Contr. Tert. Quatern. Geol. 30(3-4) 155-175 7 figs, 1 tab, 3 pis. Leiden, December 1993

Kuphus melitensis a new teredinid bivalve from the , Late Oligocene Lower Coralline Limestone of Malta

George Zammit+ Maempel

Birkirkara, Republic of Malta

— Zammit Maempel, George. Kuphus melitensis, a new teredinid bivalve from the Late Oligocene Lower Coralline Limestone of Malta.

Contr. Tert. Quatern. Geol., 30(3-4): 155-175, 7 figs, 1 tab., 3 pis. Leiden, December 1993.

Coralline Anew species of teredinid bivalve, Kuphus melitensis, is described on the basis of valves, tubes and pallets from the marine Lower

Limestone Formation (Late Oligocene) of Malta (central Mediterranean). The valves are preserved as moulds, the pallets as stalks without

This is the first fossil record of valves and of the 1770. The hard of the an awn. pallets genus Kuphus Guettard, part morphology new

is with that of its Recent arenarius The abundant fossil tube material available has species compared congener, Kuphus (Linné, 1758). provided not only a documentation of such features as shell impressions, localised swellings, calcareous encrustations, bifurcation, repair, periodic angulation, internal cameration, anterior and internal closure oftubes, but also clarification of the stages of development of its calcareous siphonal tubes. These unique tube features are described and interpreted in the light of the animal’s ability to deposit and resorb calcium carbonate. As all previous fossil Kuphus records are based solely on tubes, they are essentially indeterminate Teredinidae,

identified level. almost world-wide distribution in and and only Maltese material is positively to species From an tropical temperate

Cainozoic shallow is restricted the brackish and of the Indo-Pacific seas, Kuphus now chiefly to waters mangrove swamps region.

words Late hard Key Bivalvia, Teredinidae, Oligocene, Malta, new species, part morphology

G. Zammit Maempel, MQR, D.Sc. (honoris causa), MD, 53 Main Street, Birkirkara, Malta BKR 05,

Contents quelle interne, di impressioni sul guscio di fossili

adiacenti e di encrostazioni organiche, ma anche Riassunto 155 p. evidenze che chiariscono i varii stadi di sviluppo dei Introduction p. 155 tubi calcarei. Tutte le siphonali precedenti seg- Distribution and stratigraphy p. 156 nalazioni specifiche di Kuphus fossile sono basate 157 Systematic palaeontology p. soltanto sulle — dimensioni e forme dei tubi criteri Palaeoecology p. 167 abbastanza validi. Di si consid- non conseguenza, Palaeogeography p. 168 erano questi come Teredinidae o Kuphus indeter- 168 Acknowledgements p. minabili si e considera positivamente identificabile References p. 169 al livello specifico soltanto il materiale Maltese

basato su valvole, palette e tubi. Da una distrib- Riassunto mondiale nei uzione quasi bassifondi tropicali e

— Dal 'Lower Coralline Limestone' una for- durante il temperate l'epoca terziaria, genere Kuphus mazione del Oligocene Superiore dell'isola centro e presentemente ristretto, quasi unicamente, alle

mediterranea di Malta — si salmastre ai della segnala Kuphus aque e 'Mangrove swamps' melitensis, una nuova specie di mollusco bivalva regione Indo-Pacifica. appartenentealia famiglia dei Teredinidae. Questa

anche la delle valvole delle e prima segnalazione e INTRODUCTION del nello stato fossile. La palette genere Kuphus mor- fologia delle parti duri del mollusco terziario mal- Hitherto, all fossil records of the teredinid bivalve

del tese e qui comparata a quella analogo vivente, Kuphus have been of the calcareous tubes only.

arenarius Linne. L'abbondante materiale di Described in the Kuphus present paper are not only the tubi di melitensis ha non soltanto tubes of fossil but also its valves and Kuphus proveduto, Kuphus , pallets, documentazione delle varie chiusure interne ed found for the first time in the fossil state and in esterne del tubo, dei fori associati colle chiusure association with the calcareous tubes. In Malta,

delle associate alcune di such tubes esterne, gonfiature con penetrate the limestone strata more or 156

less vertical in life but occa- — incrassatus 1873 also as (PI. 1, Fig. 1) may Kuphus Gabb, (p. 246; see

30-40° the vertical. In from the sionally be inclined to places, Gabb, 1881, p. 342, pi. 44, fig. 12a-e), there has been a certain amount of destruction and earthy shales of Guayabin (Santo Domingo), reworking of the fragmented tubes into the horizon- 'Middle Miocene', now thought to be Late tal position during Oligo-Miocene times. Except for Miocene.

few isolated derived tube — a fragments, they are Teredo (Cuphus) primigenia Cossmann, 1921 (p. 13,

encounteredin numbers from the almost always very large or pi. 11, fig. 21) 'Stampien' (Oligocene)

of colonies, forming horizontal beds, 50-150 cm thick, Cerens (France).

referred These units arenarius Linnaeus of herein to as ’Kuphus Beds.' are - Cuphus Cottreau & Collig- limited the Lower Coralline Limestone Forma- non from the of to (1924, pp. 278, 279) Burdigalian

Sousset tion (Chattian, Late Oligocene) and occur through- (French Mediterranean coast). out its whole extent. The associated fauna indicates - Teredo (Kuphus) aff. polythalamia (L.) of Cox (1927,

marine environment. from the Miocene of a pp. 62-64, pi. 8, fig. 1)

The horizon' Pemba These tubes ’Magilus grandis described by Torn- (Zanzibar Protectorate). are quist (1904) and by Cottreau & Collignon (1924) as very similar to those of Kuphus melitensis.

in the - occurring Burdigalian (Miocene) deposits on Kuphus arenarius (Linnaeus) of Douglas (1927)

'lie de Mokamby' (SW Africa) and in Madagascar, from the Miocene (Asmari Limestone) of Iran.

' the so-called Vermetus Beds' in the These similar Maltese Burdigalian specimens too are very to

Asmari Limestone of Iran (Douglas, 1927) and the material.

'Beds of Kuphus’ reported by Cottreau (1912) from — Kuphus ?arenarius ?lankae Deraniyagala, 1969 (pp.

Miocene of the from the the Mediterranean coast of France 39-41) Late Oligocene or Early

Bouche du all Miocene of Sri Lanka. This be misiden- (Sousset area, Rhone), are analogous may a

' the Beds'. Fossil tubes to Maltese Kuphus Kuphus are tification, as the tubes are described as widening known mainly from the Miocene of the Caribbean upwards and as occurring always in pairs.

(Gabb, 1873, 1881; Maury, 1925; Oostingh, 1925; - Kuphus polythalamia (Linne) ofjung (1971, p. 161)

Rutsch, 1942;Jung, 1971), France (Cottreau, 1912), from the tuffaceous facies of the Grand Bay

Zanzibar (Cox, 1927), Iran (Douglas, 1927; Cox, Formation (early Middle Miocene) of Curasao

and the Oli- 1927), Sri Lanka (Deraniyagala, 1969) (West Indies). The specimens I examined at the

beds of Malta Naturhistorisches smaller gocene (Spratt, 1852; Jones, 1866; Museum, Basel, are

Zam- much delicate than K. Adams, 1870; Oppenheim, 1916; Felix, 1973; and more melitensis. mit Maempel, 1977, 1979, 1982; Bosence et al.,

1980; Geys & Beeusaert, 1982; Pedley et al, 1990). DISTRIBUTION AND STRATIGRAPHY Sufficient material has been recovered to allow the clarification of the of of the There is work the stages development no previous detailed on Maltese calcareous tubes in fossil and to Cainozoic siphonal Kuphus Kuphus and the first publication to illus- demonstrate its various internal and external fea- these tubes and the in which trate strata they occur

valves, internaland anterior is Zammit In the tures, including pallets, by Maempel (1977, p. 22, pi. 15).

shell and Maltese the closures, pittings, swellings, impressions Islands, Kuphus tubes are restricted to overgrowth ofencrusting organisms. In Teredinidae Attard Member of the Lower Coralline Limestone the classification of the is based the which is genera on mor- Formation (Fig. 2), now considered to be of

of the soft and phology parts pallets (Turner, 1969, Late Oligocene (Chattian) (Giannelli & Salvatorini,

The is pp. N722, 723). genus Kuphus recognised by 1972; Felix, 1973) rather than Eocene (Forbes in the characteristic bifurcated, end of or Miocene long, siphonal Spratt, 1854) Early age (Spratt, 1852, Teredinidae obli- the tube. In addition, other are 1854). Local stratigraphy is summarised in Fig. 2 wood but is known burrow gate borers, Kuphus to and localities yielding the fossil tubes are shown in

in only muddy or sandy substrates. Fig. 1.

identification in teredinids is based on Specific In Malta, Kuphus-bearing beds are best seen at

of and valves. As all fossil joint features tubes, pallets Wied Incita, Attard (central Malta) and San Pawl

Kuphus 'species' recorded hitherto in the literature tat-Targa (along the Great Fault) where limestone

workers refer the tubes are based solely on the size and shape of tube frag- quarry to as sigarri (cigars), ments, their validity is doubted and only Maltese terha (sash, band) or sallur (eels), depending on size material is identified positively to species level. and shape ofthe tubes (Zammit Maempel, 1989, pi.

Recorded include: Other have been recorded in the Mar- species 2). exposures 157

Beds in thickness at different Kuphus vary

localities in the Maltese Islands, but at Wied Incita

below surface (Fig. 3), where the bed is about 23 m

level, it is 90-150 cm thick. It is traceable over the

for distance of faces of the adjoining quarries a

several hundred metres as a soft, rubbly, rust-

stained horizontal band wedged between white

hardness. In limestone of much greater places, the

bed is faulted several metres.

SYSTEMATIC PALAEONTOLOGY

The special morphological terminology used below

is based on Turner (1966, pp. 6, 7, figs 2, 6, 7a, b)

To indicate and Turner (1969, p. N740, fig. E211).

the repositories of specimens studied the following

abbreviations used: Outline of the Maltese Islands recorded are Fig. 1. map showing

- San Pawl tat- Kuphus sites, 1 - Wied Incita, Attard; 2 MCZH — Museum of Comparative Zoology,

- Targa; 3 - Maqluba, Qrendi; 4 - Marsaxlokk; 5 Migra Harvard Cambridge, Massachusetts. - - Ta’ University, lima, Fawwara; 6 - Hal Far; 7 Has Saptan; 8

— - NHML Museum London Kandja Water Galleries; 9 L-Ilsna, Maghlaq (in Natural History (for-

alluvial Pleistocene downthrown deposit overlying merly British Museum [Natural History]), London.

Upper Coralline Limestone Formation); 10 - Ta’ Qali — NHMM National Museum of Natural History,

Pumping Station; 11 - Mosta; 12 - Hondoq ir-Rum- Geology and Palaeontology Section, Malta.

- Cenc. mien; 13 - Mgarr-x-Xini; 14 Ta’ RGM — National Museum of Natural History,

Palaeontology Department, Cainozoic Mollusca

Miner- (formerly Rijksmuseum van Geologie en the sides of the saxlokk area (Jones, 1866, p. 152), on alogie), Leiden (The Netherlands). Maqluba subsidence structure, limits of Qrendi SMF — Naturhistorisches Museum, and Senckenberg (Adams, 1870, p. 270, footnote 19) Migra lima, Frankfurt-am-Main. of Fawwara 'A SE Dingli, limits (Felix, 1973, p. 19).

UZM — author's collection below the (author's original regis- horizon rich in tubes, lying 24-27 m base tration numbers are retained) at the University Zoo- encountered of the Globigerina Fimestone', was by logical Museum, Biology Department, Malta. when T.O. Morris (pers. comm., 1973) striking a

ZM — author's collection. Malta. private shaft at Ta' Qali Pumping Station, central

from the Wied Its location is only two kilometres Family Teredinidae Rafinesque, 1815 and the Incita quarries probably represents exten- Subfamily Kuphinae Tryon, 1862 sion of the In the Kuphus Beds in that area. addition, Genus Kuphus Guettard, 1770 author has encountered isolated specimens in Cai-

— arenaria subse- nozoic rocks at Mosta, Hal Far, Has-Saptan, Ta' Type species Serpula Linne, 1758, by

of Linne Kandja water galleries and (reworked) in the Qua- quent designation Gray (1847, p. 188).

Solen alluvial In the based this name on arenarius of ternary deposit at 1-Ilsna, Maghlaq. (1758, p. 787) described nearby island of Gozo, the author could not locate Rumphius (1705, p. 124, pi. 41, fig. d, e)

the ix-Xini where Adams from Amboina, Moluccas occurring in any tubes in Mgarr gorge (Indonesia),

footnote encountered them 'in mud with (1870, p. 271, 19) gravel.

— animal with large numbers', and only isolated specimens were Diagnosis Non-wood-boring strong muscular collar it behind the shell found at Ta' Cenc and at Hondoq ir-Rummien. encircling just

All the above-mentioned sites have been visited valves; intestine passes through the large ventricular

of the the the Wied bulb heart; caecum by the author but only exposures at elongated lacking; very

reduced adductor Incita quarries have been studied in detail. The posterior muscle; long separate

and covered receding working face of this quarry complex was siphons solid, non-segmented pallets by

off repeatedly investigated during the period 1964-79. periostracum which characteristically peels post

from this site that all the described A calcite tube It was specimens mortem. stout subcylindrical heavy and in this collected. bifurcated encloses the animal figured paper were posteriorly, elongate 158

Lithostratigraphy Chronostratigraphy

Stage

Thickness Formations Members Series Giannelli & Felix, Mazzei, in Pedley, m Salvatorini, 1973 1986 1989 1972

- Gebel Mbark - - EM

Tal-Pitkal - - - T Upper Coralline 0-165 Mioc. Limestone

Mtarfa - - - T

Ghajn Melel - - - T

Greensand - 0-15 Mioc. T - - T

Upper T T T T Clays and

Marls Middle 70 Mioc. S S S S

Fm.

Lower L L L L

Upper 22-70 EL L A/B B Division

Phosphorite 0.1- B B A B Level 2 0.5

Globigerina Middle 10 217? Mioc. B A A B Limestone Division

Phosphorite 0.2- A A A A Level 1 0.6

Lower ? A A A A Division

Il-Mara

Xlendi Lower Coralline >140 Olig. C C Limestone Attard

Maghlaq

2. The Cainozoic of the Maltese Islands. Formations after members after A - Fig. exposed sequence Murray (1890), Pedley (1989),

- - - Aquitanian; B - Burdigalian; Ch Chattian; EL - Early Langhian; L - Langhian; S Serravallian; T - Tortonian; EM Early

Messinian. 159

Schematic of Coralline Limestone Formation the Wied Incita Fig. 3. lithostratigraphy and faunal content the Lower as exposed at

quarry (Attard, Malta) (not to scale).

and attached it the muscles of the — is to solely by Nomenclature Kuphus was first figured by Rumphius

and of the who illustrated pallets siphons. (1705, p. 124, pi. 41, figs d, e), a

— of from Habitat Recent Kuphus live in mangrove areas shelly tube Amboina, Moluccas (Indonesia) the Indo-Pacific where in burrows in and called it Solen Seba they occur arenarius or 'sandpyp'. (1758, mud sand with known the animal and or gravel. They are not to p. 182, pi. 94) figured Kuphus called

it bore into wood. They feed by extending long sepa- 'Tuyaux intestines'. Linne (1758, p. 787, para-

above the bottom and filter from — rate siphons feed graph 699) perhaps considering Kuphus to be a

the Indo-Pacific — referred the animal water (Turner, 1966, 1969). The serpulid to as Serpula arenaria,

of Maltese Cainozoic and other which he affinity Kuphus some a name subsequently (1767, p. 1266)

fossils In local Oligo-Miocene was first described by changed to Serpula polythalamia. 1770, Guettard

Zammit Maempel (1979), but if the 'canaliti introduced the genus Kuphus for the worm-like mol- geniculati vel plus minus reflexi' referred to in lusc Solen arenarius of Rumphius and Kuphus poly-

thalamia became the entrenched for the Bettz's unpublished manuscript (1794, p. 19) are name living

then such noted this ICZN the Kuphus tubes, affinity was by species. However, according to rules,

is valid member of the Order of St. John of Jerusalem 1758 name despite Linne's own correction

in 1767. the animal almost two centuries earlier (Zammit Maempel, Hence, should be referred to as

1982). Kuphus arenarius {Linne, 1758). 160

melitensis tube is less Kuphus sp. nov. acute, thicker, more prominently and

PI. valves 1, PI. 2, Figs 1-10, 12, PI. 3, Figs 5-15; more regularly annulated. The are much

text-figs 4, 5 smaller, more quadrangular, much thicker and with

more and distributed or 1655 of Malta 90. prominent evenly ridges petrified serpents — Worm, p. — ribs that in bunches. The ? 1794 canaliti vel minus reflexi 19. occasionally occur geniculati plus Bettz, p. pallets

? 1805 tubes — de 138. much with stalk sea Boisgelin, p. are more delicate, straighter, flatter

1852 tubes — cylindrical Spratt, 8. and without and with p. preserved an awn, a proximal 1866 of Teredo 153. large species —Jones, pp. 152, cylindrical rod (handle) proportionately larger than 1870 of Teredo — 266 large species Adams, pp. [footnote 19], that of K. arenarius, one third to one fourth 271. forming

of 1916 107. the Magilus grandis Oppenheim, p. length pallet.

with walls and — The tubes 1973 vertical borings showing rings pus- Description gently taper posteriorly

tules on their side — Felix, 19. and p. (upwards), are thick walled thinly laminated (PI.

1977 — Zammit 15. Kuphus Maempel, p. 22, pi. 1, Fig. 3), calcitic, often tortuous, slightly shiny, 1979 — Zammit Kuphus, Kuphus n. sp. Maempel, pp. 4, 7, pi. greyish-green, smooth surfaced with external annu- 2, figs 4, 10. lations 1980 cf. — Bosence with a dull Kuphus polythalamia et al., p. 36. (growth rings) (PI. 1, Fig. 5), — 1982 & 9. smooth internal Kuphus Geys Beeusaert, pp. 44, 45, fig. whitish-yellow, surface. Structures

1989 sigarri, swaba, terha, dudu — Zammit Mae- Kuphus, on and within the tube record information on its 10. mpel, pp. 4, 7, 11, pi. 2, figs 4, environment and past history. Adjoining shells and

1990 — 5a. Kuphus Pedley et al., p. 15, fig. their pebbles leave imprint on the tube's exterior

also (PI. 1, Figs 2, 4; see Savazzi, 1983, p. 297, fig.

— is tube with disarticu- Type Holotype a fragment 1 le). The tube comprises six principal regions (Fig.

lated valves moulds at the end preserved as open (PI. 4):

3, Fig. 12); University Zoological Museum, Biology - the anterior (lower) end, usually open, but Department, Malta, registration number UZM/K. occasionally with hemispherical closure (PI. 1, Fig. 206 (leg./coll. G. Zammit Maempel). 7); Paratypes: — the main undivided tube tapering posteriorly — NHML, no. 40006 (leg./coll. G. Zammit Maem- (upwards) (PI. 1, Figs 1, 2, 4, 5, 7; PI. 2, Figs 3, 12; pel ZM/K. 106). PI. 3, There be conical internal Fig. 7). may one — NHMM/Z.M. Collection (leg./coll. G. Zammit closure (PI. 2, Fig. 7), and several external swellings Maempel), Wied Incita (Attard, Malta), L.C.L. of tube wall (PI. 1, Fig. 6; PI. 2, Fig. 7). Externally, quarries: K.ll, K.20, K.27, K.42, K.47, K.52, the main undivided tube commonly bears impres- K.96, K.200, K.202, K.227, K.229-338. sions of formerly adjacent gastropods, bivalves and — RGM (leg./coll. G. Zammit Maempel, Wied pebbles (PI. 1, Figs 2, 4) and organic calcareous Incita (Attard, Malta): RGM 393 030-393 048 (ex encrustations around its lower (anterior) region (PI. K.5, K.69, K.89, K.203, K.306-318, K.321, PI. there 1, Fig. 7; 2, Fig. 12). During growth, was K.325). often resorption of earlier deposits of the tube; — UZM (leg./coll. G. Zammit Maempel), Wied the internally longitudinally divided tube Incita (Attard, Malta): K.l, K. 10, K. 13, K. 14, without evidence (siphonal end) at surface prox- K.23, K.24, K.30, K.39, K.48, K.53, K.60, K.67, imally (PI. 2, Figs 3, 8-10). Separating the main K. 101, K. 109, K.l 12, K.204, K.210, K.213, K.215, undivided tube from the distal divided siphonal end K.221, K.225, K.302, K.319-326, K.328. is a diaphragm with two adjacent, subequal, subcir- - ZM (author's private collection): K.6, K.22, cular/ovoid openings; K.28, K.37, K.51, K.54, K.57, K.72, K.201, K.207, — the but tubes externally divided attached more K.213, K.339-350 and numerous other specimens. distally (PI. 2, Figs 1, 2);

— the region of external bifurcation (PI. 2, Figs 1,

— After Malta. Derivation of name 2). Extruded siphons leave the tube through a

— Stratum typicum Kuphus Beds, Attard Member resorption area;

Coralline — (Lower Limestone Formation), Late Oli- the posterior end of the tube with two dis-

gocene (Chattian). tinctly separate siphonal tubes (PI. 2, Figs 1, 2), their

Locus typicus — Wied Incita (Attard, Malta), grid exterior covered with irregular, protruding lamellae reference 495714. (PI. 2, Figs 1, 3); the exhalent tube is usually smaller

— In to arenarius the than the inhalent both be in Diagnosis comparison Kuphus one; may irregular 161

4. Schematic reconstruction of melitensis with indication oflevels where and valves have been located Fig. Kuphus sp. nov. tube, pellets

in different tubes (not to scale). 162

5. Reconstruction of the valve of melitensis and internal based artificial Fig. right Kuphus sp. nov. (external views) on casts.

in each small direction; the lumen tube may be as as lacks the terminal, flattened, lateral protuberance

1 mm (PI. 2, Fig. 4). (awn) present in other (Recent) Teredinidae(Fig. 6).

— Discussion The following discussion is largely of melitensis much reduced The valves Kuphus are restricted to a comparison of K. melitensis with (Fig. 5), subquadrangular in lateral view and Recent material of K. arenarius from the Indian and strongly convex; the anterior slope tapering to a Pacific Oceans. Comparative hard parts morphol- point ventrally and maximum width lying at mid is outlined in Table with the addi- ogy 1 following than maximum point, being always greater [dorso- tional information: The umbones The ventral] height. are prosogyrate.

external surface has a sculpture of ventral ribs curv-

ing from umbo to ventral point, the ridges are

unequal and occasionally bunched forming distinct

the shell steps on surface. The inner surface bears a

thick arch on its upper edge, which terminates dor-

sally in a lantern-like apophysis whose surface is

abraded in all available The inner specimens. sur-

of the ventral of the ventral face part shell bears a

lies condyle, an inward-projecting boss which on the

median line; the remaining inner shell surface is

muscle be identified. smooth, no scars could Fig. 6. Terminology of right pallet of Kuphus arenarius (Linné,

based on no. 1933.7.14.1. Measurements — Height, diameter of valves in 1758) specimen NHML, This lacks the and medial of pallet gentle ventral curve apposition and diameter of tube at level of lodge- the stalk noted in other Recent specimens. ment of valves, all in millimetres [and in that order]

the values: 17.5 gave following 11.0, 12.5, [UZM/

K.203]; 12.5, 16.0, 21.0 [UZM/K.48]; 13.0, 20.0, Tube — A Recent tortuous K. arenarius tube in the

40.0 [UZM/K.204]; 14.0, 10.0, 40.0 [UZM/K.89]. author's collection at the Zoological Museum (Biol-

of K. melitensis consist of of Pallets (PI. 3, Figs 5-11) a ogy Department, University Malta, UZM/KK.l)

rod and flat- has of The cir- proximal solid cylindrical (handle) a a vertical height [length] 675 mm.

The cumference and external diameterof tened, lamellated, distal portion (stalk). straight its uppermost

times the size of the handle and stalk is about three [posterior] end are 62 and 18 mm, respectively. The 163

Kuphus arenarius Kuphus melitensis

Pallets

Size bulkier more delicate

Composition aragonitic originally aragonitic, now calcitic

*

Maximal length 80 mm 51 mm

Awn present absent.

Handle/pallet-ratio 1:3-4 1:2.5-5

Stalk course lateral and dorsal con- straight vexity

Stalk structure laminated round and ec- laminated round and eccen-

centric solid rod tric solid rod

Stalk cross-section subcircular subtriangular

Maximal diameter of stalk 4.5 mm 2.8 mm

Tube

Shape more acute less acute

Surface rough, lustreless smooth, shiny

Organic encrustations absent present in lowerlower regionregion

Annulations weak prominent

Ridges wide, irregular narrow, more regular

Inside surface smooth, with external smooth ridges sometimes reflec-reflec- ted

Maximal diameter 41 mm 34 mm (49-53 mm if encrusted)

2 4 Wall thickness max. 2 mm (locally 4-7 max. 4 mm (more with mm) encrustations)

External anterior closure hemispherical hemispherical,'drumstick'- like flattened convexity

Valves

Size max. 20 mm max. 14 mm

Thickness max. 1.0 mm max. 2.1 mm

Shape subtriangular subquadrangular

Ribs close together, notpromi- widely spaced, prominent, nent, regular distribution bunched in areas

Denticulations variable not apparent on peels

of arenarius and K. melitensis Data based several Recent Table 1. Comparison ofhard part morphology Kuphus (Linné, 1758) sp. nov. on

specimens from the Solomon Islands, Philippines, Sumatra, Andaman Islands, and Indian Ocean, and on fossil material

ofthe latter collected by the author from the type locality species. 164

its lower called 'two arched laminae'. corresponding measurements at open (1927, p. 3) These are

[anterior] end are 145 mm and 33-41 mm (internal very clearly evident in the Kuphus arenarius tube

The diameter 31-36 thickness of tube wall at no. 1857.11.10.30.12. with mm). NHML, Possibly, age,

its lower end is 2-5 and is 3.9 40 the animal loses its open mm mm at power of resorbing the thin

above the No of K. melitensis calcareous seal else be in mm edge. specimens or may need of extra

the bifurcated end and the closed lower end preserving posterior (upper) protection provided by or,

closed anterior end been recovered (lower) have perhaps, there is no need for further burrowing into

from the Kuphus Beds at Wied Incita. the sediment.

In Kuphus arenarius, the closed anterior end of the In addition to the 'external' (anterior-end) closure

tube is associated with invariably an adjoining 1-2 just mentioned, Recent Kuphus sometimes develops

cm wide of scattered sieve-like series of somewhat ring irregularly per- a similar, but internal, dia-

forations tube. of the These are analogous to those phragms forming 'camerations'. In view of the

lower of observed on the end [anterior part] mem- infrequent occurrence of these structures, such clo-

the bers of Clavagellacea (genera Clavagella sures probably represent abnormal growth halts, i.e.

1818 and Penicillus of Lamarck, Bruguiere, 1789) and, periods great stress or times of severe hostile

enable the lower end of the environmental conditions presumably, sealed-up either above or below the

animal to keep in direct contact with its environ- sediment/water interface. The phenomenon of

ment also & In cameration first noticed (see Pojeta Sohl, 1987, pp. 2, 4). a was by Home (1806) on the

Recent tube of K. arenarius wormshell' tube (NHML, no. Sumatran 'giant brought back by

1857.11.30.12) these perforations extend also over Griffiths (1806; see also Queckett, 1860). Junghuhn

the two calcareous sheets formed noted similar convex by secre- (1854, pp. 67-70, figs 1-9) a phenome-

tion from the the tube seal inside two opposite sides of to non a fragment of a Recent tube from Java,

the lower end. When of the tube is ascribed him growth resumed, by to a new species of mollusc, called

the perforations are turned into pits which may 'Kurang surumbang'. In spite of his enigmatic

remain evident markedly (specimen NHML, no. description of the tube being 'habitus ceratatoides

In Recent tube sed the 66.5.3.1). some specimens, e.g. a sipho nullius', tube is still considered to be

from Solomons and Ngela Islands, (UZM/KK. 1) that of Kuphus arenarius, for his 'sipho nullius' proba- one from in the Wilkinson Collection Java (NHML, bly refers to the specimen available rather than to

such scattered other the no. 1968.396/2), pits are over species.

besides areas to the site of external A series of regions adjacent posteriorly (upwardly) receding cam- closure or 2, Clustered in (past present) (PI. Fig. 12). erations was observed two comparable fossil

detected on the calcareous encrustation envel- tubes pits (NHML, no. L.25131, post-Pliocene, Gulf of

the closed lower end of K. and oping [anterior] a Suez, NHML, no. L.70352, Pleistocene, Tan- melitensis tube (UZM/K.303) seem to be features of zania) in which cameration is associated with the of the sponge-like surface organic overgrowth localised external tube thickening, massive invasion and related the tube surface. not to underlying by encrusting organisms and with pronounced

annular constrictions in between. X-radiography of

Tube closure — Authors such Griffiths as (1806), Kuphus tube NHML, no. 25132 has shown the swell-

Gray (1858), Douglas (1927), Cox (1927) and Tav- ings to coincide always with the chambers and the ani the end of (1948) referred to lower (anterior) constrictions with the diaphragms (PI. 2, Figs 13,

Kuphus tubes as being 'closed'. Investigations have 14). It is possible that the above-mentioned revealed, however, that Recent Kuphus tubes are encrustations could have been the stimulus inducing sealed actually open, becoming up only peri- the animal's upward withdrawal and the conse-

take when odically. The first closure seems to place quent formation of the upward-progressing dia- the tube is about 50 mm long (observed range: phragms. Another possible stimulus could have

30-63 mm). All adult K. arenarius tubes examined been the choking effect produced by the sudden and showed evidence of had 5-6 excessive of having irregularly deposition sediment on the sea floor. To spaced closures which probably represent periodical relieve itself of this life-threathening situation, the

halts' of 'growth or periods rest with added protec- animal probably extended its already unduly tion to the animal. adults Possibly, only reaching lengthened (progressively tapering) tortuous sipho- maturity and maximum development have their nal tubes, rendering them still less functional but lower end sealed permanently by what Douglas more susceptible to breakage. It is probable that 165

ani- than Protruding detachment of these fragile parts stimulated the higher diaphragm (UZM/K. 213).

tubes from its surface are mal to rebuild wider siphonal and possibly perpendicularly numerous,

conical tubercles with in the false scattered a also to move upwards tube, building a irregularly height 2.5 and maximum base diameter of bottom (diaphragm) beneath it. This might also of about mm a resemble 1.8 mm PI. Fig. 11). They explain the abundance of narrow-bored siphonal (PI. 1, Fig. 7; 2, central internal in stalactitic 'thorns' and like them exhibit a tube fragments (less than 1 mm diameter) broken of the Attard or when or the uppermost section ['C' in Fig. 3] opening (sometimes two three) abraded. The tube wall surface is neither Kuphus Beds (PI. 1, Fig. 1). underlying

nor in affected the encrusting Somewhatanalogous, but more pronounced, reg- pitted any way by

that their ular and uniform, encircling swellings and constric- growth (PI. 2, Fig. 11), so openings cannot

be the observed in tions of the tube occurred in two Maltese Cainozoic related to numerous tiny borings

Massive of the the hard infillof the fragmented cylinders (PI. 3, Fig. specimens (PI. 1, Fig. 6). infilling The encrustation resembles that by calcareous tubes prevented X-ray investigation. The swellings 12).

but its is still unknown. Cal- showed evidence of the usual encrustations (? cal- algae, exact nature

of the conical with lower ends. careous filaments some careous algae) associated [anterior] connecting

bear structures 2, overgrowth to be of Some other Maltese tubes very pronounced, (PI. Fig. 11) suggest

thickness organisms with localised, irregular thickenings (maximum organic origin, possibly encrusting encrustation shell that has since Such 10 mm) due to layers of the organic (PI. aragonitic disappeared. Recent in Red Sea encrustations have been recorded on 2, Fig. 12), noticed also specimens not

A tubes. (NHML, no. 25131; PI. 2, Fig. 13). completely Kuphus

of tube closure Calcium carbonate and — different (conical) type internal resorption deposition

which has been recorded before in Recent or of and never Kuphus possesses great powers resorption

in Mal- The of frac- fossil tubes, was discovered two fragmentary deposition of calcium carbonate. repair

The None of the tube bears tures is (NHML, no. 1968.396/2). tese specimens. fragments preserved

is extruded which leave the main tube encrustation but in one the closure associated siphons, any become covered with with localised of the wall a area, a slight swelling (PI. 2, Fig. through resorption

of this tube shows that the lustreless calcite (PI. 2, Fig. 1). Resorption 7). A longitudinal section rough, of the closure is attained by the wedge-like over-develop- also takes place at strong angulations

translucent undivided tube or tubes K. arenarius ment of one of the innermost dark, siphonal (e.g.

'Growth halts' with white NHML, nos 1968.392/2 and 5). layers that alternate irregularly opaque

the wall and in growth) and subsequent re-activa- layers to form tube (PI. 2, Figs 7-10) (interruptions also the tube After that the localised tube swelling is the result of the tion are recorded on surface. a

tube does from the additional over-development of the white layers. halt, regeneration not proceed

external edge of the old tube but from its inner Preservation of tubes — As stated, K. melitensis tubes short distance from the surface at a edge ('cone-in- end and with bifurcated upper (posterior) complete This results in cone' type structure). a very slight closed lower end are not known from a (anterior) with the of overhang of the old wall production a Maltese Cainozoic deposits. All tubes are frag- characteristic encircling fissure and a localised con- with in size from 40 to 300 mented, parts ranging striction of the tube surface (PI. 2, Fig. 12). In addi- from mm. vibrations explosives Strong resulting tube tion, the calcareous inner lining deposited on used in quarrying the limestone often dislodge the wall plugs the ring of sieve-like perforations above their hard matrix and increase tubes from very frag- the edge of the tube, turning these into mere pits mentation. Tubes that have not been loosened by arenarius tube (NHML, no. 1857/11.30.12, Kuphus the cannot be extracted from the indu- explosives K. with closed lower end; NHML, no. 66.3.5.1, rated matrix of the lowermost division ['A' in Fig. 3] of arenarius tube with lower As a result the open end). of the Beds 3, Fig. 7). Kuphus (PI. outwards seal animal's renewed activity, the convex

— lower end is shattered Calcareous encrustations of tube A thick cal- closing the forcibly (probably

surface with the assistance of of the careous encrustation, having a sponge-like partial resorption seal)

localised and the thin eggshell-like material texture and forming uneven swellings (PI. fragmented very

all lower from the shattered seal is at times found adhering to 2, Fig. 11), envelops regions (anterior

outside of the tube wall at this level arenarius parts) of K. melitensis tubes (e.g. UZM/K.302and K. the (K.

tube since does it extend to a level NHML, no. 1910.4.8.1). Also, penetra- 206). Only very rarely 166

tion of the seal is the tube shows a more the anterior of Recent valves eccentric, slope some Kuphus vary

less marked such in or angulation just beyond 'growth shape according to theirposition on the valves. In

with halt' external closure (PI. 2, Fig. 11). the dorsal region they are fine triangular projections

arising from the external border of the ribs, directed Abnormality — A partitioned siphonal tube with in the middle of the outwards; region valve, they are patent canals (K.72) exhibits sudden dilatation of nodular and swell on either side of the rib, whilst in their tube walls with formation of an internal sac- the ventral region they are flattened, round-edged, culation - first diameter 10 8 on one (maximum x blade-like structures arising from the inner side of mm) and then immediately on the other (maximum the rib and directed inwards towards the gape. They diameter 12 x 10 mm). This resulted in partial probably serve as graters, beaters and scrapers/ detachmentof the two siphonal tubes with evidence It scoopers of mud, respectively. was not possible to at surface. The abnormality is thought to reflect such the rubber of identify any structures on peels underlying pathology of the animal's siphons (? the Maltese Kuphus valves. localised and would have growth) presumably pre- Preservation of valves — melitensis Kuphus valves are vented the animal from extending or withdrawing all preserved as moulds, indicating that the its siphons. compo- sition was probably aragonitic. Cavities left by the

Valves — Previous authors valves {e.g. Gray, 1858; aragonitic provide a weak area in the other-

commented the absence of wise Caiman, 1926) have on solidly infilled tubes which tend to break at this

shell valves from Recent tubes. No evidence level to the internal/external Kuphus expose moulds of the

of such found within the 405 in structures was mm valves. Most valves are apposition (UZM/K.48,

tube closed lower end with no. located either long (NHML, 89, 200-204), high up in the tube

when this time almost 1857.11.30.12) was opened at the blocking its internal diameter (PI. 3, Figs

attached As the lower down (see note to above-mentioned tube). 13-15) or occupying halfits width (PL 3,

calcium possibility of the phenomenon of carbonate Fig. 12). The position of the fossil valves undoubt-

in adult tubes with closed lower the resorption Kuphus edly depends on position and level of the ani-

end was not then Caiman well the tube considered, (1926) sug- mal, as as on the inclination of at the

that gested the Kuphus animal represented a meta- time of the animal's death. Post-mortem shrinkage

adult form of another teredinid of the animal morphosed species, may also be responsible for the high

such mannii which in of the as Dicyathifer (Wright, 1866), position some of fossil valves inside the tube

life bore into wood with the but which In the early valves, (PL 3, Figs 13-15). life, however, valves are

on adult form, shed these and close the lower attaining structures normally to (anterior) end, and were its substrate from changed wood to mud or sand found to be still occupying this position in specimen

with Not all with closed lower gravel. Kuphus (ante- UZM/K.204, with closed anterior (lower) end.

end lose their rior) valves, e.g. NHML, no. The eight Recent Kuphus arenarius animals (from

Another 70 1857.11.30.12. mm long K. arenarius Solomon Islands) examined by the author have

with closed lower end in the (anterior) valves that are much than those of of the larger any Museum Natuurhistorisch Rotterdam (Kastan- Kuphus melitensis (ZM/K.34, ZM/K.48, ZM/K. 106

both valves jesingel Collection), however, yielded (= NHML, no. 40006), ZM/K.200, ZM/K.201,

and when pallets opened by Arie Janssen, the cura- ZM/K.202, ZM/K.203, ZM/K.206 (holotype = of Cainozoic Mollusca tor at the Natural History UZM/K.206), and ZM/K.213).

Museum Leiden (pers. comm., 1971). It is possible

that 1857.11.30.12 and the — The of NHML, no. specimen Pallets pallets Kuphus arenarius are internal

referred to Caiman contained old animal and by an structures attached to the rest of the animal by the

that the Rotterdam sleeve specimen was merely a young muscles and to the tube by their retractor animal with closure a temporary of its lower (ante- muscles. Presumably, when danger threatens, first

end the of of its the and rior) enjoying advantages one tem- siphons subsequendy the pallets, are with-

halts'. It is ascertain the porary 'growth not possible to drawn. The distal ends of two awns are placed in whether of the Cainozoic with closed with each any Kuphus contact other forming a protective can- lower end to tubes did lose their valves, but valves the animal level the opy over at a just beneath in the fossil tube with evidence of were present only diaphragm. The lateral convexity of the Recent a closed lowerend and whose inside could be stalk exam- Kuphus helps facilitate such an apposition and ined Denticulations (UZM/K.204). on the ribs of accommodate the animal in protecting itself against 167

laminations external threats. Each pallet of K. arenarius consists by the accretion of (thick at centre,

rod off around eccentric of a short, wide, proximal cylindrical (handle) tapering laterally) an (medial) encircles and extension solid core. A calcareous laminated a narrower cylindrical (stalk) sepa- deposit

fossil UZM/K.126 rated from each other by an obliquely placed encir- a pallet (PI. 3, Fig. 9) obliquely,

hood-like of cling fibrous band. At the distal end of the stalk is a forming a projecting structure on one

that its sides at the of stalk/handle. It to triangular flattened development (awn) gener- junction seems

end of be the situated and ally encloses at its distal (base triangle) one or analogous to similarly similarly

The stalk fibrous band in more cavities (cups, pockets) (Fig. 6). angulated, tough, present Kuphus

also into the awn without also arenarius no. 1933.7.14.1; see passes discontinuity (see pallets (NHML,

and A number of outwards Moll, 1952, p. 81), gently curving ventrally Fig. 6). grooves radiating

and from the base of the handle medially along its course (NHML, no. upwards convex

The distal better attachment for the sleeve muscles of 1857.11.30.12; PI. 3, Fig. 1). cavity or provided the K. melitensis In Recent better within the is divided pallet. Kuphus, cup awn thereby completely attachment of the sleeve muscles is procured by a (NHML, no. 1933.7.14.1) or partially (NHML, no. and dorsal and more handle by one or more anteriorly- 1857.11.30.12) into a larger a smaller rugged

of the stalk within the directed of the of the handle ventral pocket. The course projections edge (Fig.

It also that brackish be marked was observed water Kuphus awn can usually followed as a pro- 6).

inner wall arenarius tend to have much thicker and larger pal- tuberance running along its flat (medial)

PI. often lets than specimens from marine environments and (NHML, no. 57.11.30.12; 3, Fig. 1), termi- melitensis differ mark- knob its that the of marine K. nating as a distinct at distal edge (NHML, pallets from the but The lateral wall of the is in size and homologous no. 1933.14.7.IV). awn edly shape

inflated than the medial bulkier structures possessed by its modern congener shorter and more one, giv-

from brackish waters. the In some (PI. 3, Figs 1,5-11; text-fig. 6) ing rise to 'cup' (PI. 3, Fig. 1; text-fig. 6).

the lateral no. 1933.7.14.1), specimens (NHML, — Preservation of pallets Pallets are invariably pre- wall bears median cleft indentation at its distal a or with served as calcite stalks handle; most rest on the this is margin. In some specimens, accompanied by lithified infill of the unpartitioned tube (PI. 3, Fig. somewhat lateral within the cup, sug- developments in of found to 7); one case, a set pallets was adhering of the two tube-like structures gestive developed by tube the inside of the uppermost region of an empty

some of the Teredo Linne, 1758. The species genus cemented in minute of position by sparry crystals absence of from all fossil stalks recovered an awn of calcite; other sets jam the siphonal canals tube fossil the nar- and the presence of pallets jamming fragments UZM/K.213 (PI. 3, Fig. 5), UZM/K.215 row-bore siphonal canals of UZM/K.213, 215, 219 (PI. 3, Fig. 11), UZM/K.219 and UZM/K.221 (PI. that and 221 PI. 3, 5, 6, suggest pallets (see Figs 11) whilst 3, Fig. 6), a few others were found free-lying. of K. melitensisnever had an awn. They could conse- The handles of pallets UZM/K.213and 221 project have had different function to quently quite a per- the anteriorly about 1 cm beyond diaphragm into absence of form, an also the opinion supported by the held in empty unpartitioned tube, being position the characteristic lateral and dorso-ventral curva- by minute crystals ofcalcite. Some free-lying pallets in of tures noticed Recent pallet stalks. Sectioning (UZM/K.214, Kuphus Bed, Middle Division = 'B' in the stalk of Recent K. arenarius pallet a (NHML, no. calcite Fig. 3) have their handle encrusted with crys- and of the awn of 1933.7.14.IV) radiography pallet free of such tals and their stalk any encrustation,

another such no. 1933.7.14.1) specimen (NHML, that thus indicating they too once occupied a posi- have revealed that the stalk increases in diameter by tion in the siphonal canals similar to that of UZM/ the addition of successive laminations along its K.213 and 221, but became liberated from their external side (PI. 3, Fig. 2), and thus indicates that tube. tissue the pallets are covered, during life, by compa-

rable with mantle tissue, capable of carbonate PALAEOECOLOGY whilst does the addition of excretion, the awn so by for author successive laminations along its distal border (PI. 3, Three entire Kuphus animals collected the

fracture of fossil stalk from Island Fig. 3). An accidental a pallet by J. Pepys-Cockerell Ngela (British

Solomon Islands where the animal is (UZM/K. 126; PI. 3, Fig. 10) revealed its structure Protectorate),

and called found at the to be identical to that of the Recent Kuphus (NHML, eaten 'lombio', were

both formed entrance to a small brackish water stream no. 1933.7.14.IV; PI. 3, Fig. 2), being running 168

The animals col- situ while after death through mangrove swamps. were Kuphus was living or shortly or

lected at approximately 2000 metres from Biota on after the tops had been broken off and reworked

the northern the Boli in the into the sediment. In entrance to passage overlying the last case, the

of islands. in mud broken could have been Ngela group They were living tops exposed on the sea

containing gravel and small pebbles (1 cm diameter) floor for a considerable period before being buried.

in of a water 30-60 cm with the of their In none of the Maltese Beds wood depth tip Kuphus was any

shell and their soft tissue about 5 wood-like siphons projecting or material found associated with Kuphus

cm above the sediment surface. Although the ani- tubes. Unlike other Teredinidae, Recent and fossil

mal's body is of a gun-metal colour, the projecting Kuphus is not known to (have) bore(d) in wood. The

fossils siphons are bright orange-brown, much resembling associated with Maltese Cainozoic Kuphus

the bivalve roots of decayed trees and therefore well camou- (echinoid Echinocyamus, Chlamys, gastropod

flaged (J. Pepys-Cockerell, pers. comm. 12.3.1973). Cypraea and scaphopod Dentalium at Wied Incita,

and A 370 mm long unbroken Kuphus tube from Java Echinocyamus and a coral reef at San Pawl tat-

has attached to indicative of (NHML, no. 1968.386/1) epibionts Targa) are a shallow, warm, fully its upper [posterior] and middle third. These marineenvironment. Associated fossils of other Cai- include attachment of the nozoic normal post-mortem bryozoan occurrences also suggest a salinity

and Membranipora sp. non-porous barnacles that marine environment (Cottreau, 1912; Cottreau & need fully marine conditions for survival. This Collignon, 1924; Rutsch, 1942;Jung, 1971). shows that Kuphus tubes (even of considerable

PALAEOGEOGRAPHY length) may become partly exhumed or completely dislodged intact from their substrate. Fragmented Geographical and ecological conditions for the tubes be buried 'a few feet for may down', a note by genus Kuphus have altered considerably since early Bertram B. Osmaston of the Imperial Forest Ser- and middle Cainozoic times. From an almost world- vice, the collector and donor of the large end of wide distribution in tropical and subtropical shallow arenarius tube Kuphus NHML, no. 1910.8.4.1, seas the is restricted (Fig. 7), genus now chiefly to records that it few feet below the was 'dug up a the brackish waters and of the mangrove swamps surface, on the beach in estuarian mud, at Hope Indo-Pacific region (Zammit Maempel, 1979, fig. Town, Post Blair, Andaman Is., Indian Islands'. Tavani stressed effect 7). (1948, p. 9) the climatic Old K. 'man- records quote arenarius inhabiting changes had in shifting the distribution of Kuphus grove swamps' (von Martens, 1883, p. 218; south of the equatorial line. Recent Kuphus is most Oppenheim, 1916, p. 108, footnote 14), 'mud with abundantaround the Solomon Islands and the East SE gravel' (Rumphius, 1705, p. 124, Ceram), 'sand Indies, but it does not figure in the faunal list of the with gravel' (Dillwyn, pp. 1087, 1088; forests of the Indo-West-Pacific mangrove swamp Fischer, 1887, pp. 1158, 1159; Oostingh, 1925, p. region compiled by Macnae (1968). Kuphus has dis- 319, Malay Archipelago), 'hard sand' (Sivickis, appeared from the Mediterranean and the Carib- and 1928, pp. 293, 294, pi. 3, fig. 14, Mindaro) bean since the middle Cainozoic. As in those times, 'coral reefs' (Junghuhn, 1854, pp. 64-70, Java, pos- modern still Kuphus inhabits shallow warm sea sibly transported; Hedley, 1895, pp. 464-466; 1898, but it waters, has now moved south of the equatorial As insufficient information p. 96). ecological is given line and has adapted itself, almost exclusively, to in old finds and these records of Kuphus as they brackish conditions and water mangrove swamps. the absence of ignore completely presence or epi-

it is known if these the time bionts, not were living at ACKNOWLEDGEMENTS of exhumation whether dead or they were living on or live tubes. Deep gratitude is expressed to Drs S.R.A. Kelly

invaded also of Epibionts a number fragmented (Cambridge), P.S. Crowther (Bristol), R.P.S. JefF- tubes of K. melitensis in the division of the N. C.P. upper eries, Morris, and Nuttall (all Natural His-

Beds Incita. This R.D. Kuphus at Wied unit represents an tory Museum, London), Turner (Harvard), extensive elongation of an algal-rich calcareous sed- T.J. Palmer (Aberystwyth) and A.W. Janssen iment laid down under normal marine conditions. (Leiden) for help and advice; Drs R.D. Turner

subvertical and buried in The tubes were mostly the (Cambridge, Massachusetts), N. Tebble (then Lon- sediment with their upper parts exposed. It is not don), A. Zilch (Frankfurt) and P. Jung (Basel) for clear whether colonisation took in collections in their the late by epibionts place allowing access to care; 169

Ottobre 1794. Valletta (National Library), Malta, manu-

349, 34 (unpublished). script no. pp.

Boisgelin, L. de, 1805. Ancient and Modern Malta. London

1: lviii 326 14 2: vi + xxxi (Richard Philips), + pp., map, pis;

258 + pp.

Field Guide Bosence, D.W.J., H.M. Pedley & E.P.F. Rose, 1980.

to the Mid-Tertiary carbonate facies ofMalta. London (Pal-

88 aeontological Association), pp.

of of the Caiman, W.T., 1926. Exhibition a specimen giant

Teredo. — Proc. linn. Soc. London, 467: 5.

de Cossmann, M., 1921. Synopsis illustre des mollusques

Pliocene et de l'Oligocene en Aquitaine. — Mem. Soc. geol.

Fr., Paleont., 55: 23-116, pis 11-18. Recent and 7. distribution of (black dots) Fig. Geographical Observation les terrains tertiaires de la Cottreau, J., 1912. sur fossil (open dots) Kuphus. du Cote entre Sousset et l'Anse du Grand Vellat (Bouches

Rhone). — Bull. Soc. geol. Fr., (4)12(5-6): 331-342.

Cottreau, J., & N. Collignon, 1924. Les couches dites a 'Magilus

grandis' de l'ile de Mokamby, province de Majunga - leur P. for two expeditions Pepys-Cockerell setting up leur extension la Cote Ouest a l'extreme Sud de J. age, sur et

locate colonies and collect — 24. (1972-73) to Kuphus speci- Madagascar. Bull. Soc. geol. Fr., (4)24: 278-280, pi.

Dr H.E. Coomans Cox, L.R., 1927. Neogene and Quaternary Mollusca from mens for the author; (Amster- of the Zanzibar Protectorate. In: Report on palaeontology dam), Dr E. Gittenberger (Leiden), A.W. Janssen the collection made Zanzibar Protectorate based mainly on (Leiden), A. Capart (Brussels), R. Kelias (Berlin), by G.M. Stockley A.R.C.S., D.I.C., F.G.S., Government C.E. Palmer Knudsen (Glasgow), J. (Copenhagen), Geologist, 1925-26, Zanzibar. Zanzibar (Government of

Miss Dobson R.M.C. 13-102, 3-19. (London), Eagar (Manches- Zanzibar), pp. pis

1969. The fossil Teredo of Ceylon ter), J.V. Harrison (Oxford), A. Radwanski (War- Deraniyagala, P.E.P., giant Soc. 13 new subspecies. —J. r. Asiat. (Ceylon Branch), n.s., saw), T. Harrison (Kuching, Malaysia), Rev. P.L. 39-41, 1 fig., 1 pi. Seitz (Kontum), Nguyan Van Xuan (NhaTrang) Dillwyn, L.W., 1817. A descriptive catalogue of Recent shells and Lr. for data mate- Truong on museum the Linnean 1-2. London, (Dalat) arrangedaccording to method, pp.

Prof. R.L. L.A. 1-580 581-1092 + 29 rial; J.A. Douglas, Merklin, (1), pp. (2).

Korobkov and G. Tavani for field data from Iran, Douglas, J.A., 1927. Contribution to Persian palaeontology: Kuphus arenarius from the Miocene of Persia. London/ Russia, Cyrenaica and Somalia; Dr L.E. Eames Oxford (Anglo-Iranian Oil Company Ltd.), 5 pp. and Dr P.E.P. Lanka) (Middlesex) Deraniyagala (Sri and Gozo. Felix, R., 1973. Oligo-Miocene stratigraphy of Malta for information about fossil material recovered by 104 8 7 Wageningen (Veeman), pp., figs, pis. Drs L. them from Iraq and Sri Lanka, respectively; Fischer, P., 1880-87. Manuel du conchyliologie et de paleon-

naturelle des for ou histoire mollusques Giannelli and G. Salvatorini (Pisa) reporting on tologie conchyliologique

vivents et fossiles. Suivi d'tin appendice sur les brachiopods the micropalaeontology of the Kuphus Beds sedi- 600 D.P. Oehlert, avec 23 planches contenant figures of par ment, Dr R.J. Davis (London) on the mineralogy desinnees S.P. Woodward. Paris xxv + 1369 par (F. Savy), Maltese tubes and Mr C.P. Palmer Kuphus (London) 1188 24 pp., figs, map, pis. Prof. Dr C.-D. on the associated scaphopods; Gabb, W.N., 1873. On the topography and geology of Santo

the — Am. Soc. 15: 49-259, 2 Reuther (Berlin) for helpful discussions on tec- Domingo. Trans. phil. Phil., n.s.,

tonics affecting Kuphus Beds at Wied Incita, Prof. maps. Gabb, W.N., 1881. Descriptions of Caribbean Miocene fossils. Schembri, Miss M. Dimech and Miss Brown for J.P. — 44, 45. J. Acad. nat. Sci. Phil., (2)8: 337-348, pis

in one of the P. Testrafer- help drawings; Marquis J. Geys.J.F., &J.F. Beeusaert, 1982. Malta en Gozo: Oligoceen en

and — rata Bonici for the sons Ivan, Mioceen een mediterrane Gea, photography my op eilandengroep. 15(2):

their assis- 18 Mark and Joe and daughter Anna for 41-49, figs. foraminiferi Giannelli, L., & G. Salvatorini, 1972. I planctonici tance in the collection of specimens (1964-79). dei sedimenti terziari dell' arcipelago maltese, 1. Bio- Their help is gratefully acknowledged. stratigrafia del 'Globigerina Limestone'. — Atti Soc. Tosc.

Sci. nat., Mem., (A)79: 49-74.

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1895. — Proc. linn. Soc. 6 Hedley, C., Conchological notes. NSW, 449-488, figs, map.

23(1): 91-96. Oostingh, C.H., 1925. Report on a collection of Recent shells

Hedley, C., 1898. Further notes on Australasian shipworms. — from Obi and Salmahara (Moluccas). — Meded. Landb.

Proc. linn. Soc. 91-96. 382 + 1 NSW, 23(1): Hogeschool Wageningen, 29(1): pp.

Observations the shell of the Ober die Home, E., 1806. on sea worm Oppenheim, C.H., 1916. Tertiarbildungen von

Deutsch-Ostafrika. — found off the coast of Sumatra, proving it to belong to a Z. dt. geol. Ges., 68: 103-110.

ofthe of Teredo Controls species ofTeredo; with an account anatomy Pedley, H.M., 1989. onCenozoic carbonate deposition — navalis. Phil. Trans, r. Soc. Lond., 96: 276-292,pi. 12, fig. in the Maltese Islands: review and reinterpretation. — Mem.

4; pi. 13. Soc. geol. It., 38 (1987): 81-94.

Notes from Malta. H.M., D. Bosence, C. Dart S. Pratt Field Jones, T.R., 1866. onsome specimens Appen- Pedley, & [1990]. — dix to: F.W. Hutton. On the physical geology of Malta. Guide to the Cenozoic Platform Carbonates of the Maltese

Geol. 152. Islands. Field 54 22 hand- Mag., 3(22): 151, Trip Guide, pp., pis. (unpublished

Jung, P., 1971. Fossil mollusks from Carriacou, West Indies. — out).

Bulls Am. Paleont., 61(269): 147-262, 2 tabs, 24 pis. Pojeta, J. Jr., & N.F. Sohl, 1987. Ascaulocardium armatum

1854. In: L. Arnold seine new the ultimate Junghuhn, F., (ed.). Java, Gestalt, (Morton 1833), genus (late Cretaceous):

Pflanzendecke und innere Bauart, 3. Leipzig (J.K. Hasskarl), variation on the bivalve paradigm. Mem. Paleont. Soc., 24:

316 58 77 42 pp., figs. pp., figs.

C. 1758. 1. 1860. of the of the Museum of Linne, von, Systema naturae per regna tria, (10th Queckett, J., Catalogue contents

Holmiae iv + 824 the of of 1. Plants and edition). (Laurentii Salvii), pp. Royal College Surgeons England,

invertebrate Linne, C. von, 1767. Systema naturae, 1(2) (12th edition). animals in the dried state. London (Taylor and

Holmiae: 533-1327. Frances).

Account ofthe fauna and flora of 1705. D'Amboinische Rariteitkamer. Amster- Macnae, W., 1968. mangrove Rumphius, G.E.,

and forests in the Indo-West-Pacific In: F.S. dam, 30 + 340 + 43 60 swamps region. pp., pis.

Russell & M. Yonge (eds). Advances in marine Biology, 6. Rutsch, R.F., 1942. Die Mollusken der Springvale Schichten

London/New York (Academic Press): 73-270, 76 figs, 2 tabs. (Obermiocaen) von Trinidad (British West Indies). — Verh.

naturf. Ges. 54: 96-186, 2 3-9. Martens, E. von, 1883. Die Weich- und Schaltiere gemain- Basel, figs, pis

fasslich dargestellt. Leipzig (Fentag)/Prag (Tempsky), iv + Savazzi, E., 1983. Adaptations to tube dwelling in Bivalvia. —

327 205 15: 21 pp., figs. Lethaia, 275-297, figs.

1925. A further contribution to the Seba, A., Maury, G.J., palaeontology 1758. Locuplatissimi rerum naturalium thesauri, 3.

of Trinidad. Bulls Am. Paleont., 96, 18, 9, Amsterdam 212 116 10(42): pi. figs (Jausonio-Waesbergiosa), pp., pis. — 10. Sivickis, P.B., 1928. New Philippine shipworms. Phil. J. Sci.,

1986. The Miocene ofthe Maltese Islands: 3 Mazzei, R., sequence 37(3): 285-298, pis.

biostratigraphic and chronostratigraphic references based on Spratt, T.A.B., 1852. On the Geology of Malta and Gozo. Malta — nannofossils. Atti Soc. ital. Sci. nat., Mem., Governor = Wm. Mail 24 (A)92(1985): (The Reid) Daily Office, pp.,

3 1 tab., 3 165-197, figs, pis. maps.

PLATE 1

melitensis Lower Coralline Limestone Formation Wied Incita Attard Kuphus sp. nov., (Late Oligocene, Chattian), quarry, (central Malta).

1. Detail of Beds; tubes Scale Fig. Kuphus penetrating strata subvertically. bar equals 53 mm (= diameter of white ring on lens-

cover).

Growth Fig. 2. annulations, pittings and deep accidental scars on tube surface (UZM/K.345), x 0.6.

Fig. 3. Siphonal tube fragment (UZM/K.10). Hard (laminated) tube wall enclosing concentric calcite laminations around

siphonal canals, exposed by accidental oblique wear, x 1.5.

4. of bivalve Anadara tube wall x 1. Fig. Deep impression a ribbed, subquadrangular (? genus Gray, 1847) on (UZM/K.65),

Fig. 5. Annular ridges and coloration oftube are limited to outermost thin laminations (UZM/K.305), x 1.

Fig. 6. Annular swellings oftube UZM/K.225 with calcareous encrustations (compare Pl. 2, Figs 7, 13), x 1.

7. Calcareous encrustations round Fig. on closed lower (anterior) end of tube (UZM/K.112), x 1.

Fig. 8. A smooth-surfaced, posteriorly-tapering juvenile Kuphus melitensis tube (UZM/K. 113), with ? evidence ofinternal division

Scale anteriorly. bar equals 1 cm.

9. Detail of 8. Note 2. Fig. Fig. rounded, externally-undivided, closed, posterior end, x PLATE 1 171\1 172

1854. On the of Malta and 2nd 1655. Museum Historia Spratt, T.A.B., Geology Gozo, Worm, O., Wormianum seu rerum rar-

edition. Governor = Wm. 16 iorum Hafniae Danorum in aedibus auctoris (The Reid) Whitefriars, pp., quae servantur.

Batavorum map. Lugduni (Officina Seviriana).

1948. Osservazioni sul Guettard. — Zammit 1977. outline of Maltese Tavani, G., genere Kuphus Maempel, G., An geology.

Atti Soc. Tosc. Sci. 1-10, 1 tab. Malta 44 6 29 nat., Mem., (A)55: (The author), pp., figs, pis.

1904. einer Fauna der Westkiiste Zammit 1979. The Indo-Pacific Tornquist, A., Uber eocene Maempel, G., affinity of some

von Madagascar. — Abh. Senckenb. naturf. Ges., 27(2): Maltese Tertiary fossils. — Centr. Medit. Nat., 1(1): 1-12.

323-327, Zammit G., earliest pi. 46, fig. 11. Maempel, 1982. The 'Treatise' on Maltese

Turner, R.D., 1966. Survey and illustrated catalogue of the fossils. — Melita Historica, 8(2): 133-148. — Teredinidae (Mollusca: Bivalvia). Cambridge (Museum of Zammit Maempel, G., 1989. The folklore of Maltese fossils.

Harvard vii 269 25 in Mediterranean Social Comparative Zoology, University), + pp., Papers Studies, 1: 29 pp., 4 pis.

figs, 64 pis.

Turner, R.D., 1969. Teredinidae. In: R.C. Moore (ed.). Treatise

on Invertebrate Paleontology, Part N, Mollusca, Bivalvia,

6(2): N 722-N 741, figs E196-E213. Boulder and Lawrence

(The Geological Society ofAmerica and University ofKansas Manuscript received 27 February 1993, revised version accepted

Press). 8 October 1993.

PLATE 2

12. melitensis Late Wied Incita for Figs 1-10, Kuphus sp. nov. tubes, Oligocene, (Malta), except Fig. 6, which is from the Kuphus Beds,

Division C, at Attard.

1 - Extrusion of different levels and siphons at subsequent bifurcation oftubes. Thick, lustreless, uneven, calcareous deposit

covers emerging siphons (UZM/K.23), x 1.

2 - Siphonal tubes attached proximally, bifurcating distally (UZM/K.24), x 0.7.

3 - Longitudinal section of tube in region of diaphragm, showing section of main undivided tube, diaphragm and internal

division without evidence at surface (UZM/K.67), x 1.

4 - 86 mm narrow-bored, tortuous tube UZM/K.53 with dull and calcite Detached, long, siphonal irregular laminations, x

0.9.

5 - Transverse section of tube UZM/K. 14 formation of second showing incipient siphonal tube, x 3.5.

6 - tube UZM/K.30 with Cliona 1. Siphonal fragment multiple (? post-mortem) sponge borings, x

7 - section of tube UZM/K.210, with dark Longitudinal unpartitioned fragment crystalline layer in its wall showing an

almost conical internal closure complete and swelling of tube wall (compare Pl. 1, Fig. 6), x 1.5.

8 - Transverse section of tube UZM/K.109. of dark very thick-walled, unpartitioned fragment Regional thickening

laminations is crystalline suggestive of incipient conical internal closure, x 1.5.

9 - Transverse section of siphonal end of tube UZM/K. 1, with thick wall, irregularly alternating light and dark coloured

laminations and canals encircled first and then 1.5. narrow subequal siphonal individually jointly (cable-like), x

10 - Transverse section of siphonal end of tube UZM/K.300, demonstrating wide, subequal, thin-walled siphonal canals.

Narrower siphon about to be extruded, x 2.

12 - Unpartitioned Kuphus tube fragment UZM/K.302, with heavy organic calcareous encrustation and perforated conical

structures times interconnected calcareous filaments, a - at by exposed tube wall unaffected by the encrustation, x 1.3.

11. arenarius Solomon Islands Protectorate. Lowermost of UZM/ Fig. Kuphus (Linné, 1858), Recent, Ngela Island, segment tube

KK. c - site of closure of - of turritellid - 1, x 0.4, sutures marking previous tube; g deep impression a gastropod;o open

lower - - - accidental (anterior) end; p pittings; r growth rings; s deep scars.

14. Post-Pliocene beach South Red Sea. 13 - tube with Figs 13, Kuphus sp. deposit, Gaysum Islands, Gulf of Suez, NHML, no. 25131,

calcareous encrustations Pl. - organic analogous to 2, Fig. 12, x 1.3; 14 radiograph of same tube revealing internal

closures and camerations, x 1.3. PLATE 2 173\2 174

PLATE 3

Figs(Linné,1-4. Kuphus arenarius 1738), Recent, Solomon Islands.

1 - External view ofleft pallet (NHML, no. 57.11.30.12), lacking proximal end (handle). Note dorso-ventral curvature of stalk

and its extension into the into and smaller 2. awn dividing cup a larger (dorsal) a (ventral) pocket, x

2 - Transverse section ofleft pallet stalk NHML, no. 1933.7.14.IV,showing laminations around a medial hard core (compare

structure of K. melitensis pallet, Pl. 3, Fig. 10).

3 - Radiograph ofpallet stalk of NHML, no. 1933.7.14.I,showing growth of awn by deposition ofhorizontal laminations on

distal edge.

4 - Anterior view of valves of in characteristic wide 2.5. NHML, no. 1933.7.14.I, apposition, showing pedal gape, c x

5-15. melitensis Lower Coralline Limestone Attard Late Wied Figs Kuphus sp. nov., Kuphus Beds, Formation, Member, Oligocene,

Incita, central Malta.

5, 6 - Tube fragments UZM/K.213 and UZM/K.221, respectively, with both pallets jamming siphonal canals and with handle

protruding into unpartitioned tube. Note abnormal development ofpallet on left in Fig. 5, x 1.

7 - Crushed tube (UZM/K.212) with both pallets (distal ends slightly chipped) embedded upright in infill of unpartitioned

tube section, x 2.

8 - Straight-shafted pallet with an unduly short handle, recovered from an unpartitioned tube fragment (UZM/K.223),

Division C of Kuphus Beds, Attard, x 2.

9 - Pallet UZM/K. 126 with deformed broken and calcareous its handle/stalk slightly stalk, tip a developmentover junction, x

2.

10 - Detail of end of shown in 9. Note eccentric hard in Recent arenarius upper specimen Fig. (medial) core as Kuphus (compare

Pl. 3, Fig. 2).

11 - of tube with removal of tube wall each Fragment internally partitioned (UZM/K.215) partial to expose a pallet jamming

of the canals. Note handles into undivided 2.2. two siphonal protruding tube, x

12 - tube UZM/K.206 with both valves embedded in matrix 2. Note thickness of valves. Holotype, (moulds) at open end, x

13 - Latex peel from paratype NHML, no. LL.40006 (ex Zammit-Maempel Collection, K. 106) showing commarginal

ornament.

14 - Anterior view of tube UZM/K.48 with internal mould ofboth valves in apposition, x 1.4.

15 - Rubber peel of internal mould of both valves in apposition in tube RGM 393 033 (UZM/K.203) showing abraded

apophysis, x 1.9. PLATE 3 175\3