Mipus tomlini (van Regteren Altena, 1950)

Oliverio, Marco, 2008, Coralliophilinae (Neogastropoda: Muricidae) from the southwest Pacific, Mémoires du Muséum national d'Histoire naturelle 196, pp. 481-586 : 568-585

Mipus tomlini (van Regteren Altena, 1950)

Figs 142-144

Coralliophila (Mipus) tomlini van Regteren Altena, 1950: 217-218 , text fig. 10.

Other reference:

Latiaxis sp. - Clover 1982: pl. 18, fig. 308.

TYPE MATERIAL. — Holotype RGM 46737, figured by Kosuge (1986: pl. 25, fig. 8); paratype from “horizon above layer I: sheet 110B, M273” (van Regteren Altena 1950: 217). Late Pliocene.

TYPE LOCALITY. — Java, Indonesia, “Poetjangan layers (volcanic facies): sheet 110B, C71”. Late Pliocene .

MATERIAL EXAMINED. — Indonesia. KARUBAR: stn CP 73, 360 m, 1 dd (Fig. 142).

08°29’S, 131°33’W, 840-855 m, 1 dd (Fig. 143). Solomon Islands. SALOMON 1: stn CP 1794, 09°16’S, 160°08’E, Tonga. BORDAU 2: stn DW 1583, 18°37’S, 174°03’W, 327- 494- 504 m, 1 lv (Fig. 144).

DISTRIBUTION. — Indonesia, Pliocene (Java) to Recent ( Gulf of Boni , Celebes, 492 fms: Kosuge 1986d; KARUBAR stn CP 73, 840-855 m). Tonga, one empty shell in 327-360 m, Solomon Islands, live in 494-504 m. Range extension eastward in the Pacific .

REMARKS. — Similar to Mipus eugeniae , with a very angled last whorl and fine spiral sculpture. Originally described as a Pliocene fossil from Java. Clover (1982) figured the first Recent record (USNM 239474, from Celebes), as Latiaxis

FIGS 142-144. Teleoconchs of Mipus tomlini (van Regteren Altena,1950) . 142, BORDAU 2 stn DW 1583, H 15.8 mm; 143, KARUBAR stn CP 73, H 39.4 mm; 144, SALOMON 1 stn CP 1794, H 46.4 mm.

FIGS 145-168. Protoconchs of Coralliophila . 145, C. violacea (Kiener, 1836) , LIFOU 2000 stn 1451. 146, 147, C. squamulosa (Reeve, 1846) , LAGON stn 726. 148, C. bulbiformis ( Conrad, 1837) , LIFOU 2000 stn 1432. 149, C. costularis (Lamarck, 1816) , LIFOU 2000 stn 1461. 150, C. erosa (RÖding, 1798) , LIFOU 2000 stn 1425. 151, C. candidissima n. sp., paratype MNHN 20188, LAGON stn 1432. 152, C. amirantium Smith, 1884 , LAGON stn 517. 153, C. norfolk n. sp., paratype MNHN 20195, BATHUS 3 stn CP 811. 154, C. xenophila n. sp., paratype MNHN 20205, MUSORSTOM 8 stn CP 1087. 155, C. cf. ovoidea (Kosuge, 1985) , BATHUS 2 stn DW 726. 156, C. squamosissima ( Smith, 1876) , LAGON stn 280. 157, C. monodonta (Blainville, 1832) , LIFOU 2000 stn 1451. 158, C. curta G.B. Sowerby [third of the name],1894,LIFOU 2000 stn 1429. 159, C. mitraeforma Kosuge, 1985 , VOLSMAR stn DW 41. 160, C. roseocephala Kosuge, 1986 , SMIB 8 stn DW 190. 161, Coralliophila pulchella (A. Adams, 1854) , BATHUS 1 stn DW 692. 162, C. carolae D’Attilio & Myers, 1984 , LAGON stn 933. 163, C. fimbriata (A. Adams, 1954) , LIFOU 2000 stn 1435. 164-166, C. robillardi (Lienard, 1870) , LIFOU 2000 stn 1460; 167, 168, Rhizochilus cf. antipathum Steenstrup, 1850 ; 167, NORFOLK 1 stn DW 1712; 168, LIFOU 2000 stn 1465. Scale bars 200 Μm unless otherwise indicated.

FIGS 169-192. Protoconchs of Babelomurex , Hirtomurex , Latiaxis and Rapa . 169, B. yamatoensis Kosuge, 1986 , VOLSMAR stn DW 41. 170, B. fusiformis ( Martens, 1902) , MUSORSTOM 6 stn DW 481. 171, B. depressispiratus n. sp., paratype MNHN 20214, MUSORSTOM 4 stn DW 210. 172, B. glaber Kosuge, 1998 , BORDAU 1 stn DW 1496. 173, B. armatus (G.B.Sowerby [third of the name], 1912),MUSORSTOM 7 stn DW 513. 174, B. shingomarumai (Kosuge,1981) , SMIB 8 stn DW 159. 175, B. purpuraterminus ( Kosuge, 1979) , SMIB 5 stn DW 95. 176, B. virginiae Kosuge & Oliverio,2004 , MUSORSTOM 6 stn DW 471. 177, B. pallox n. sp., paratype MNHN 20229, NORFOLK 1 stn DW 1680. 178, B. lischkeanus (Dunker, 1882) , BORDAU 2 stn DW 1543. 179, B. cf. tuberosus ( Kosuge, 1980) , SMIB 8 stn DW 182- 184. 180, B.diadema (A.Adams,1854) , SMIB 5 stn DW 95. 181, B. macrocephalus n. sp., holotype MNHN 20232,MUSORSTOM 4 stn DW 227. 182, Babelomurex kuroharai ( Habe,1970) ,MUSORSTOM 8 stn CP 1084. 183, B. kawanishii ( Kosuge,1979) ,SMIB 5 stn DW 76. 184, B. wormaldi (Powell, 1971) , CHALCAL 2 stn DW 77. 185, B.miyokoae Kosuge,1985 ,MUSORSTOM 5 stn 329. 186, Hirtomurex sp.D , CHALCAL 2 stn DW 76. 187, H.filiaregis (Kurohara,1959) ,SMIB 8 stn DW 187. 188, Hirtomurex sp.C , MUSORSTOM 4 stn DW 226. 189, Hirtomurex sp. A , BATHUS 2 stn CP 743. 190, Latiaxis pilsbryi Hirase,1908 , NORFOLK 1 stn DW 1679. 191, L. hayashii Shikama,1966 ,MUSORSTOM 5 stn 304. 192, R. rapa ( Linné, 1758) , LAGON stn 912.Scale bars 200 Μm.

FIGS 193-205. Protoconchs of Mipus ; 193, M. isosceles (Barnard, 1959) , BORDAU 1 stn DW 1497. 194, M. sugitanii Kosuge, 1985 , CHALCAL 2 stn DW 69; 195-196, M. tonganus n. sp. 195, paratype MNHN 20245, BATHUS 4 stn DW 887; 196, paratype MNHN 20246, BORDAU 1 stn CP 1506. 197, M. alis n. sp., holotype MNHN 20241, BORDAU 2 stn DW 1631. 198-205, M. boucheti n. sp., larvae, from a capsule in the holotype’s mantle cavity, MUSORSTOM stn CP 1018. Scale bars 100 Μm (193, 196, 197), 200 Μm (185-192, 194, 195), 500 Μm (192).

FIGS 206-218. Teleoconchs of Coralliophila , Babelomurex and Hirtomurex . 206, 207, Coralliophila violacea (Kiener,1836) . 206, LIFOU 2000 stn 1418, H 26.0 mm. 207, “Oceanie”, MHNG 1098/70/1, H 39.4 mm. 208, C. curta G.B. Sowerby [third of the name], 1894, LIFOU 2000 stn 1457, H 6.6 mm. 209, 210, Babelomurex yamatoensis Kosuge, 1986 . 209, VOLSMAR stn DW 41, H 23.5 mm; 210, SMIB 8 stn CP 162, H 27.6 mm. 211, 212, B. neocaledonicus Kosuge & Oliverio 2001 , holotype MNHN 0354, SMIB 4 stn DW 51, H 26.8 mm. 213, 214, B. glaber Kosuge, 1998 . 213, MUSORSTOM 8 stn CP 1017, H 22.1 mm; 214, MUSORSTOM 8 stn CP 963, H 28.7 mm. 215, 216, B. kuroharai ( Habe, 1970) , MUSORSTOM 8 stn CP 1084, H 14.9 mm. 217, B. virginiae Kosuge & Oliverio, 2004 , holotype MNHN 2190, SMIB 3 stn DW 18, H 23.3 mm. 218, Hirtomurex winckworthi ( Fulton, 1930) , BORDAU 1 stn CP 1501, H 28.7 mm.

FIGS 219, 220. Babelomurex natalabies n. sp. 219, a stamp of the New Caledonian postal service (1989) depicting the holotype, courtesy of C. Berthault. 220, holotype MNHN 20841, CHALCAL 2 stn DW 81, 23°20’S, 168°03’E, 311 m, H 16.4 mm.

sp., and Kosuge identified the USNM shell with van Regteren Altena’s taxon. Protoconch missing in all examined shells and in the holotype (Kosuge 1986: pl. 25, fig. 8) and the other Recent Indonesian shell (Kosuge 1986: fig. 7). Enigmavasum enigmaticum Poppe & Tagaro, 2005 , was described (as a new genus and new species in the Turbinellidae ) on the basis of empty shells collected in 800 m, along the coast of Cotabato (Mindanao, Philippines). The shell is very similar to M. tomlini , but to verify its status more material from the Philippines is needed. It at least raises doubts about the placement of E. enigmaticum in the Turbinellidae .

CONCLUSION

This is the first attempt to revise a local tropical fauna of Coralliophilinae on the basis of a very large number of samples (well over a thousand specimens studied).

In total, 97 coralliophiline species have been identified so far in the area of the southwest Pacific herein surveyed, representing 35-50% of the estimated 200-250 species of the subfamily globally. This is a very good representation of global diversity in this group, compared with the 240 taxa of the other muricid subfamilies recorded from the same localities, which represent only about 16% of the 1500-1550 non-coralliophiline muricids of the world (R. Houart, pers. comm.). In total, therefore, over 330 species of Muricidae , with Coralliophilinae constituting about 28%, have been recorded from the region. The coralliophiline species have been classified in 9 genera, including all currently employed genera with the exception of Emozamia , which has been recorded from the Kermadec Ridge (Marshall & Oliverio in prep.). However, to deal with supraspecific biogeography, a better assessment of coralliophiline phylogeny is needed.

Thirteen species are described as new, 4 had earlier been described (Kosuge & Oliverio 2000, 2004) from the same area, 2 have uncertain status but probably represent undescribed species ( Hirtomurex sp. C and Hirtomurex sp. D ), 3 are being described from the nearby New Zealand Exclusive Economic Zone ( Hirtomurex sp. A and Hirtomurex sp. B : Marshall & Oliverio in prep.) and 1 from the Marquesas ( Coralliophila nukuhiva: Oliverio 2008 ). This makes a total of 22 new taxa recently discovered (23% of the total) during this first revision of the southwest Pacific coralliophilines and related work (Table 1).

TABLE 1. — Summary of the distribution of the species of Coralliophilinae included in the present revision, based on data from the present work and from the literature (see text). Depth ranges are internal ranges based on live-taken specimens; †: based on empty shells only. Habitats are classified as D (deep water) and S (shallow water) based on whether the range of the species is centered below or above 150 m, respectively.

Coralliophila cancellarioidea n. sp. Southwest Pacific

Coralliophila candidissima n. sp. West Pacific

Coralliophila carnosa Kosuge, 1986 West Pacific

Coralliophila carolae D’Attilio & Myers, 1984 Indo-West Pacific

Coralliophila clathrata (A. Adams, 1854) Indo-West Pacific

Coralliophila costularis (Lamarck, 1816) Indo-West Pacific

Coralliophila crebrilamellosa (Sowerby III, 1913) Indo-West Pacific

Coralliophila curta G. B. Sowerby III, 1894 Indo-West Pacific

Coralliophila erosa (RÖding, 1798) Indo-West Pacific

Coralliophila fearnley ( Emerson & D’Attilio, 1965) West Pacific

Coralliophila fimbriata (A. Adams, 1854) Indo-West Pacific

Coralliophila cf. fritschi (Martens, 1874) Indo-West Pacific

Coralliophila inflata (Dunker in Philippi, 1847) Indo-West Pacific

Coralliophila jeffreysi Smith, 1879 West Pacific

Coralliophila mitraeforma Kosuge, 1985 West Pacific Hirtomurex kawamurai (Shikama, 1978) West Pacific

167°E 22°S 400-620 † D

164°E - 167°E 20°S 8-55 † S

135°E - 167°E 34°N - 21°S 31-236 † D

130°E - 164°E 34°n - 20°S 90-100?

30°E - 135°W 34°N - 30°S 5-20 † S

34°E - 170°E 0° - 30°S 13-40 (5-65 †) S

30°E - 168°E 34°N - 16°S 200-260 † D

56°E - 167°E 21°N - 20°S 0-20 † S

34°E - 167°E 35°N - 20°S 0-57 S

120°E - 170°E 34°N - 20- 25°S 40-48 † S

40°E - 140°W 31°N - 24°S 90-305 (12-447 †) S

18°E - 167°E 35°N - 20°S 347-373 † D

30°E - 168°E 34°N - 22°S 270-400 † D

130°E - 180°E 34°N - 40°S 28-30 S

128°E - 178°W 32°N - 24°S 245-400 D 125°E - 171°E 33°N - 23°S 402-450 † D

Coralliophila nodosa (A. Adams, 1854) West Pacific 125°E - 155°W 24°N - 24°S 230-262 † D

Coralliophila norfolk n. sp. Southwest Pacific 168°E - 174°W 19°S - 23°S 389-400 (281-523 †) D

Coralliophila nukuhiva Oliverio, 2008 Southwest Pacific 163°E - 140°W 8°S - 23°S 220-416 (+ 444 †) D

Latiaxis hayashii Shikama, 1966 West Pacific 133°E - 168°E 32°N - 24°S 280-385 † D

Mipus coriolisi Kosuge & Oliverio, 2004 Southwest Pacific

Mipus isosceles (Barnard, 1959) Indo-West Pacific

Mipus mamimarumai Kosuge, 1981 West Pacific

Mipus matsumotoi Kosuge, 1985 Indo-West Pacific

Rhizochilus cf. antipathum Steenstrup, 1850 West Pacific

Total = 97 species New species = 13

158°E - 167°E 18°S - 22°S 547-620 † D

30°E - 178°W 18°S - 30°S 335-350 † D

120°E - 167°E 33°N - 21°S 600 D

30°E - 169°E 33°N - 30°S 495-550 D

135°E - 155°W 34°N - 33°S 45-245 † D

lv: 0-600 m 25 S dd: 0-855 m 70 D 2?

In addition to the new taxa, the material contains range extensions for 37 species (38% of the total fauna): one reported for the first time from the Pacific Ocean; 14 reported for the first time from the southwest Pacific, and 22 representing significant other extensions of ranges. Along with the new taxa, this survey has thus added significant new biogeographical information for 59 species (61% of the species). This gives an idea of the poor state of knowledge of marine biodiversity and of the importance of intensive exploration using adequate means. At least 17 species (17.5%) are known only from the surveyed area, and 19 (19.5%) from a slightly larger area including northern New Zealand (to be treated by Marshall & Oliverio in prep.). Of the 89 species listed by Tsuchiya (2000) for Japanese waters, 54 (61%) are also present in the southwest Pacific area surveyed herein. Before attempting to evaluate the rate of endemism, a deeper taxonomic insight and better geographical coverage are required, given the poor level of knowledge of coralliophiline geographic patterns.

Coralliophilines have successfully radiated in deep waters, an unusual and interesting pattern for a group associated with corals. Deep habitats in the Indo-West Pacific are considered those deeper than 100-150 m, the lower limit for the hermatypic scleratinians, at which they are outcompeted for space by Alcyonacea, Stylasteridae and Porifera. Twenty-five species (26%) have been collected at shallow stations (i.e. at less that 100-150 m), including one new species. Seventy species (72%) have been sampled from deep waters (more than 100-150 m depth), including all remaining new species. A similar depth distribution is observed in other areas. Among the Japanese species, 70% are from deep waters (Tsuchiya 2000). In the eastern Atlantic at least 72% of the coralliophiline fauna inhabits deep waters (MO, unpublished data).

It would be interesting to compare the vertical trends of diversification of the coralliophilines with those of their anthozoan hosts. The azooxanthellate scleractinian fauna of Vanuatu and Wallis and Futuna studied by Cairns (1999) proved rich and diversified, particularly in deep waters. The New Caledonian fauna is suggested (Cairns 1999) to also be extremely diverse and similar to that of the Vanuatu region. More data must be gathered on geographical variation in the vertical patterns of diversity in both groups, before this comparison can be made constructively.

Planktotrophic development has been documented with certainty in 68 species (70% of the total, but 87% of the species with documented larval type). At least 10 species (10% of the total and 13% of the documented species) have a paucispiral protoconch and protoconch characters indicating either a severe shortening of the pelagic larval phase (possibly entirely lecithotrophic) or that development is entirely intracapsular. In the remaining 19 species (27% of the total) the type of development could not be ascertained for the lack of a protoconch sufficiently well preserved for study (either in the present material or in samples from the rest of the species’ range) (Table 2). Once a larger and more complete dataset is available, this feature will certainly prove important in understanding biogeographic patterns and developmental dynamics of coralliophilines. It is also very important from the taxonomic point of view (Kosuge 1986) in a subfamily in which intraspecific teleoconch variability is often high. Unfortunately – as in many other gastropod groups – most type specimens lack a useful protoconch, hampering correct identification in several instances. In the few cases in which the holotype is lost, unless other information can help in the identification process, neotype selection will be the sole way to stabilize the nomenclature. In the remainder, examination of the protoconchs in as many topotypic specimens as possible would be of great help, and the only way to solve this problem.

TABLE 2. — Protoconch characteristics (number of whorls, height and width) and inferred mode of larval development

(P: planktotrophic; NP: non-planktotrophic;?: unknown).

Hirtomurex kawamurai (Shikama, 1978) Leptoconchus sp. (spp.?)

Hirtomurex teramachii (Kuroda, 1959) - - - P

Hirtomurex winckworthi ( Fulton, 1930) 2-2.2 - - P

Magilus antiquus (Montfort, 1810)

Mipus alis n. sp.

Mipus boucheti n. sp.

Mipus coriolisi Kosuge & Oliverio, 2004

Mipus isosceles (Barnard, 1959)

Mipus mamimarumai Kosuge, 1981

Mipus matsumotoi Kosuge, 1985

Mipus sugitanii Kosuge, 1985

Mipus tomlini (Van Regteren Altena, 1950)

Mipus tonganus n. sp.

Mipus tortuosus (Azuma, 1961) - - -?

- - - P

- - -?

1.8 520 710 P

1.8 780 700 NP

- - -?

1.6 780 840 P

- - -?

- - -?

2.8 720 710 P

- - -?

1.3 680 780 NP

- - -?

ACKNOWLEDGEMENTS

I thank Philippe Bouchet (MNHN, Paris) who allowed me to revise the coralliophiline material from the French expeditions to the southwest Pacific. Financial support of the European Commission’s Research Infrastructure Action via the SYNTHESYS Project, is here acknowledged. A PARSYST visit grant permitted a visit to the MNHN and the entire malacological staff contributed in making my various visits to the MNHN both fruitful and comfortable. Particularly, Virginie Héros and Philippe Maestrati (MNHN) are heartily thanked for their assistance with sample management, locality data and recovering type material. BenoÎt Fontaine and Delphine Brabant (MNHN) and Raimondo Villa (Rome) are thanked for assistance with photography. SEM work was carried out at the LIME centre of “Roma Tre” University and Andrea Di Giulio (“Roma Tre” University) is heartily thanked for assistance. This research also benefited from a Centenary Grant (1999) from the Malacological Society of London and MURST (1999) funds for the study of geographic variation and biodiversity. Yves Finet (MHNG), Bruce Marshall (NMNZ), Hiroshi Saito (NMT), Ole S. Tendal (ZMUC), Kathie Way and Amelie MacLellan (BMNH), Ryohei Yamanishi (OMNH) and Matthias Glaubrecht (ZMB) are thanked for bibliographic support and type material information and loans. Terumichi Kimura ( Japan) provided valuable information on Japanese Coralliophilinae , Riccardo Giannuzzi-Savelli (Palermo), Bret Raines (Victorville) and Jean Tröndlé (La Force) are thanked for bibliographic support. Philippe Bouchet, Harry G. Lee (Jacksonville), Bruce

Marshall and Riccardo Giannuzzi-Savelli are thanked for discussion of nomenclatural issues. Alan Beu and Didier Merle read the manuscript critically and offered very useful hints. Virginie Héros and Robert H. Cowie did very careful editorial work, correcting a huge number of imprecisions in the text.

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