BREVIORA Museum of Comparative Zoology NN oom — = — vy "*Addseeg datas 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 HouartT! 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: 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 "Research Associate, Institut royal des Sciences natur- elles de Belgique, rue Vautier, 29, 1000 Bruxelles, Belgium; e-mail: roland.houart@skynet.be Mollusca; Gastropoda; Murex s.s; Indian Ocean; new species 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 Muséum national d’Histoire naturelle, Paris, France, and in the Senckenberg Museum, Frankfurt, Germany. © The President and Fellows of Harvard College 2011. 2 BREVIORA TABLE |. 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) Pl shoulder cord primary cords of the convex part of the teleoconch whorl . secondary cords of the convex part of the teleoconch whorl (example: sl = secondary cord between P1 and P2; s2 = secondary cord between P2 and P3, etc.) adapertural primary cord on the siphonal canal MP median primary cord on the siphonal canal abapertural primary cord on the siphonal canal 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 extreme abapertural secondary cord on the siphonal canal (Example: eabs1 = secondary cord between EABP! and EABP2) Aperture ID infrasutural denticle D1-D6 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). No. 521 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: Muséum 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 al., 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; | paratype IRSNB IG 31624/MT.2304; 2 paratypes RH (Iv). Other Material. Kalba, United Arab Emirates, 4 dd, coll. D. Rolfe; Damman, Saudi Arabia, 2 dd, RH., Bahrain, | 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 s1 S2 s3 s5 s6 ads ms abs eabs1 Figure 1. impressed. Protoconch large, whorls round- ed, irregularly shaped. Last whorl weakly angular abapically. Maximum width 1600-— 1800 um. 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 TWO NEW SPECIES OF MUREX SS. 3 Terminology of spiral cord morphology in Murex echinodes n. sp. (holotype). 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 4 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°08.5’E-12°16.0'N, 44°09.5’E, 472-479 m, 170.9 mm, holotype SMF 311509/1 (photos ZMF). Scale bars, 0.5 mm. TWO NEW SPECIES OF MUREX SS. 2011 JSOSUOT OS ‘YVIAA “‘(uayorq diy) SUC] Od pur Cg ‘adAjojoy UT yds Ajqeqoid ‘y10ys pq ‘SUOCT €d ‘Woys Tq ‘autds jsosuo] [g “s[Ioym jsv] pue YY uo jusiedde SSOT “[IOYM YIINOJ 0} puoses WOl} spvely) [esids ssojnuvis 3} SSOID YOIYM ([JOYM puooas UO Z[—-O][) SPp10d [eIDAIS “Ayjeatdeqe yore ‘AyTeordepe TTSus JO MSE-Oe Jsowlye UO JUSIOYpe “MOLIVU “YOOUIS diy 1ejjaumnyjoo ‘a}vao ‘peoig "xaAauod ATYVaM ‘9}eAO ATPeOIg ‘(adAjofoy) 309019 “uTyy diy jeuIWIaT, ‘oinjns dsop & YIM S[IOYM SIV] “YSIY ‘popunol ¢ oUON ‘jeuvo jeuoydis uo og ‘Aljeotdeqe YASUI] UI SUISBdIOUI OG—Ed ‘a1afosqo Tq ‘aulds ysoq10Ys [q "YJOOUS s[IOYM IOYIO ‘S[IOYM YAINOJ 0} puosas WOIJ SQII MOT AIDA € IO Z ‘Aypeordeqe yore ApyVom ‘ATTeordepe ITSUS JO %08 ysowye UO JUSIOYpe “YJOOUS ‘JIOyM jsv] Jo sourds 9g pure 6g JO aseq YIM pasny ‘peoiq dij r1e]jaunyjoo ‘a}vAo ‘peolg XOAU09 A[suoIs ‘papunol ‘prog ‘(poutuexa syouo0s0}01d Q) poster ATYeom “UT ‘qysres diy [VUIUIAT, ‘spIoyM podeys puv pozis Ajiepnsor1 ‘peolg “osIe] ¢°7-S 17 suON ‘Cd uey} I9}10Ys Apyeom og ‘ourds ysasuoy] Cd ‘oulds \soqloys pq ‘JOYS €d ‘191080 Td “WOYs Td "YJOOUIS JSOW]S [IOYM ISP] 01 YY TOYM YIN; 0} PUO0dIs WOIJ SQII MOT € IO Z “AyTeoideqe yooro “AT[eo1depe ISU JO MSE OE Joule uo JUIOYpe ‘YOOUWS ‘MoIIeU diy reyjaurnjos ‘ayeAo ‘perorg Jejnsue ATyeom ‘popunol ‘peoirg ‘(poururexa Sy2uo0d0}01d g) prog ‘ysry dy yeuruiay [roym ISI pous}epf “[[ewUs YIM S[JOYM poooy A[BUOIS ¢/°7—-Z “SOSUO] QS ‘a}9[OSqo JSOWTR “Yeon, ‘$d ueY} JoJIOYs ATYVOM Og ‘BUO] Cd “Woys pd “Buoy Ed \soq10Ys Tq ‘outds jsasuo] [dq “S[IOYM YOUODOITO} ISP] 0] PUOdES WOIJ SQII MOT € IO Z ‘Ayeoideqe yoors ‘ATTeardepe TPYS JO %SE VE jsoulye UO JUSIOYpR ‘YJOOUS ‘moIIeU dij 1e]Jaunyjoo ‘a}eAo ‘proig ‘rejnsue Apyeam ‘ayeAo ATpeoIg ‘(pourwmexa sysuo0s0}01d ¢) PoaInd pue poster Apyeom AI] jeurumoy “Ajjesideqe se[nsue ATYVIM [IOYM JsevT] “Iepnso1I adeyg ‘sj1oym popunol ¢/[—-9'| poym youosog[9} Ise] OU} uo soutds Arepuosas poym YyoUod09]9} ISP] OU} uo sourds Aleut g ainjdynos [BIXe [BolIeAIO}UyT ainjiody youosogyay oy} JO WO youos0}01g xadpsau ‘Wy SNIDUOS “We xpdojoos ‘We Sapoulysa “We Jojoereyo “SUALOVUVHO TIAHS AWOS AO SNOSIUVUNOZ) = °¢ ATAV EE 6 BREVIORA spiral cords ending as long, acute spines on varices and on siphonal canal. First teleo- conch whorl with visible P] and P3; P2 first appears on second whorl, third whorl with adis, IP, PlI—P3, more apparent on axial nodes; fourth and fifth whorls with adis, IP, abis, Pl, sl, P2, s2, P3; sixth whorl occasion- ally with additional tertiary cord between P1 and s1; last whorl with t, adis, IP, abis, P1, (t), sl, P2, s2, P3, s3, P4, s4, PS, s5, P6, s6, ADP, ads, MP, ms, ABP, abs, EABPI, 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 (= P1) 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 PS, 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 No. 521 a “longer more adapically directed shoulder spine” (Pl) 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 Pl, P2, and P3. P2 never reaches the strength of Pl and P3 on the abapical teleoconch whorls, but decreas- es in strength to be rather similar in strength and height to the secondary cords sl 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 Pl and P3, P2 starting only from the end of the second teleoconch whorl. Then P2 becomes stronger and higher, almost reaching the strength of P1 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 SS. 7 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. 8 BREVIORA whorls. The protoconch of M. scolopax (Figures 19-21) is consistently larger, with 22.75 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 Pl 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 ™. 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 No. 521 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, pl. 53, figs. 4-5 (not Murex carbonnieri). Type Material. India, Gujarat, 63 miles W of Mangrol, 21°11’N, 69°16’E, 70-71 m, Anton Brunn cruise 4B Exped., stn. 210B, 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. 218A, 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 23115, 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. 2011 TWO NEW SPECIES OF MUREX S:S. 9 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 15 30 45 60 75 TWO NEW SPECIES OF MUREX SS. 1] 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 P1 and P3, starting P2 on second whorl. Third whorl with visible Pl, P2, and P3, starting sl; P3 occasionally obscured by succeeding whorl. Fourth whorl with adis, IP, abis, Pl, t, sl, P2, s2, P3, s3, P4, s4, P5, s5, P6, s6é, 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; Pl, P3, and PS spines longest; P4 short; P2 and P6 very short or lacking. Fifth whorl with t, adis, IP, abis, Pl, t, sl, t, PQ tyS2et oats Soy te haat, S40 tlbont. So. C, P6, t, s6, t, ADP, t, ads, t, MP, t, ms, t, ABP, t, abs, t, EABP1, t, eabsl, t, EABP2, eabs?2, 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 15 30 45 60 75 90 105 120 135 150 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 Conical with 3+ whorls with narrow, weak 2.25—2.5 rounded whorls, last whorl weakly keel abapically, terminal lip thin, erect, angular abapically, terminal lip thin, of sinusigeral type (more than 30 curved (18 protoconchs examined). protoconchs examined). Form of the teleoconch Broadly ovate, weakly angular. Broadly ovate, shouldered. Aperture Broad, ovate, columellar lip narrow, Broad, ovate, columellar lip narrow, smooth, adherent on almost 30% of smooth, adherent on almost 25-30% of shell adapically, erect abapically. shell adapically, erect abapically. Intervarical axial Onset of varices from fourth whorl with 2. Onset of varices from third whorl with 2 sculpture intervarical nodes, fifth whorl with 3 or 4 intervarical nodes, fifth and sixth whorls nodes or nodose ridges, sixth whorl with with 2-4 nodes, last whorl with 3 or 4 4-6 low, nodose ridges, last whorl with 6 high, broad, nodose ridges. very low nodose ridges. Primary spines on the _P1 longest, P2 shortest, P3 long, P4 short, Same as for M. indicus. last teleoconch whorl _P5 long, P6 short. Secondary spines on the Obvious, s! shortest, s4 long, s6 longest. Short or almost obsolete. s6 most obvious. last teleoconch whorl Occasional presence of tertiary spinelets. No tertiary spinelets. 2011 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. 75A, 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 TWO NEW SPECIES OF MUREX SS. 13 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 Héros, Muséum 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 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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