNYVYNNVYNNNNNNNNNNNNNRP RPE RRP REP NRFPRRFPOOCOCCCSO 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) 15 16 34(0), 37(2) 17 2(1), 38(2), 40(2) 18 22(1), 25(2), 32(1) 19 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 3 G 6 2 9 10 5 16 Zz 15 19 22 18 26 25 28 30 29 32 33 35 36 0 23 34 37 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 in some Present in some Present in some Present 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 (© %hugae Kuroda, 1953) Lirasyrinx Powell, 1942 (= 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 oa 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) REFERENCES Adams, H. & Adams, A. 1853. The genera of Recent Mollusca Volume 1. John van Voorst, London Andrews, E.B. 1991. The fine structure and function of the salivary glands of the dogwhelk Nucella lapillus (Gastropoda: Muricidae). Journal of Mollus- can Studies 57: 111-126. Auffenberg, K. & Lee, H.G. 1988. A new species of intertidal Terebra from Brazil. Nautilus 102: 154-158. Bandel, K. 1984. The radulae of Caribbean and other Mesogastropoda and Neogastropoda. Zoologische Verhandelingen 214: 1-188. Bogdanoy, I.P. 1985. 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Annals of the Natal Museum 27: 633-720. — 1988. Turridae (Mollusca: Gastropoda) of southern Africa and Mozam- bique. Part 4. Subfamilies Drilliinae, Crassispirinae and Strictispirinac. Annals of the Natal Museum 29: 167-320. 1989. Notes on Prychobela and Brachytoma, with the description of a new species from Mozambique (Mollusca: Gastropoda: Turridae). Annals of the Natal Museum 30: 185-196. Kohn, A.J. 1959. The ecology of Conus in Hawaii. Ecological Monographs 29: 47-90. Leviten, P.J. 1970. The structure and function of the alimentary system of Polystira albida (Gastropoda: Toxoglossa). M.Sc. Thesis, University of Miami. Maes, V.O. 1983. Observations on the systematics and biology of a turrid assemblage in the British Virgin Islands. Bulletin of Marine Science 33: 305-335. Marincovich, L. 1973. Intertidal molluscs of Iquique, Chile. Science Bulletin. Natural History Museum, Los Angeles County 16: 1-49. Marcus, E. & Marcus, E. 1960 On Hastula cinerea. Boletim da Faculdade de Filosofia, Ciencias e Letras. Universidade de Sao Paulo 23: 25-66. Marsh, H. 1970. Some preliminary studies of the venoms of some vermivorous Conidae. Toxicon 8: 271-277. 1971. The foregut glands of vermivorous cone shells. Australian Journal of Zoology 19: 313-326. McLean, J.H. 1971. A revised classification of the family Turridae, with the proposal of new subfamilies, genera, and subgenera from the eastern Pacific. Veliger 14: 114-130. Miller, B.A. 1970. Studies on the biology of some Indo-Pacific Terebridae. Ph.D.Thesis, University of New Hampshire, Dover. 213 pp. — 1971. Feeding mechanisms of the family Terebridae. Reports of the American Malacological Union Pacific Division 1970: 72-74. —— 1975. The biology of Terebra gouldi Deshayes and a discussion of life history similarities among other terebrids of similar proboscis type. Pacific Science 29: 227-241. — 1980. The biology of Hastula inconstans (Hinds, 1844) and a discussion of 170 life history similarities among other hastulas of similar proboscis type. Pacific Science 33: 289-306. Miller, J.A. 1989. The toxoglossan proboscis: structure and function. Journal of Molluscan Studies 55: 167-182. — 1990. The feeding and prey capture mechanism of Turricula nelliae spurius (Hedley) (Gastropoda: Turridae). pp. 979-992. In: Morton, B. (Ed.) Proceedings of the Second International Marine Biological Workshop: The Marine Flora and Fauna of Hong Kong and Southern China, Hong Kong, 1986. Hong Kong University Press, Hong Kong. Mills, P.M. 1977. On the venom gland of terebrid molluscs. Proceedings of the Third International Coral Reef Symposium, Miami, Florida. , 631-637. Morrison, J.P.E. 1966. On the families of Turridae. Annual Report of the American Malacological Union for 1965.: 1-2. Nybakken, J. 1990. Ontogenetic change in the Conus radula, its form, distribution among radula types, and significance in systematics and ecology. Malacologia 32: 35-54. Olivera, B.M., Rivier, J., Clark, C., Ramilo, C.A., Corpuz, G.P., Abogadie, F.C., Mena, E.E., Woodward, S.R., Hillyard, D.R. & Cruz, L.J. 1990. Diversity of Conus neuropeptides. Science 249: 257-263. Pace, S. 1901. On the anatomy of the prosobranch genus Pontiothauma, E.A. Smith. Zoological Journal of the Linnean Society 28: 455-462. Pearce, J.B. 1966. On Lora treveliana (Turton) (Gastropoda: Turridae). Ophelia 3: 81-91. Ponder, W.F. 1970. Some aspects of the morphology of four species of the neogastropod family Marginellidae with a discussion of the evolution of the toxoglossan poison gland. Journal of the Malacological Society of Australia 2: 55-81. : — 1973. Origin and evolution of the Neogastropoda. Mala-cologia 12: 295-338. — & Warén, A. 1988. Classification of the Caenogastropoda and Heterostro- pha — a list of the family-group names and higher taxa. Malacological Review Supplement 4: 288-326. Powell, A.W.B. 1942. The New Zealand Recent and fossil Mollusca of the Family Turridae. With general notes on turrid nomenclature and systemat- ics. Bulletin of the Auckland Institute and Museum 2: 1-192. — 1966. The molluscan families Speightiidae and Turridae. An evaluation of the valid taxa, both Recent and fossil, with lists of characteristic species. Bulletin of the Auckland Institute and Museum 5: 1-184. Robinson, E. 1960. Observations on the toxoglossan gastropod Mangelia brachystoma (Philippi). Proceedings of the Zoologi-cal Society of London 135: 319-338. Rudman, W.B. 1969. Observations on Pervicacia tristis (Deshayes, 1859) and a comparison with other toxoglossan families. Veliger 12: 53-64. Saunders, P.R. & Wolfson, F. 1961. Food and feeding behavior in Conus californicus Hinds, 1844. Veliger 3: 73-76. Schultz, M.C. 1983. A correlated light and electron microscopic study of the structure and secretory activity of the accessory salivary glands of the marine gastropods, Conus flavidus and Conus vexillum (Neogastropoda, Conidae). Journal of Morphology 176: 89-111. Sheridan, R., Van Mol, J.J., & Bouillon, J. 1973. Etude morphologique du tube digestif de quelques Turridae de la region de Roscoff. Cahiers de Biologie Marine 14: 159-188. Shimek, R.L. 1975. The morphology of the buccal apparatus of Oenopota levidensis (Gastropoda, Turridae). Zeitschrift fur Morphologie de Tiere 80: 59-96. J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV — 1983a. Biology of the northeastern Pacific Turridae. I. Ophiodermella. Malacologia, 23: 281-312. — 1983b. The biology of the northeastern Pacific Turridae. II. Oenopota. Journal of Molluscan Studies, 49: 146-163. — 1983c. The biology of the northeastern Pacific Turridae. III. The habitat and diet of Kurtziella plumbea (Hinds, 1843). Veliger 26: 10-17. & Kohn, A.J. 1981. 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(Zool.) © The Natural History Museum, 1994 Zoology Series ISSN 0968 — 0470 Vol. 60, No. 1, pp. 1-104 The Natural History Museum Cromwell Road London SW7 5BD Issued 23 June 1994 Typeset by Ann Buchan (Typesetters), Middlesex 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 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 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 5 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- 7 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 8 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 pi 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 10 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 AN S NS SSS Shor oss SD 14 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). 12 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 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 23 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. 24 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 a) 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 26 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 i) x: ao) = S 6 63.7 3372 S7a* 19SIGE Sl S81 5 61/4 62.9 82.3 36.7 18.2 49.8 79 5 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- 51 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 We » ogee? BSc 0.01imm | 1 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 53 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. 54 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 56 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 ss ——— oe 2 3s ——_ S 1S) o es ° 5 5 = EEU_UAAZZZ™" a? 0 ——_——E SSS SSS 58 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 rp ( q- | / » +2 ; =3°0 | = St Sitka fs OF G YE a —~ \ oe NS ra KC) Pouttsle sane ft 60 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 61 62 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. 64 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 65 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 67 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; 68 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 oO 69 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. : i 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 71 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 72 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 WO 13 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. Nn ON RECENT SPECIES OF SPIRASERPULA REGENHARDT, 1961 iS Gif PE SN FA ew) 5 A Liza a: | Gaz Wr 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. 76 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 { { | J \ 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 | NX \ 4 i : NY : | \ \ \ G 0.01mm \\ \ " i \ \ N\\ (tS Wl \ \ " \ 4 C \ | D \ 78 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. 80 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 82 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). D 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- N Se Sy jem J 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 84 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 85 86 T.G. PILLAI AND H.A. TEN HOVE jo - mm = ih 2 a 3 5) 1 Vane Wes = “H { een i) L rife Nake CS \ fe i ay ” i & S; we 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. 87 88 T.G. PILLAI AND H.A. TEN HOVE R 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 89 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 90 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 91 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. 92 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 a SS SEN =cwiteire 94 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 95 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 99 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. REFERENCES Amoureux, L., F. Rullier & L. Fishelson, 1978. Systématique et Ecologie D’Annélides Polychétes de la Presqu’il du Sinai. Israel Journal of Zoology, 27: 57-163. Ben-Eliahu, M. N., and H. A. ten Hove, 1989. Redescription of Rhodopsis pusilla Bush, 1905, a little known but widely distributed species of Serpulidae (Polychaeta). Zoologica Scripta 18, 3: 381-395. Bianchi, S.N. 1981. Policheti Serpuloidei. Guide per il riconoscimento delle specie animali delle acque lagunari e costiere italiane. Consiglio Nazionale delle Ricerche, AQ/1/96: 1-182. Hove, H. A. ten. 1979. Different causes of mass occurrence in serpulids p.282-298, in: G. Larwood & B. R. Rosen (eds), 1979. Biology and systematics of colonial organisms. Syst. Ass. Spec. Vol.II, xxxv: 589pp. Hove, H. A. ten. 1984. Towards a phylogeny in serpulids (Annelida; Polycha- eta). Proceedings of the First International Polychaete Conference, Sydney, edited by P. A. Hutchings. Published by The Linnaean Society of New South Wales, 181-196. Hove, H. A. ten, and M. J. A. Jansen-Jacobs. 1984. A revision of the genus Crucigera (Polychaeta: Serpulidae); a proposed methodical approach to serpulids, with special reference to variation in Serpula and Hydroides. Proceedings of the First International Polychaete Conference, Sydney, edited by P. A. Hutchings, published by The Linnaean Society of New South Wales, 143-180. Hove, H. A. ten, and M. O. M. Aarts. 1986. The distribution of Serpulidae (Annelida Polychaeta) on the warm-temperate and tropical Eastern Atlantic shelf. Netherlands Institute for Sea Research Publication Series, 13: 34-35. Hove, H. A. ten, and Pillai, T. G. ——. A cladistic analysis of Spiraserpula Regenhardt, 1961 (Serpulidae, Polychaeta); characters and character states. (Paper read at the Fourth International Polychaete Conference, to be published). Jager, M. 1983. Serpulidae (Polychaeta sedentaria) aus der norddeutschen hoéheren Oberkreide — Systematik, Stratigraphie und Okologie. Geol. Jb., (A)68: 3-219. Jager, M. 1993. Danian Serpulidae and Spirorbidae from NE Belgium and SE Netherlands: K/T boundary extinction, survival, and origination patterns. Contr. Tert. Quatern. Geol. , 29(3—4): 73-137. Land, J. van der. 1987. Report on the CANCAP-Project for Marine Biological Research in the Canarian-Cape Verdean Region of the North Atlantic Ocean (1976-1986). Part I. List of Stations. Zodlogische Verhandelingen, Rijksmuseum van Natuurlijke Historie, Leiden, CANCAP Project Contribu- tion, 24; 1-94. Land, J. van der, and Sukarno, 1986. Theme IV Coral Reefs. Part one. R. V. Tyro and K. M. Samudera September — November 1984. The Snellius II Expedition. Progress Report. Royal Netherlands Academy of Arts and Sciences. Indonesian Institute of Sciences, 76 + 71 pp. Lommerzheim, A. 1979. Monographische Bearbeitung der Serpulidae (Poly- chaeta sedentaria) aus dem Cenoman (Oberkreide) am Stdwestrand des Minsterlander Beckens. Decheniana (Bonn), 132: 110-195. Pillai, T. G. 1993. A Review of some Cretaceous and Tertiary serpulid polychaetes of the genera Cementula and Spiraserpula Regenhardt, 1961, Laqueoserpula Lommerzheim, 1979 and Protectoconorca Jager, 1983. Pala- Ontologische Zeitschrift, Stuttgart, 67: 69-88. Regenhardt, H. 1961. Serpulidae (Polychaeta sedentaria) aus der Kreide Mitteleuropas, ihre Gkologische, taxonomische und stratigraphische Bewer- 104 tung. Mitteilungen aus dem Geologischen Staatsinstitut in Hamburg, 30: 5-115. Scrutton, S.T. 1975. Hydroid-Serpulid Symbiosis in the Mesozoic and Tertiary. Palaeontology, 18 (2): 255-274. Straughan, D. 1967. Marine Serpulidae (Annelida: Polychaeta) of eastern Queensland and New South Wales. Australian Journal-of Zoology, 15: 201-261. Uchida, H. 1978. Serpulid Tube Worms (Polychaeta.Sedentaria) from Japan T.G. PILLAI AND H.A. TEN HOVE with the Systematic Review of the Group. Bulletin of the Marine Park Research Station, 2: 1-98. Zibrowius, H. 1968. Etude morphologique, systématique et écologique, des Serpulidae (Annelida Polychaeta) de la région de Marseille. Recueil des Travaux de la Station Marine d’Endoume, Bulletin 43(59): 81-252. Zibrowius, H. 1972. Une espéce actuelle du genre Neomicrorbis Rovereto (Polychaeta Serpulidae) découverte dans l’étage bathyal aux Acores. Bull. Mus. Hist. Nat. Paris (3) 39, Zool. 33: 423-430. GUIDE FOR AUTHORS Policy. The Bulletin of the British Museum (Natural His- tory) Zoology, was established specifically to accommodate manuscripts relevant to the Collections in the Department of Zoology. 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Leg- ends, brief and precise, must indicate scale and explain symbols and letters. Reprints. 25 reprints will be provided free of charge per paper. Orders for additional reprints can be submitted to the publisher on the form provided with the proofs. Later orders cannot be accepted. 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 Zoology Series S)2 THE NATURAL HISTORY MUSEUM VOLUME 60 NUMBER 2 24 NOVEMBER 1994 The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum (Natural History)), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology. The Zoology Series is edited in the Museum’s Department of Zoology Keeper of Zoology: Dr C.R. Curds Editor of Bulletin: Dr N.R. Merrett Assistant Editor: Dr B.T. Clarke Papers in the Bulletin are primarily the results of research carried out on the unique and ever- growing collections of the Museum, both by the scientific staff and by specialists from elsewhere who make use of the Museum’s resources. Many of the papers are works of reference that will remain indispensable for years to come. All papers submitted for publication are subjected to external peer review for acceptance. A volume contains about 160 pages, made up by two numbers, published in the Spring and Autumn. Subscriptions may be placed for one or more of the series on an annual basis. Individual numbers and back numbers can be purchased and a Bulletin catalogue, by series, is available. Orders and enquiries should be sent to: Intercept Ltd. P.O. Box 716 Andover Hampshire SP10 1YG Telephone: (0264) 334748 Fax: (0264) 334058 Claims for non-receipt of issues of the Bulletin will be met free of charge if received by the Publisher within 6 months for the UK, and 9 months for the rest of the world. World List abbreviation: Bull. nat. Hist. Mus. Lond. (Zool.) © The Natural History Museum, 1994 Zoology Series ISSN 0968 — 0470 Vol. 60, No. 2, pp. 105-172 The Natural History Museum Cromwell Road London SW7 5BD Issued 24 November 1994 Typeset by Ann Buchan (Typesetters), Middlesex Printed in Great Britain at The Alden Press, Oxford 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- © The Natural History Museum, 1994 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. Crassarietellus 10 0 0 0 0 OO OO TOO TOO O00) 0 OO, ONO OG sl lO) OW, © 00 OD OD @ 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) MATERIAL EXAMINED. ©, Zoological Museum, University 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 Ak 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 a z ————— yy S yee iy —— \ 20 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 MATERIAL EXAMINED. ©’, allotype, The Natural History 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 3 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 ee ‘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 | y 4 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 y at ry / way Parrot, j ~ Oe ang aes Ancestor 161 Fa “OTN } seminal receptacle ay ====> 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. Iga) \o ml PHYLOGENY OF ARIETELLID COPEPODS “OSEIDYISIV INOYIIM JUSWBS SoJLOIPUI YSLI9}SY “JUSWITaS YORS UO dPJ_aS JO JOQUINU dy} So}BOIPUI JOquINU sy] “YPjadD]Iq ‘H ‘snuvnjpovjay ‘oD ‘snjudvsnvavg ‘4 ‘snjjaiaiuy ‘y ‘viaydosnuvavg *q ‘snjnaasojngg ‘> ‘snjjajaiavsavg *g ‘snjjajalavssvéy “Vy “e19U99 PI[[A}a1Ie JY} JO SojnuUaJUe a]eUOJ JO siNjeUIe pue susa}jed UOISNy [e}UOWZas Jo UONENSNI[I oNeWIaYIS “6E “BIA eee 2 el lllelteleleleltelelzlelele le lel = = ©] m pear oP Sic spapaeilengreucier tele (e Sieieteiele S86] © fee zie ecsltslellelleltelellelztelelle le telielelelellelle @ ¢ ¢| 4 * KK * KK KKK eee ¥ ei Js) G)@ ASPIRE eRe es 8 : * KK ree ¥ |i)s)AREEPEREEEEOCEEEE © q * OK * Fee =) s) EASIER © 3 * *K fe + Fie) s) )EIRIPIPIEIEEIEEPPIEREIIE s © e * KK Fee ¥ |i) EERE ERASE Av INAXX MAXX IAXX AXX AIXX WIXX MIXX IXX XX XIX IIAK WAX IAX AX AIX TNX =X «IX x xl IMA WA IA A A\ Ul iH | S. OHTSUKA, G.A. BOXSHALL AND H.S.J. ROE 164 I SSIEPSSS~S=SS REEL e “OSEJOYISOL OU YJIM JUIWBAS B SO}LIIPUT YSIS “JUOWIBas YORI UO 9eIIS JO JOQUINU dy) SoJeOIPUI JOQuINU dy] “‘snuD|vIODIA- *D ‘snjudvsnv.avg ‘4 ‘sapiojudvsnvang “q ‘snjjaiauy ‘q ‘viuydosnuvavg ‘dD ‘snjjajaiavsspdd *g Svldaunduy,) ‘VW *e19U98 pI[a}alIe dy} JO sanuUDjUe JjeW JO dINJeWLe pue suJo}ed UOISNY [e}USWIAs Jo UONRINSN][I ONeWIDYOS «Op “BIY eee lt oo REE Oey PIE += © OK VK eee el ol PEELE) 2 Peele ele OGRE EP EEE 5) 3 Peel PEE PEPER 5) * OK & Peer ae EEE EEE so Feel oo/ IER EELS & ee oe alt gp|pplBEE pCR a MAXX MAXX IAXX AXX AIXX IIXX IIXX IXX XX XIX INAX HAX IAX AX AIX INK IX IX x Xl A HA OIA A Al ul iT I 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 Pelagic Hyperbenthic Pelagic Hyperbenthic Hyperbenthic Fig. 45. Habitat cladogram of arietellid genera. Substitution of habitat type of each genus onto cladogram shown in Fig. 44. 4a 4b Sa Sb 6a 6b Ta 7b 8a 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). REFERENCES Alvarez, M.P.J. 1985. A new arietellid copepod (Crustacea): Pilarella longicor- nis, gen. n., sp. n., from the Brazilian continental shelf. Revista Brasileira de Zoologia, 28: 189-195. Andronovy, V.N. 1974. 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Phylogeny of calanoid copepods. Syllogeus, 58: 191-196. Roe, H.S.J. 1972. The vertical distributions and diurnal migrations of calanoid copepods collected on the SOND Cruise, 1965. 1. The total population and general discussion. Journal of the Marine Biological Association of the United Kingdom, 52: 277-314. Rose, M. 1933. Copépodes pélagiques. Faune de France 26: 1-374. Sars, G.O. 1901-1903. Copepoda. Calanoida. An account of the Crustacea of Norway 4. Bergen Museum, Bergen, 171 pp. — 1905. Liste preliminaire des Calanoidés recueillis pendant les campagnes de S.A.S. le Prince Albert de Monaco avec diagnoses des genres et des espéces nouvelles (1 re partie). Bulletin du Musée Océanographique de Monaco, 26: 1-22. — 1924, 1925. Copépodes particuli¢rement bathypélagiques provenant des campagnes scientifiques accomplies par le Prince Albert ler de Monaco. Résultats das Campagnes Scientifiques Prince Albert 1, 69: text (1925), 408 pp., atlas (1924), 127 pls. Scott, A. 1909. The Copepoda of the Siboga Expedition. Part I. Free- swimming, littoral and semi-parasitic Copepoda. Siboga Expeditie, 29a: 1-323, 69 pls. Scott, T. 1892. Additions to the fauna of the Firth of Forth. Part IV. Tenth Annual Report of the Fishery Board for Scotland 3: 244-272. 1894. Report on Entomostraca from the Gulf of Guinea. Transactions of the Linnnean Society of London, series 2, 6: 1-161, 15 pls. Vervoort, W. 1965. Pelagic Copepoda Part II. Copepoda Calanoida of the families Phaennidae up to and including Acartiidae, containing the descrip- tion of a new species of Aetideidae. Atlantide Report 8: 9-216. Wilson, C.B. 1950. Copepoda gathered by the U.S. Fisheries Steamer *Alba- tross’ from 1887-1909, chiefly in the Pacific Ocean. United States National Museum Bulletin 100, 14: 141-441, pls 2-36. Wolfenden, R.N. 1911. Die marinen Copepoden der deutschen Stidpolar- Expedition 1901-1903. II. Die pelagischen Copepoden der Westwinddrift und des stidlichen Eismeeres mit Beschreibung mehrerer neuer Arten aus dem Atlantischen Ozean. Deutsche Stidpolar-Expedition, 12 (Zoologie 4): 181-380, pls 22-40. GUIDE FOR AUTHORS Policy. The Bulletin of the British Museum (Natural His- tory) Zoology, was established specifically to accommodate manuscripts relevant to the Collections in the Department of Zoology. It provides an outlet for the publication of taxo- nomic papers which, because of their length, prove difficult to publish elsewhere. Preference is given to original contribu- tions in English whose contents are based on the Collections, or the description of specimens which are being donated to enhance them. Acceptance of manuscripts is at the discretion of the Editor, on the understanding that they have not been submitted or published elsewhere and become the copyright of the Trustees of the Natural History Museum. All submis- sions will be reviewed by at least two referees. 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