Zootaxa 3911 (4): 560–570 ISSN 1175-5326 (print edition)

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Copyright © 2015 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.3911.4.5
http://zoobank.org/urn:lsid:zoobank.org:pub:A73BEF82-6D47-40FD-8EF0-206B1AB948F7

New Indo-Pacific species of the genus Teretia Norman, 1888

(Gastropoda: Raphitomidae)

MAURO MORASSI1 & ANTONIO BONFITTO2,3

1
Via dei Musei 17, 25121, Brescia, Italy
2
Department of Biology, Geology and Environmental Sciences (BiGeA), University of Bologna, via Selmi 3, 40126, Bologna, Italy
3
Corresponding author. E-mail: antonio.bonfitto@unibo.it

Abstract

Four new species are assigned to the genus Teretia Norman, 1888 in the family Raphitomidae Bellardi, 1875 and herein
described: Teretia neocaledonica sp. nov., T. sysoevi sp. nov., T. tongaensis sp. nov. from the southeastern Pacific and
Teretia tavianii sp. nov. from the Gulf of Aden. The new species represent the first Indo-Pacific record of a genus previ-
ously known in the recent molluscan fauna by only two species from the Atlantic Ocean-Mediterranean Sea and Southern
Africa. A possible Tethyan origin for the genus Teretia is suggested.

Key words: Systematics, Mollusca, biodiversity

Introduction

The conoidean family Turridae sensu Powell, 1966 is the largest family in Mollusca with about 4,000 named living
species (Tucker, 2004) and 358 recognized genera and subgenera (Bouchet et al., 2011). Despite this large number
of described species, the total number of extant “turrid” species is likely to be much higher with Bouchet et al.
(2008) suggesting that 5,000 turriform gastropods occur in the South Pacific alone, a considerable part of which is
still underexplored. In this paper we describe four new species from the Indo-Pacific that resemble members of
Teretia Norman, 1888, a genus well represented in the Tertiary of Europe but previously known in the Holocene by
only two species from the Atlantic-Mediterranean Sea and Southern Africa. Although the discovery of the new
species suggests a remarkable recent diversification, we were unable to find records of fossils of the genus Teretia
in the Indo-Pacific. Based on these findings and information available from literature, it is hypothesized that
Teretia had a wide west-east distribution thus providing an additional example of the well-documented faunal
exchanges between different sectors of Tethys Realm as defined by Harzhauser et al. (2002). Inclusion of Teretia in
the family Raphitomidae follows the revised classification of the Conoidea proposed by Bouchet et al. (2011) in
which the traditional polyphyletic taxon Turridae is resolved as 13 monophyletic families.

Material and methods

Three species were dredged off New Caledonia (T. neocaledonica sp. nov., and T. sysoevi sp. nov.) and Tonga
Islands (T. tangaensis sp. nov.) by the French Tropical Deep-Sea Benthos Programme, a joint project of the Institut
de Recherche pour le Développement (IRD) and the Muséum National d’Histoire Naturelle, Paris. Teretia tavianii
sp. nov. was dredged off the Gulf of Aden in September 1992. Descriptions and measurements are based on shells
oriented in the traditional way, spire up with the aperture facing the viewer. SEM micrographs were taken using a
Hitachi S-2400. The following abbreviations are used in the text: MZB = Museo di Zoologia dell’Università di
Bologna; MNHN = Muséum National d’Histoire Naturelle, Paris; l = length; b = shell width; a = aperture length; a/
l = ratio of aperture length to total shell length; b/l = ratio of shell width to total length; dd = empty shell(s); stn =

560 Accepted by T. Duda: 9 Dec. 2014; published: 21 Jan. 2015

station. All measurements were made using a Leica Stereomicroscope for incident light and 10X ocular
micrometer.

Taxonomy

Superfamily Conoidea Fleming, 1822

Family Raphitomidae Bellardi, 1875

Genus Teretia Norman, 1888

Type species: Pleurotoma anceps Eichwald, 1830

Diagnosis. Shell up to about 12 mm in length, narrowly fusiform to fusiform, sculpture predominant spiral
consisting of spiral cords of nearly equal strength or with a peripheral cord stronger than other cords. Axial growth
lines forming at most weak plicules on subsutural ramp. Aperture lanceolate, outer lip simple forming a deep to
very deep sinus immediately beneath the suture. Protoconch planktotrophic, narrowly conical with diagonally
cancellate sculpture.
Remarks. The genus Teretia Norman, 1888 was proposed as nomen novum for Teres Bucquoy, Dautzenberg
and Dollfus, 1883 non Boettger, 1878, with Pleurotoma anceps Eichwald, 1830, a European Miocene-Pliocene
species, selected as type-species (see Tucker 2004 for detailed references). The genus Teretia is documented in
Europe at least from the early Miocene. Janssen (1984) reported six species from the Burdigalian of Winterswijk
(Netherlands): Teretia fusianceps Nordsieck, 1972, T. anceps Eichwald, 1830 and four unidentified species. In his
revision on the Pliocene “turrids” from southern Spain, Vera-Peláez (2002) described Teretia pentacarinifera Vera-
Peláez, 2002 and Teretia policarinarum Vera-Peláez, 2002 from Málaga, two species morphologically similar to
Teretia anceps and T. teres (Reeve, 1844). According to Vera-Peláez (2002: 223), the specimen from the Pliocene
of Roero (Italy) figured by Cavallo & Repetto (1992: 134, fig. 356) under the name Stenodrillia sulciensis
(Bellardi, 1877) belongs to Teretia pentacarinifera. The online checklist of Italian Pliocene fossil “turrids” of the
Italian Society of Malacology (SIM) (SIM Editorial Board, 2014) recognizes seven Teretia species, including the
very poorly known T. nana (Hornung, 1920), and lists T. intermedia (Foresti, 1874) as a full species while Brunetti
& Vecchi, 2003 treated this latter taxon as a “form” of T. anceps. The SIM checklist, however, makes no mention of
T. monterosatoi (Cipolla, 1914) considered a valid species by Brunetti & Vecchi (2003). The latter authors stated
that T. fusianceps from the Miocene of Netherlands is very similar to the Miocene-Pliocene T. turritelloides
(Bellardi, 1847) but did not explore this resemblance further. Brunetti & Vecchi (2003: pl. 2 fig. 6) figured a
specimen from Monteveglio (Bologna, Italy) under the name T. turritelloides, but this specimen differs from the
syntype of T. turritelloides photographed by Ferrero Mortara et al. (1981: plate 18, fig. 6) in having more
prominent spiral sculpture and fewer secondary cords, and seems more similar morphologically to the specimen in
the photograph of T. fusianceps from Winterswijk (Gelderland, Netherlands) published by the Natural History
Museum of Rotterdam (NMR) (NMR Editorial Board, 2014). Baluk (2003: 67) considered T. fusianceps a
synonym of T. anceps and figured three specimens under this name, two of which (Figs. 1–2) are comparable to T.
fusianceps while the other has “typical” features of T. anceps. Two additional fossil species described by Seguenza
(1880:258), namely Homotoma cincta and H. multicingula, both originally compared to Teretia anceps, are
currently accepted as members of Teretia (Brunetti & Vecchi, 2003). More recently, Schnetler (2005) described
Teretia guersi from the Gram Formation, Late Miocene of Denmark, which was based on two specimens retaining
only 1½ teleoconch whorls but morphologically distinctive. Barnard (1958) described Acrobela acus on the basis
of examination of a few shells from Southern Africa. Although the species has a typical raphitomine diagonally
cancellate protoconch, it was assigned to genus Acrobela Thiele, 1925, which is currently regarded as a synonym
of Microdrillia Casey, 1903 (see WoRMS Register) in the family Borsoniidae Bellardi, 1875. In his revision of the
borsoniine gastropods from Southern Africa, Kilburn (1986: 635) noted this discrepancy and suggested allocation
of A. acus in Teretia. Althougth Barnard (1958) did not compare the Southern African species with Teretia teres,
the two taxa are actually very similar and examination of the type material of A. acus would be necessary to more
precisely define its morphological features and confirm specific distinction from T. teres. Following current
literature (WoRMS Register), A. acus is here treated as a valid species belonging to the genus Teretia.

NEW INDO-PACIFIC TERETIA SPECIES Zootaxa 3911 (4) © 2015 Magnolia Press · 561
Teretia neocaledonica sp. nov.
Figures 1.A–J

Type material: Holotype (MNHN IM-2000-28374) and 22 paratypes (21 paratypes MNHN IM-2000-28376; one
paratype MZB60095)
Type locality: South of New Caledonia, Bank Eponge (mont 8), 24°55S 168°22E, 502–532 m [N/O “Alis”
SMIB 8 stn DW146-147].
Material examined: South of New Caledonia: Bank Eponge (mont 8), 24°55S 168°22E, 502–532 m [N/O
“Alis” SMIB 8 stn DW146-147] (23 dd, holotype and 22 paratypes); Bank Eponge (mont 8), 24°55.20'S
168°21.73'E, 514–522 m [N/O “Alis” SMIB 8 stn DW146] 27 January 1993 (1 dd). Coral Sea, Bank CAPEL:
24°44.83 S 159°41.00 E, 285 m [N/O “Coriolis” MUSORSTOM 5 stn DW 274], Bouchet, Metivier& Richer coll.
09 October 1986 (1 dd). New Caledonia, Loyalty Ridge: 24°45,55'S 170°07,40' E, 850–855 m [N/O “Alis”
BATHUS 3 stn DW 780], Bouchet, Richer &Warén coll. 24 November 1993 (1 dd); 23°47,50'S 169°48,75'E,
731–751 m [N/O “Alis” BATHUS3 stn DW 793], Bouchet, Richer &Warén coll. 26.11.1993 (1 dd); 23°35,12' S
169°36,73' E, 655 m [N/O “Alis” BATHUS 3 stn DW 800], Bouchet, Richer &Warén coll. 26 November 1993 (1
dd); 23°39,39'S 167°58,94'E,650–730 m [N/O “Alis” BATHUS3 stn DW 809], 27 December 1993 (1 dd).
Description. Shell (Figs. 1.A–D) fusiform (b/l 0.41; a/l 0.33) with high spire, strongly excavated base and
relatively long anterior canal. Teleoconch up to about 4¾ whorls (four in most specimens) with shallowly
impressed, linear suture. Subsutural ramp wide, shallowly concave. Earlier two teleoconch whorls with a
prominent peripheral cord below mid- height of the whorl and two closely spaced weak cords bordering abapical
part of subsutural ramp. On third whorl the adapical cord on subsutural ramp becomes bisected and a second main
cord develops at level of abapical suture (Figs. 1.E–F). Last whorl with a third main cord and additional weak
cords: one on abapical part of subsutural ramp and 1–2 in each interspace between main spiral cords. Last whorl
with 26 spiral cords on base and rostrum. Subsutural ramp with two spiral threads bordering adapical suture.
Teleoconch whorls sculptured by fine axial growth lines forming weak, widely spaced, arcuate plicules on
subsutural ramp, rendering somewhat nodulous the two spiral threads bordering adapical suture (Fig. 1.E). Under
SEM (Figs. 1.E–G), surface is seen to be covered by a sculpture of microgranules. Aperture lanceolate, outer lip
thin. Anal sinus very deep, narrow, asymmetrical, with its deepest point on sutural ramp (Fig. 1.D). Protoconch
(Figs. 1.H–J) tall, narrowly conical of 3½–4 whorls; first whorl tilted with microgranules arranged in spiral rows,
subsequent whorls with diagonally cancellate sculpture terminating a short distance above abapical suture where
fine slightly prosocline riblets occur. Maximum diameter about 0.48 mm. Shell yellowish-white to white with a
yellowish-brown band on periphery of whorls: weak to absent on first whorl, distinct on penultimate and last
whorls, and (in some specimens) also at level of limit base-rostrum. Some specimens with few weak, light brown,
subsutural blotches on last whorl. Protoconch brown. Dimensions: Holotype 5.9 x 2.4 mm, aperture height 2.6 mm.
Remarks. Teretia neocaledonica sp. nov. resembles the Miocene-Pliocene T. turritelloides (Bellardi, 1847) in
shape and disposition of spiral cords but differs from the latter in details of protoconch and teleoconch sculpture.
More in detail, Teretia neocaledonica sp. nov. has slightly fewer protoconch whorls than T. turritelloides (3.5–4
versus 4–4.5) with a different sculpture; in the new species the axial riblets strongly decussate on periphery of each
protoconch whorl, particularly on last whorl, while in T. turritelloides decussation occurs on whorl surface except
for a narrow area near adapical suture (see Brunetti & Vecchi, 2003: plate 3, fig. 3). Furthermore, judging from the
syntype of T. turritelloides from the Pliocene of Colli Astesi, figured by Ferrero Mortara et al. (1981: pl. 18, fig.
6a–6b), the fossil species has a secondary sculpture more prominent than T. neocaledonica. The Italian Pliocene
Teretia elegantissima (Foresti, 1868) differs distinctly from the new species in having a much more prominent
peripheral keel formed by numerous (4–6) closely spaced spiral cords (fig. 3.G).
Etymology. Neocaledonica = alluding to the fact that the type material was dredged off New Caledonia.

Teretia sysoevi sp. nov.

Figures 1.L–T

Type material. Holotype (MNHN IM-2000-28375) and two paratypes (paratype 1 MNHN IM-2000-28377
(coated); paratype 2 MZB60096).

562 · Zootaxa 3911 (4) © 2015 Magnolia Press MORASSI & BONFITTO
FIGURE 1.A–J. Teretia neocaledonica sp. nov. A. Holotype (MNHN IM-2000-28374), New Caledonia, Cruise SMIB 8, stn
DW146-147, 5.9 x 2.4 mm. B–J. Paratype (MZB60095), New Caledonia, Cruise SMIB 8, stn DW146-147. B. Shell, 4.9 x 2.1
mm. C–D. Coated specimen, frontal and lateral view, scale bar 2 mm. E. Penultimate whorl, scale bar 500 µm. F–G.
Microsculpture of penultimate whorl, scale bar 100 µm. H–I. Protoconch, scale bar 500 µm. J. Details of protoconch 1, scale
bar 100 µm. L–T. Teretia sysoevi sp. nov. L–M. Holotype (MNHN IM-2000-28375), South-West Pacific, Cruise
MUSORSTOM 7, stn DW507, 5.8 x 2.3 mm. N. Paratype 1 (MNHN IM-2000-28377), New Caledonia, Cruise BATHUS 3, stn
DW786, 7.2 x 2.8 mm. O–T. Paratype 2 (MZB60096), New Caledonia, Cruise MUSORSTOM 4, stn DC 168. O. Shell, 7.5 x
2.9 mm scale bar 1 mm. P. Penultimate whorl, scale bar 600 µm. Q. Microsculpture of penultimate whorl, scale bar 200 µm. R.
Microsculpture of penultimate whorl, scale bar 80 µm. S–T. Protoconch, scale bar 200 µm.

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 Type locality. Wallis & Futuna Islands, 14°19,6'S 178°06,7'E, 419–425 m [MUSORSTOM 7 stn DW 507], 11
May 1992.
Material examined. Wallis & Futuna Islands, 14°19,6'S 178°06,7'E, 419–425 m [MUSORSTOM 7 stn DW
507], 11 May 1992 (1 dd, holotype); New Caledonia: 18°48,20’S 163°10,80’E, 720 m [MUSORSTOM 4 stn DC
168] 16 September 1985 (1 dd, paratype 1); Loyalty Ridge: 23°54.46 S 169°49.15 E, 699–715 m [N/O “Alis”
BATHUS 3 stn DW 786], Bouchet, Richer & Warén coll. 25 November 1993 (1 dd, paratype 2).
Description. Shell (Figs. 1.L–O) fusiform (b/l 0.39; a/l 0.38) with high spire, strongly excavated base and
relatively short anterior canal. Teleoconch of five whorls (six in paratypes) with shallowly impressed, linear suture.
Subsutural ramp wide, moderately deeply concave. Earlier two teleoconch whorls with a prominent peripheral cord
below mid- height of the whorl and two weak cords bordering abapical part of subsutural ramp (the abapical one
bisected). On third whorl a second main spiral cord develops at level of abapical suture increasing in strength on
subsequent whorls (Fig. 1.P). In one paratype the second main cord develops from first whorl onwards. Last whorl
with a third main spiral cord near abapical suture and one weak cord in the interspace between peripheral and
second main cords. Last whorl with 19 spiral cords on base and rostrum. Subsutural ramp lacking spiral sculpture
(holotype) or with up to four low threads (paratypes) on later two whorls. Teleoconch whorls sculptured by fine
axial growth lines forming weak, irregularly spaced, arcuate plicules on subsutural ramp (Fig. 1.P). Under SEM
(Fig. 1.R), surface is seen to be covered by a sculpture of dense microscopic pustules. Aperture lanceolate, outer lip
thin. Anal sinus very deep, narrow, asymmetrical; its deepest point on sutural ramp. Protoconch (Figs. 1.S–T) of
2½+ whorls; tip missing, remaining whorls with diagonally cancellate sculpture terminating a short distance above
abapical suture where fine slightly prosocline riblets occur. Maximum diameter about 0.48 mm. Colour of
teleoconch whitish to pale buff, protoconch light brown. Dimensions: Holotype 5.8 x 2.3 mm, aperture height 2.2
mm. Largest Paratype (MZB): 7.5 x 2.9 mm, aperture height 3.1 mm.
Remarks. Teretia sysoevi sp. nov. is morphologically similar to Teretia neocaledonica sp. nov. from which it
differs in numerous details of shell morphology. Teretia sysoevi sp. nov. has more numerous teleoconch whorls
(5–6 versus 4–4¾), a more deeply concave subsutural ramp lacking spiral cords near adapical suture and the
growth lines consist of very fine lines rather than axial plicules (compare Figs. 1.E–F with Fig. 1.P–Q). Under
SEM, T. sysoevi has a distinctive sculpture of dense microscopic pustules (Fig. 1.R) while T. neocaledonica has
sparser granules (Fig. 1.G). The two species further differ in colour: T. sysoevi has a whitish to pale buff shell
lacking the brown band typically occurring on periphery of spire whorls of T. neocaledonica.
Etymology. Named after Alexander Sysoev of Zoological Museum, Moscow State University (Russia) in
recognition to his contribution to our knowledge of the turriform gastropods.

Teretia tavianii sp. nov.

Figures 2.A–G

Type material. Holotype (MZB60097) and two paratypes (paratype 1 MZB60098 (coated); paratype 2 MNHN
IM-2000-28378).
Type locality. Gulf of Aden (Indian Ocean), 11º55'95"N, 44º22'70"E to 11º55'82"N, 44º22'53"E, 795–810 m],
September 1992 [N/O “Marion Dufresne”, RED SED 92, stn 1].
Material examined. Three dd (holotype and two paratypes) from the type locality.
Description. Shell (Figs. 2.A–C) widely fusiform (b/l 0.46; a/l 0.50) with high spire, strongly excavated base
and relatively long anterior canal. Teleoconch up to about 3½ whorls, with shallowly impressed, linear suture.
Subsutural ramp rather wide, concave. Spiral sculpture on first teleoconch whorl consisting of a prominent
peripheral cord below mid-height of the whorl and two closely spaced weak spiral cords bordering abapical part of
subsutural ramp (the abapical one much stronger). On subsequent whorls a second cord develops at level of
abapical suture; last whorl with one spiral thread in the interpace between two cords. Adapical part of subsutural
ramp with a weak spiral cord near suture. Last whorl with six and nine spiral cords on base and rostrum
respectively. Teleoconch whorls sculptured by fine axial growth lines forming relatively strong, widely spaced,
arcuate plicules on subsutural ramp, rendering somewhat nodulous the spiral thread bordering adapical suture.
Under SEM, the surface is seen to be sculptured by microgranules (Fig. 2.D). Aperture lanceolate, outer lip thin.
Anal sinus deep, narrow, asymmetrical, its deepest point on subsutural ramp (Fig. 2.B). Protoconch of three whorls;

564 · Zootaxa 3911 (4) © 2015 Magnolia Press MORASSI & BONFITTO
first whorl tilted with microgranules arranged in spiral rows, subsequent whorls with diagonally cancellate
sculpture terminating a short distance above abapical suture where fine slightly prosocline riblets occur. Maximum
diameter about 0.38 mm. Shell white, protoconch brown. Dimensions: Holotype 3.2 x 1.5 mm, aperture height 1.6
mm.
Remarks. Teretia tavianii sp. nov. differs from the Southern Africa Teretia acus (Barnard, 1958) in having
fewer, less prominent, spiral cords (two versus three on spire whorls) and in number of protoconch whorls (3 rather
than 4½). T.tavianii sp. nov. differs from T. anceps and T. teres in possessing fewer, less prominent cords (two
versus respectively 4–5 and 3–4 main cords on spire whorls; compare Figs. 2.C–D and Fig. 3.B–C) and in
protoconch features (compare Figs. 2.E–G with Figs. 3. D–F). More in detail, T. tavianii sp. nov. has fewer
protoconch whorls (3 rather than 4–4 ½) and while in the new species the diagonally cancellate sculpture tends to
be restricted to whorl periphery, in both T. anceps and T. teres it occurs on whorl surface except for a narrow area
near adapical suture (compare Fig. 2.E with Fig. 3.D). Teretia fusianceps Nordsieck, 1972 is distinguished from T.
tavianii sp. nov.in having more prominent spiral cords and a more narrowly fusiform shell.
Etymology. Named after Marco Taviani, Geologist of Institute of Marine Science, National Research Council
ISMAR-CNR, that made possible the participation of the one of the authors (AB) to the cruise RED SED ’92.

Teretia tongaensis sp. nov

Figures 2.H–N

Type material. Holotype (MNHN IM-2000-28379).

Type locality. Tonga Islands, 20°42' S 174°54' W 650–676 m [BORDEAU 2 stn DW 1553]
Material examined. One dd (holotype) from type locality.
Description. Shell fusiform (Figs. 2.H–J) (b/l 0.40; a/l 0.44) with high spire, strongly excavated base and
relatively short, broad, nearly straight, siphonal canal. Teleoconch of up to 5½ whorls with shallowly impressed,
linear suture. Subsutural ramp wide, shallowly concave. Earlier two teleoconch whorls with two weak cords
bordering abapical part of subsutural ramp; a relatively prominent peripheral cord and a second main cord at level
of abapical suture. On third whorl abapical cord on subsutural ramp becomes bisected, and two additional weak
cords appear: one on abapical part of ramp, the other between the two main cords; from this whorl onwards main
cords only slightly stronger than weaker cords, with the peripheral cord at about mid- height of the whorl. On
penultimate whorl additional weak cords appear: two in the interspace between main cords and two in the abapical
part of whorl near suture. Last whorl with one interstitial spiral thread in each interspace between cords; there are
about 28 spirals on base and rostrum. Subsutural ramp with two spiral threads bordering adapical suture.
Teleoconch whorls sculptured by fine axial growth lines forming relatively strong, widely spaced, arcuate plicules
on subsutural ramp, rendering somewhat nodulous the two spiral threads bordering adapical suture. Under SEM
(Fig. 2.N) surface between spiral cords with a sculpture of microscopic pustules. Aperture oblanceolate, with outer
lip simple and fragile, curved in lateral view and forming a very deep, asymmetrical anal sinus, its deepest point on
sutural ramp (Fig. 2.J). Protoconch damaged of 3+ whorls; tip missing, subsequent whorls with diagonally
cancellate sculpture terminating a short distance above suture, abapical part of whorl occupied by fine prosocline
riblets (Figs. 2.O–P). Maximum diameter about 0.48 mm. Shell yellowish-white with a narrow pale yellowish-
brown peripheral band on three earlier whorls. Protoconch yellowish-brown. Dimensions: Holotype 8.2 x 3.4 mm,
aperture height 3.4 mm.
Remarks. Although presently known only from a single specimen, this species in our opinion warrants formal
description because of its distinctive morphological features. T.tongaensis sp. nov. differs from T. neocaledonica
sp. nov. in shape (whorls rounded rather than angulated by the peripheral cord), presence of much more numerous
and less prominent spiral cords (the peripheral cord is only barely stronger than other cords in T. tongaensis sp.
nov.) and remarkably shorter siphonal canal. Among described fossil species, T. tongaensis sp. nov. resembles
Homotoma multicingula Seguenza, 1880 from the “Pliocene” (actually Pleistocene) of Gallina (Reggio Calabria,
Italy) in shape and presence of numerous spiral cords, but the latter is much larger (15 mm in length versus 8.7
mm), with longer siphonal canal and, judging from the original description, has even more numerous spiral cords
(Seguenza, 1880: 258). Teretia guersi Schnetler, 2005 from the late Miocene of Denmark lacks angulated earlier
whorls but otherwise differs in possessing stronger and more prominent spiral cords.
Etymology. tongaensis = alluding to the fact that the type material was dredged off Tonga Islands.

NEW INDO-PACIFIC TERETIA SPECIES Zootaxa 3911 (4) © 2015 Magnolia Press · 565
FIGURE 2.A–G. Teretia tavianii sp. nov., Gulf of Aden, Cruise RED SED 92, stn RS1. A–B. Holotype (MZB60097), 3.2 x 1.5
mm. C. Paratype 1 (MZB60098), penultimate whorl, scale bar 1mm. D. Surface microsculpture of penultimate whorl, scale bar
200 µm. E–F. Protoconch, scale bar 200 µm. G. Details of protoconch 1, scale bar 50 µm. H–N. Teretia tongaensis sp. nov.
H–N. Holotype (MNHN IM-2000-28379), Tonga, Cruise BORDAU 2, stn DW1553, 8.2 x 3.4 mm H–J. Frontal and lateral
view of holotype, scale bar 2 mm L. Penultimate whorl, scale bar 500 µm. M. Microsculpture of penultimate whorl, scale bar
500 µm. N. Microsculpture of penultimate whorl, scale bar 100 µm. O–P. Protoconch, scale bar 200 µm.

566 · Zootaxa 3911 (4) © 2015 Magnolia Press MORASSI & BONFITTO
FIGURE 3. A–F. Teretia teres (Reeve, 1844), Jonian Sea, from 39°33.3’N–18°04.7’E to 39°33.6’N–18°54.5’E, 1017–1060 m
[N/O “Bannock”, Cruise J74, stn 12] (MZB11791). A. Specimen A, shell, 6.9 x 3.3 mm. B–F. Specimen B coated B.
Penultimate whorl, scale bar 500 µm. C. Surface microsculpture of penultimate whorl, scale bar 100 µm. E–F. Protoconch,
scale bar 200 µm. G. Details of protoconch 1, scale bar 100 µm. G. †Teretia elegantissima (Foresti, 1868), Italy, Calabria,
Vallone Catrica (RC) (MZB14856) H. Pleurotomella packardi Verril, 1872, Atlantic Ocean, Cruise BIOGAS, stn CP37,
47°34’N–08°39W, 2175 m.

Discussion

Because of the close morphological resemblance between species, and the scarcity of adequate material, the
identity of most Teretia has traditionally been rather controversial and confusing (e.g., Brunetti & Vecchi (2003)
point out some misidentifications of fossil taxa). In particular, numerous authors (Bellardi, 1877; Bucquoy,
Dautzenberg & Dollfus, 1883; Chirli, 1997; Nordsieck, 1977; Poppe & Goto, 1991; Powell, 1966; Vera-Peláez,
2002) consider Teretia anceps a synonym of the recent (but ranging back to Pliocene according to Brunetti &
Vecchi, 2003) Pleurotoma teres Reeve, 1844, a species distributed from Eastern Atlantic (Norway to Angola) to the
Mediterranean Sea (WoRMS Editorial Board, 2014), while according to other authors (Bouchet &Warén, 1980;
Brunetti & Vecchi, 2003; Cipolla, 1914) these two taxa represent distinct species. Powell (1966: 126) assigned the
Atlantic recent Pleurotoma megalembryon Dautzenberg & H. Fischer, 1896 to Teretia, which according to Bouchet
& Waren (1980) belongs to genus Pleurotomella (see Bouchet & Warén, 1980 and Tucker, 2004 for references).
According to the authors, P. megalembryon superficially resembles Teretia teres in type of spiral sculpture but
otherwise differs distinctly from the latter in shape and much broader protoconch (Bouchet & Waren, 1980: 38).
However, the proper generic allocation of P. megalembryon is problematic, the species almost certainly belongs to
a genus distinct from Pleurotomella. Nordsieck (1968) proposed Azorilla with P. megalembryon selected as type
species, but this generic name was considered a synonym of Pleurotomella by Bouchet & Waren (1980). More
recently, Beu (2011:108) suggested recognition of Azorilla as a valid genus to allocate P. megalembryon
Dautzenberg & Fischer, 1896 and P. lottae Verrill, 1885.
Actually, the genus Pleurotomella, as used in current literature (Beu, 2011; Bouchet & Waren, 1980; Powell,
1966), is a very heterogeneous, almost undifferentiated, assortment of species grouped together entirely on the
basis of poorly defined shell features. Pleurotomella packardi Verrill, 1872 (fig. 3.H), type species of
Pleurotomella, differ distinctly from the species here assigned to Teretia in having swollen whorls, very different
sculpture which is predominantly axial rather spiral, shape (biconic vs fusiform), protoconch shape (rather broad vs
narrowly conical) and anal sinus features (see Bouchet & Warén, 1980). Similar morphological discrepancies are
detected comparing the recent species of Teretia with members of other raphitomine genera such Gymnobela
Verrill, 1884, Thesbia Jeffreys, 1867, Phymorhynchus Dall, 1908 or Xanthodaphne Powell, 1942 (see Bouchet &
Waren, 1980; Powell, 1966 for generic diagnosis).

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 Based on teleoconch sculpture, Teretia species can be assigned to different subgroups: a “Teretia anceps-
group” (including T. anceps, T. fusianceps, T. policarinarum, T. pentacarinifera, T. intermedia and the living T.
teres and T. acus) that is characterized by species with prominent, somewhat lamelliform, spiral cords (the
peripheral cord equal to slightly stronger than other cords), and a “T. turritelloides-group” (including species such
T. turritelloides, T. elegantissima (Fig. 3.G) and T. monterosatoi) that have a peripheral cord stronger than other
cords sometimes projecting like a keel and rendering angular the whorl profile. According to Brunetti & Vecchi,
2003) T. turritelloides and T. elegantissima (but not T. monterosatoi) have a well developed microscopic sculpture
of pustules or granules on shell surface, while Teretia anceps and T. teres have a smooth surface. In contrast to this
statement, under SEM the surface of T. teres is seen to be covered by a microscopic sculpture of granules. A
possible third subgroup includes species such Teretia guersi and Homotoma multicingula.
Two of the new species here discussed, namely Teretia neocaledonica sp. nov. and T. sysoevi sp. nov., are
morphologically comparable with some members of the “T. turritelloides-group” in type of spiral sculpture (a
prominent peripheral cord and two weak cords on subsutural ramp) and microsculptural features. Compared to
their fossil congeners, Teretia neocaledonica sp. nov. and T. sysoevi sp. nov., have a less prominent peripheral cord
but the significance of this difference is uncertain.
A similar situation occurs, for example, in the raphitomine genus Cryptodaphne Powell, 1942. Cryptodaphne
pseudodrillia Powell, 1942, type species of the genus, has distinctly keeled whorls while in the recent
Cryptodaphne rugosa Sysoev, 1997 the peripheral angulation varies from distinct to weak (“in the paratypes […]
the strength and relative position of the main spiral keel vary a little”) according to Sysoev, 1997: p.335) and in C.
kilburni Morassi & Bonfitto, 2006, an otherwise “typical” Cryptodaphne species, there is no evident peripheral
angulation.
Teretia tavianii sp. nov. has relatively prominent spiral cords and a (relatively) scarcely developed microscopic
sculpture of granules as for members of the “Teretia anceps-group”. Teretia tongaensis sp. nov. has a sculpture of
numerous, not very prominent spiral cords and relatively well developed microscopic sculpture on shell surface;
these features suggest that it probably belongs to the “T. guersi-group”.
The presence of different morphological subgroups within the genus and the relatively less prominent spiral
sculpture in the recent species may suggest that Teretia, as presently construed, in polyphyletic. However, in the
absence of soft parts for anatomical and/or molecular studies, we consider preferable assign the new species to
Teretia on the basis of general resemblance in shell features refraining from proposing new and doubtful genera or
subgenera. Similarly the allocation of the new Indo-Pacific species in any other raphitomine group such, for
example, genus Pleurotomella, should in our opinion be rejected on the basis of clear morphological discrepancies.
The resemblance between the recent Indo-Pacific species here discussed and members of Teretia from the
Tertiary of Europe suggests a Tethyan origin for the genus. Despite extensive survey in literature we were unable to
track records of Teretia species in the Tertiary of the Indo-Pacific region which may suggest the hypothesis that
Teretia the genus had a wide west-east distribution, i.e. from western sectors to eastern ones of the large
biogeographic unit defined Tethys Realm by Harzhauser et al. (2002), until the late Early Miocene, when the final
breakdown of the eastern sectors (Atlantic and Mediterranean) from the eastern ones (Indo-Pacific) occurred (Rögl,
1998; Harzhauser et al., 2002).

Acknowledgments

We thank Kai Ingemann Schnetler for providing the description of Teretia guersi and Philippe Bouchet for the loan
of part of the material described and discussed in the present paper. Alexander E. Fedosov and an anonymous
referee provided comments on an early draft of the present paper. This work was supported by Canziani Fund,
Department of Biological, Geological and Environmental Sciences, University of Bologna, Bologna, Italy.

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