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BREVI 






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US ISSN 0006-9698 



Cambridge, Mass. 



7 February 2011 



Number 521 



DESCRIPTION OF TWO NEW SPECIES OF MUREX S.S. (MOLLUSCA: 
GASTROPODA: MURICIDAE) FROM THE NORTHERN INDIAN OCEAN 

Roland Houart 1 

Abstract. Two new species of Murex s.s. are described: Murex echinodes from Kuwait and Murex indicus from 
western India. Both species are compared with related species from Somalia, the Red Sea, Oman, and India. 

Key words: Mollusca; Gastropoda; Murex s.s; Indian Ocean; new species 



Many new species of Murex s.s. have been 
described since the Indo-Pacific species of 
Murex s.s. were revised by Ponder and Vokes 
(1988), namely, M. (M.) hystricosus Houart 
and Dharma, 2001; Murex megapex Neu- 
bert, 1998; M. (M.) philippinensis Parth, 
1994; M. (M.) salomonensis Parth, 1994; M. 
(M) somalicus Parth, 1990; M. (M.) spinas- 
treptos Houart, 2010; and M. (M.) spinifer 
Heiman and Mienis, 2010. Another species is 
being described from Taiwan (Houart, 
2010b, in press). 

Although the two species named herein 
were partially illustrated by Ponder and 
Vokes (1988), they were misidentified and 
figured under other names. Since that 



1 Research Associate, Institut royal des Sciences natur- 
elles de Belgique, rue Vautier, 29, 1000 Bruxelles, 
Belgium; e-mail: roland.houart@skynet.be 



revision was published, the definition of 
structural homologies through spiral cord 
morphology and their ontogeny was com- 
pletely revised by Merle (1999, 2001, 2005). 
This new definition of the morphology of the 
spiral cords was also applied in recently 
described species of Murex s.s. by Houart 
and Dharma (2001) and Houart (2010a, b). 

MATERIALS AND METHODS 

The material used in this study belongs 
mainly to the collection of the Museum of 
Comparative Zoology and to the private 
collection of the author. Other comparative 
type material used here is housed in the 
Museum national d'Histoire naturelle, Paris, 
France, and in the Senckenberg Museum, 
Frankfurt, Germany. 



The President and Fellows of Harvard College 2011. 



BREVIORA 



No. 521 



Table 1 . Terminology used to describe the spiral 

cords (after Merle 1999 and 2001). Terminology in 

parentheses: erratic feature. 

P primary cord 

s secondary cord 

t tertiary cord 

ad adapical (or adapertural) 

ab abapical (or abapertural) 

SP subsutural cord 

IP infrasutural primary cord (primary cord on 

subsutural ramp) 
adis adapical infrasutural secondary cord (on 

subsutural ramp) 
abis abapical infrasutural secondary cord (on 

subsutural ramp) 
PI shoulder cord 

P2-P6 primary cords of the convex part of the 

teleoconch whorl 
sl-s6 secondary cords of the convex part of the 

teleoconch whorl (example: si = 

secondary cord between Pi and P2; s2 = 

secondary cord between P2 and P3, etc.) 
ADP adapertural primary cord on the siphonal 

canal 
MP median primary cord on the siphonal canal 

ABP abapertural primary cord on the siphonal 

canal 
EABP extreme abapertural primary cord on the 

siphonal canal (Example: EABP2 = 

between EABP1 and EABP3) 
ads adapertural secondary cord on the siphonal 

canal 
ms median secondary cord on the siphonal 

canal 
abs abapertural secondary cord on the siphonal 

canal 
eabs extreme abapertural secondary cord on the 

siphonal canal (Example: eabsl = 

secondary cord between EABP1 and 

EABP2) 

infrasutural denticle 
abapical denticles 



The identification of shell characters and 
the ontogeny of spiral cord morphology and 
position on the shell (Table 1; Fig. 1) are 
mainly based on Merle (1999, 2001, 2005) 
and Merle and Houart (2003). 



DEPOSITORY 

IRSNB: Institut royal des Sciences naturelles 
de Belgique, Bruxelles, Belgium. 

MCZ: Museum of Comparative Zoology, 
Harvard University, Cambridge, Massa- 
chusetts, U.S.A. 

MNHN: Museum national d'Histoire natur- 
elle, Paris, France. 

RH: Collection of the author. 

SMF: Senckenberg Museum, Frankfurt, 
Germany. 

Murex {Murex) echinodes, new species 

Figures 1, 2-5, 9, 18, 40; Table 2 
Murex scolopax — Bosch and Bosch, 1982: 
90, text fig.; Bosch and Bosch, 1989: 58, 
text fig.; Ponder and Vokes, 1988: 49 (in 
part), figs. 76A (only), 84A (only), 85E; 
Coulombel, 1994: 67, text fig.; Bosch et ah, 
1995: 117, fig. 465 (not Murex scolopax). 
Murex megapex — Robin, 2008: 239, fig. 7 
(not Murex megapex). 

Type Material. Kuwait, 138.6 mm (lv), 
holotype MNHN 23114, Bahrain; 1 paratype 
IRSNB IG 31624/MT.2304; 2 paratypes RH 
(lv). 

Other Material. Kalba, United Arab 
Emirates, 4 dd, coll. D. Rolfe; Damman, 
Saudi Arabia, 2 dd, RH., Bahrain, 1 lv., RH. 

Distribution. Gulf of Oman, United Arab 
Emirates, eastern Saudi Arabia, and Kuwait. 

Description. Shell large for the genus, up to 
138.6 mm in height at maturity (holotype). 
Height/width ratio, spines not included: 2.6- 
3.0. Broadly ovate, spinose, weakly tubercu- 
late. Shoulder weakly convex. 

Ivory-white or light tan with axial light 
brown flammules, more apparent on primary 
spiral cords, extending on dorsal face of 
spines. Columellar lip glossy white. Inside of 
outer apertural lip white for a short distance, 
light brown within. 

Spire high with 1.6-1.75 protoconch 
whorls and up to 7 broad, strongly convex, 
weakly nodose teleoconch whorls. Suture 



2011 



TWO NEW SPECIES OF MUREX S.S. 



s1 
s2 
s3 

s5 
s6 

ads 
ms 
abs 




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P2 
P3 

P4 
P5 

P6 

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MP 

ABP 



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EABP2 
EABP3 



Figure 1 . Terminology of spiral cord morphology in Murex echinodes n. sp. (holotype). 



impressed. Protoconch large, whorls round- 
ed, irregularly shaped. Last whorl weakly- 
angular abapically. Maximum width 1600- 
1800 urn. Terminal lip weakly raised and 
weakly curved. 

Axial sculpture of teleoconch whorls con- 
sisting of low, broad, weakly rounded varices 
with long, acute, narrowly open primary and 
secondary spines. Shoulder spine longest. 
Other axial sculpture of low intervarical ribs 
and numerous growth striae. Intervarical 



axial sculpture becoming weaker in strength 
abapically, almost disappearing on penulti- 
mate and last whorls. First teleoconch whorl 
with 7 axial lamellae, second whorl with 3 or 4 
axial lamellae and onset of varices, third with 
3 varices and 2 intervarical ribs, fourth and 
fifth with 3 varices and 2 or 3 intervarical ribs, 
sixth and last whorls with 3 varices and quite 
indistinct intervarical axial sculpture. 

Spiral sculpture of low, narrow, smooth pri- 
mary, secondary, and tertiary cords. Primary 



BREVIORA 



No. 521 




Figures 2-9. (2-5, 9) Murex echinodes new species; (2, 3) Kuwait, 138.6 mm, holotype MNHN 23114; (4, 5) 
Kuwait, 135.7 mm, paratype coll. RH; (9) Protoconch (holotype MNHN). (6-8) Murex megapex Neubert, 1998, Gulf 
of Aden, approximately 60 nm SW of Aden, 12°16.0'N, 44 o 08.5'E-12°16.0'N, 44°09.5'E, 472^79 m, 170.9 mm, 
holotype SMF 311509/1 (photos ZMF). Scale bars, 0.5 mm. 



2011 



TWO NEW SPECIES OF MUREX S.S. 



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BREVIORA 



No. 521 



spiral cords ending as long, acute spines on 
varices and on siphonal canal. First teleo- 
conch whorl with visible PI and P3; P2 first 
appears on second whorl, third whorl with 
adis, IP, P1-P3, more apparent on axial 
nodes; fourth and fifth whorls with adis, IP, 
abis, PI, si, P2, s2, P3; sixth whorl occasion- 
ally with additional tertiary cord between PI 
and si; last whorl with t, adis, IP, abis, PI, (t), 
si, P2, s2, P3, s3, P4, s4, P5, s5, P6, s6, ADP, 
ads, MP, ms, ABP, abs, EABP1, eabsl, 
EABP2, (EABP3). Secondary spines on the 
siphonal canal increasingly strongly bent 
ventrally. ADP and MP giving rise to longest 
spines on siphonal canal. Shoulder spine (= PI) 
longest spine of convex part of teleoconch 
whorls. P2 shortest; P4 medium sized. 

Aperture broad, ovate. Columellar lip 
narrow, smooth, rim partially erect, adherent 
along nearly adapical half. Anal notch deep, 
broad. Outer lip weakly erect, crenulated, 
with strong labral tooth between P4 and P5, 
smooth within. Siphonal canal long, 53-56% 
of total shell length, narrow, straight, weakly 
recurved dorsally and abaxially bent at tip, 
narrowly open, with acute, long spines 
decreasing in length abapically. Operculum 
dark brown, ovate with subapical nucleus 
and 15 or 16 concentric ridges. Attached 
surface with broad, thick callused rim. 

Radula unknown. 

Remarks. Murex echinodes n. sp. is part of 
a group of large-shelled Murex species, all 
occurring in the northwestern Indian Ocean 
(Somalia, Gulf of Aden, Red Sea, Arabian 
Sea, Gulf of Oman, and Kuwait). Other 
species of that group are Murex scolopax 
Dillwyn, 1817 (Figs. 10-12, 19-21); Murex 
somalicus Parth, 1990 (Figs. 13-17); and 
Murex megapex Neubert, 1998 (Figs. 6-8, 
40). 

Murex echinodes was previously illustrated 
and discussed by Ponder and Vokes (1988: 
49-51 [in part], fig. 84A only) but was then 
considered to be a form of M. scolopax with 



a "longer more adapically directed shoulder 
spine" (PI) and "with a secondary spine 
(actually P2) usually developed between the 
shoulder spine and the spine below." Addi- 
tionally, the inner lip (columellar lip) "is less 
expanded over the parietal wall." 

I do not fully agree with these authors 
when they wrote that these characters are not 
consistent. Some are not, but some are 
consistent (see below), and other characters, 
not cited by Ponder and Vokes (1988), are 
additional features that differentiate the two 
species. 

In fact, what they cited as a "secondary 
spine usually developed between the shoul- 
der spine and the spine below" is not a 
secondary spine, but a primary spine — more 
precisely, P2. In M. scolopax, P2 is apparent 
from the first teleoconch whorl, which in 
adult shells has visible PI, P2, and P3. P2 
never reaches the strength of PI and P3 on 
the abapical teleoconch whorls, but decreas- 
es in strength to be rather similar in strength 
and height to the secondary cords si and s2 
on the last teleoconch whorl, or sometimes 
slightly stronger. In M. echinodes n. sp. the 
first teleoconch whorl and the first part of 
the second whorl bear only PI and P3, P2 
starting only from the end of the second 
teleoconch whorl. Then P2 becomes stronger 
and higher, almost reaching the strength of 
PI and P3, with a short but obvious varical 
spine on the last whorl. 

The first teleoconch whorls are also 
obviously narrower and more angular in 
M. echinodes. 

In addition, although the varical spines are 
of similar strength on the siphonal canal in 
both species, those of the convex part of the 
teleoconch whorls are obviously longer and 
straighter in the new species. 

Two other differences cited by Ponder and 
Vokes (1988) are the less expanded inner lip 
(columellar lip) on the parietal wall and the 
more rounded protoconch with fewer 



2011 



TWO NEW SPECIES OF MUREX S.S. 




Figures 10-21. (10-12) Murex scolopax Dillwyn, 1817; (10, 11) near Dissei Island, 2 m, Red Sea, 146.3 mm, RH; 
(12) Dahlak, Red Sea, 116.8 mm, RH. (13-17) Murex somalicus Parth, 1990, South Somalia; (13, 14) 87.3 mm; (15) 
120 mm, RH; (16) South Somalia, RH; (17) South Somalia, RH. (18) Protoconch of Murex echinodes new species 
(holotype MNHN). (19-21) Protoconch of Murex scolopax, Near Dissei Island, 2 m, Red Sea, RH. Scale bars, 
0.5 mm. 



BREVIORA 



No. 521 



whorls. The protoconch of M. scolopax 
(Figures 19-21) is consistently larger, with 
2-2.75 strongly keeled whorls, with a small, 
flattened first whorl and a high, broad 
terminal lip. 

Murex echinodes also has more apparent, 
although weak, axial sculpture, especially on 
first teleoconch whorls, which is almost 
absent in M. scolopax. It is also narrower 
relative to its height, with a height/width 
ratio mean of 2.86 compared with 2.71 in M. 
scolopax. 

Murex echinodes also differs in color, 
having brown or grayish brown flammules 
over the entire shell, more apparent on the 
spiral cords and on the dorsal side of the 
spines, whereas M. scolopax has only uni- 
formly brown primary spiral cords and only 
rarely colored spines. 

Murex scolopax lives in the southern part 
of the Red Sea and the Gulf of Aden. The 
shell illustrated by Coulombel (1994: 67, text 
fig.) as M. scolopax is a specimen of M. 
echinodes n. sp.; however, its presence in 
Djibouti is very questionable. The shell 
illustrated by Coulombel could have been 
collected outside of Djibouti. 

Murex echinodes n. sp. differs from M. 
somalicus in having narrower and higher 
spire whorls; a more angular shell; longer, 
straighter, and more numerous spines; and 
PI being the longest spine in M. echinodes, 
whereas it is the shortest, broad and adapi- 
cally curved, in M. somalicus. Murex echi- 
nodes also has a narrow columellar lip, 
whereas it is broadly expanded in M. 
somalicus, even fused with the last whorl 
spines P5 and P6. Moreover, in M. somali- 
cus, P6 is situated on the siphonal canal, 
whereas it is situated abapically on the 
convex part of the whorl in M. echinodes, 
M. scolopax, and M. megapex. 

Murex megapex is only known from two 
specimens — the holotype and one para- 
type — both in the Senckenberg Museum. It 



has not been collected since its description. 
Murex echinodes differs consistently from M. 
megapex in having a different protoconch 
(Figs. 8, 9, 10), with comparatively smaller 
first whorls; a more angular shell; smoother, 
shallower, and lower spiral sculpture onset of 
the first teleoconch whorls (compare Figs. 8 
and 9) and comparatively broader spines. 
Etymology. Echinodes (L): prickly 

Murex (Murex) indicus new species 

Figures 22-26, 32-34, 41; Table 3 
Murex carbonnieri — Ponder and Vokes, 

1988: 22 (in part), figs. 75A, B (only); Rao, 

2003: 228, pi. 53, figs. 4-5 (not Murex 

carbonnieri). 

Type Material. India, Gujarat, 63 miles W 
of Mangrol, 2I 11'N, 69°16'E, 70-71 m, 
Anton Brunn cruise 4B Exped., stn. 21 OB, 17 
Nov. 1963, holotype MCZ 361891, 79.3 mm 
(lv); India, 78 miles SSW of Bassein, Maha- 
rashtra, 19°7'N, 71°41'E, 68-70 m, stn. 203A, 
14 Nov. 1963, 1 paratype MCZ 361894 (dd); 
India, 44 miles SSW of Veraval, Gujarat, 
20°23'N, 70°0'E, 71-79 m, stn. 206A, 15 Nov. 
1963, 1 paratype MCZ 361890 (dd); India, 
31 miles S of Dwarka, Gujarat, 21°49'N, 
68°55'E, 50-52 m, stn. 216A, 2 paratypes 
MCZ 361893 (2 dd); India, 48 miles SSW of 
Dwarka, Gujarat, 22°3'N, 68°19'E, 79-88 m, 
stn. 21 8 A, 18 Nov. 1963, 2 paratypes MCZ 
361892 (lv); India, 83 miles W of Mandiv, 
Gujarat, 22°32'N, 68°07'E, 58 m, stn. 221A, 
18 Nov. 1963, 2 paratypes MCZ 262523 (lv 
and dd); India, Bihar, 40 miles W of Arrah, 
17°54'N, 72°27'E, 46-55 m, 1 paratype 
IRSNB IG 31625/MT.2305, 1 MNHN 
231 15, 1 coll. R. Houart (all ex MCZ 361889). 

Other Material. India, 40 miles W of Arrah, 
17°54'N, 72°27'E, 46-55 m, stn. 201A, MCZ 
361889 (80 lv and dd juveniles); off India, by 
Taiwanese fishermen, 5 lv, coll. RH. 

Distribution. Arabian Sea, off W-NW 
India, 17°54'-22°32'N, 68°07'-72°27'E, liv- 
ing at 46-70 m. 



20i: 



TWO NEW SPECIES OF MUREX S.S. 




Figures 22-31. (22-26) Murex indicus new species; (22, 23) India, Gujarat, 63 miles W of Mangrol, 21°11'N, 
69°16'E, 70-71 m, 79.3 mm, holotype MCZ 361891; (24, 25) India, 48 miles SSW of Dwarka, Gujarat, 22°3'N, 
68°19'E, 79-88 m, 81.5 mm, paratype MCZ 361892; (26) off India, 83.8 mm, coll. RH. (27-30) Murex carbonnieri 
(Jousseaume, 1881); (27) Aden, Red Sea, 65.2 mm, lectotype MNHN (photo MNHN); (28) India, Puri, Orissa, 
74.6 mm, MCZ 277715; (29, 30) Singapore, 102.3 mm, coll. RH. (31) Murex forskoehlii Roding, 1798. Gulf of Suez, 
79.8 mm, coll. RH. 



10 



BREVIORA 



No. 521 




Figures 32-39. Protoconchs and early teleoconch whorls; (32-34) Murex indicus new species, India, 40 miles W 
of Arrah, 17°54'N, 72°27'E, 46-55 m, MCZ 361889; Figures 32 and 33 figured by Ponder and Vokes (1988). (35-39) 
Murex carbonnieri (Jousseaume, 1881); (35) paralectotype MNHN, Aden, Red Sea (photo A. Robin); (36) lectotype 
MNHN, Aden, Red Sea (photo A. Robin); (37, 38) Singapore (Figs. 29-30); (39) Sri Lanka, coll. RH. Scale bars, 
0.5 mm. 



Description. Shell medium-sized for the 
genus, up to 92.3 mm in height at maturity 
(coll. RH, "India"). Height/width ratio 2.5- 
2.7. Broad, nodose, spinose, shoulder weakly 
sloping, weakly convex. 

Light tan or light brown with small dark 
brown blotches on primary and secondary 
spiral cords, between axial nodes; columellar 
lip glossy white, inner side of outer lip white 
for a short distance within, with brown 
blotches between apertural crenulations, 
light brown within the aperture. 

Spire high with 3+ protoconch whorls (first 
whorl slightly broken in all the examined 



specimens). Teleoconch up to 7 broadly 
convex, weakly shouldered spinose and nodose 
whorls. Suture impressed, partially obscured 
by small axial lamellae of succeeding whorl. 
Protoconch small, conical, with a narrow weak 
keel abapically on penultimate and last whorls. 
Terminal lip thin, erect, of sinusigeral type. 

Axial sculpture of narrow axial lamellae 
on first and second whorl and of moderately 
high or high varices and intervarical nodes 
on succeeding whorls. First and second 
whorl with 8 axial lamellae; third whorl with 
2 or 3 axial lamellae and onset of varices with 
2 or 3 intervarical nodes; fourth whorl with 3 



2011 



TWO NEW SPECIES OF MUREX S.S. 



II 




Figure 40. Distribution map of Murex scolopax species group: Murex scolopax (open circle), M. megapex 
(square), M. somalicus (diamond), M. echinodes (circle). 



spinose varices and 2 high intervarical nodes; 
fifth whorl with 3 spinose varices and 3 or 4 
low intervarical nodes or nodose ridges; sixth 
whorl with 3 spinose varices and 4-6 very 
low nodose ridges, more conspicuous on 
primary and secondary spiral cords; seventh 
whorl with 3 spinose varices and 6 very low, 
almost obsolete intervarical nodose ridges, 
obvious only on primary and occasionally on 
secondary spiral cords. Other axial sculpture 
of numerous, low growth lamellae. Spiral 
sculpture of low or moderately high, narrow, 
nodose primary, secondary, and tertiary cord 
and threads. First and second whorls strong- 
ly shouldered with visible PI and P3, starting 
P2 on second whorl. Third whorl with visible 
PI, P2, and P3, starting si; P3 occasionally 
obscured by succeeding whorl. Fourth whorl 
with adis, IP, abis, PI, t, si, P2, s2, P3, s3, 
P4, s4, P5, s5, P6, s6, ABP, abs, MP, ms, 
ADP, ads, EABP1; P2 very narrow on first 3 



teleoconch whorls, weakly stronger and 
broader from fourth whorl, P6 narrowest 
primary cord; primary cords giving rise to 
acute spines; PI, P3, and P5 spines longest; 
P4 short; P2 and P6 very short or lacking. 
Fifth whorl with t, adis, IP, abis, PI, t, si, t, 
P2, t, s2, t, P3, t, s3, t, P4, t, s4, t, P5, t, s5, t, 
P6, t, s6, t, ADP, t, ads, t, MP, t, ms, t, ABP, 
t, abs, t, EABP1, t, eabsl, t, EABP2, eabs2, 
EABP3. Secondary spines giving rise to short 
spinelets; P2, P4, and P6 occasionally of 
same strength as secondary cords. Sixth and 
seventh whorls identical to fifth whorl, 
although with additional tertiary cords on 
subsutural ramp and between secondary 
cords of convex part of teleoconch whorl 
and of siphonal canal. Shoulder spine 
longest, P2 very short, P5 second longest 
spine, followed by P3. Spines on siphonal 
canal decreasing in length abapically, grad- 
ually bent ventrally. Additional spinelets 



12 



BREVIORA 



No. 521 




105 120 135 150 



Figure 41. Distribution map of Murex carbonnieri and M. indicus: Murex carbonnieri (circle), M. indicus 
(open circle). 



Table 3. Comparisons of some shell characters. 



Character 



M. indicus 



M. carbonnieri 



Protoconch 



Form of the teleoconch 
Aperture 



Intervarical axial 
sculpture 



Primary spines on the 
last teleoconch whorl 

Secondary spines on the 
last teleoconch whorl 



Conical with 3+ whorls with narrow, weak 
keel abapically, terminal lip thin, erect, 
of sinusigeral type (more than 30 
protoconchs examined). 

Broadly ovate, weakly angular. 

Broad, ovate, columellar lip narrow, 
smooth, adherent on almost 30% of 
shell adapically, erect abapically. 

Onset of varices from fourth whorl with 2 
intervarical nodes, fifth whorl with 3 or 4 
nodes or nodose ridges, sixth whorl with 
4-6 low, nodose ridges, last whorl with 6 
very low nodose ridges. 

PI longest, P2 shortest, P3 long, P4 short, 
P5 long, P6 short. 

Obvious, si shortest, s4 long, s6 longest. 
Occasional presence of tertiary spinelets. 



2.25-2.5 rounded whorls, last whorl weakly 
angular abapically, terminal lip thin, 
curved (18 protoconchs examined). 

Broadly ovate, shouldered. 

Broad, ovate, columellar lip narrow, 
smooth, adherent on almost 25-30% of 
shell adapically, erect abapically. 

Onset of varices from third whorl with 2 
intervarical nodes, fifth and sixth whorls 
with 2-4 nodes, last whorl with 3 or 4 
high, broad, nodose ridges. 

Same as for M. indicus. 

Short or almost obsolete. s6 most obvious. 
No tertiary spinelets. 



2011 



TWO NEW SPECIES OF MUREX S.S. 



13 



formed by secondary and occasionally by 
tertiary cords. 

Aperture moderately large, ovate. Colu- 
mellar lip narrow, smooth, rim partially 
erect, adherent at adapical extremity. Anal 
notch deep, broad. Outer lip weakly erect, 
crenulated with weak, broad labral tooth 
between P4 and P5. Siphonal canal long, 52- 
57% of total shell length, straight, with low 
growth lamellae over entire length, weakly 
bent abaxially at abapical extremity, nar- 
rowly open; broad and tapered adapically. 

Operculum not observed; all live-taken spec- 
imens have dried animal deep inside the shell. 

Radula and animal unknown. 

Remarks. The protoconch of Murex in- 
dicus n. sp. had already been illustrated in 
part by Ponder and Vokes (1988: fig. 75 A, 
B), who figured the protoconch of a speci- 
men housed in MCZ (361889, ex 262074) as 
M. carbonnieri (Jousseaume, 1881) (here, 
Figs. 32-34). 

Murex indicus differs from M. carbonnieri 
in having a conical, multispiral protoconch 
with a sinusigeral notch, denoting plankto- 
trophic larval development, in contrast to the 
protoconch of the typical M. carbonnieri, 
which denotes non-planktotrophic larval 
development and having fewer, more irreg- 
ular and abapically angular last whorls. The 
terminal lip of M. carbonnieri is not of 
sinusigeral type but is weakly convex and 
rather high (Fig. 36-39), as also seen in 
Ponder and Vokes (fig. 75C). 

Other differences in M. indicus are the 
thicker apertural varix, the more numerous 
tertiary cords, the more numerous varical 
spinelets originating from the secondary and 
tertiary cords, and the more numerous and 
more close-set spines on the siphonal canal. 

Murex indicus n. sp. also resembles M. 
forskoehlii Roding, 1798, from the Red Sea 
and invasive in the eastern Mediterranean. 
However, M. forskoehlii is very close to M. 
carbonnieri, with almost identical protoconch 



whorls, similar sculpture morphology of the 
teleoconch whorls, and the same color 
pattern with brown spots, mainly on the 
primary spiral cords. Both forms could be 
conspecific, although the protoconch whorls 
seem to be consistently larger and less 
angular in M. forskoehlii. Ponder and Vokes 
(1988: 33) already pointed out the close 
relationship between both species. Murex 
indicus n. sp. differs from M. forskoehlii in 
having the same shell character differences as 
with M. carbonnieri. 

Etymology. Indicus (L): from India. 

Note. Ponder and Vokes (1988) examined 
hundreds of specimens housed in dozens of 
Museums to complete their revision. Re- 
search at the same scale to determine the 
complete geographical distribution of the 
species included in the present study was an 
impossible task. However, it is interesting to 
have an idea on their current distribution. 

Because it is quite impossible to identify 
specimens correctly with lists only, the distri- 
bution maps are based on the type material 
and on specimens housed in my reference 
collection only. They cover partially, but at a 
large scale, the distribution maps published 
by Ponder and Vokes (1988: figs 13, 25). 

ACKNOWLEDGMENTS 

I am most grateful to Adam Baldinger, 
Museum of Comparative Zoology, Harvard 
University, Cambridge, Massachusetts, for 
the loan of specimens and the gift of 
paratypes; to Thierry Backeljau, Institut royal 
des Sciences naturelles de Belgique, for his 
continuous help and useful collaboration; to 
Virginie Heros, Museum national d'Histoire 
naturelle, Paris, for the digital photograph of 
the holotype of Murex carbonnieri; to Ronald 
Janssen, Senckenberg Museum, Frankfurt, for 
the digital photographs of the holotype of 
Murex megapex; to Alain Robin, Le Mesnil St. 
Denis, France, for the digital photographs of 



14 



BREVIORA 



No. 521 



the protoconch of the holotype of Murex 
carbonnieri; to Dave Rolfe, Northfleet, United 
Kingdom, for the loan of additional species of 
Murex echinodes n. sp.; and to John Wolff, 
Lancaster, Pennsylvania, U.S. A, for checking 
the English text. Finally, I am very thankful to 
the two referees, M. G. (Jerry) Harasewych and 
Winston Ponder, for their useful comments. 



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