Littorina antipodum Nevill, 1885: 143 Littorina (Austrolittorina) unifasciata antipodum Rosewater,1970: 423 Nodilittorina (Nodilittorina) antipodum Reid, 1989: 99 Nodilittorina antipodum Nodilittorina (Austrolittorina) antipodum Reid, 2002 a : 154 Austrolittorina antipodum Littorina caerulescens Turbo caerulescens Lamarck, 1822 Melarhaphe neritoides A. unifasciata Afrolittorina praetermissa A. africana Littoraria mauritiana M. neritoides A. unifasciata Littorina mauritiana Phasianella mauritiana Lamarck, 1822 Littoraria mauritiana A. unifasciata Littorina (Melarhaphe) mauritiana Tryon, 1887: 247 Melaraphe A. unifasciata A. cincta L. mauritiana Echinolittorina ziczac Litorina (Melarhaphe) mauritiana Suter, 1913: 188 A. unifasciata L. mauritiana Melaraphe Littorina mauritiana Litorina mauritiana Bucknill, 1924: 37 Melarhaphe unifasciata Iredale, 1915: 447 Littorina unifasciata Gray, 1826 A. unifasciata A. unifasciata Oliver, 1915: 518 Melarhaphe zelandiae Finlay, 1928: 241 A. cincta Melarhaphe oliveri Finlay, 1930: 224–225 Powell, 1937: 67 Powell, 1962: 83 Melarapha oliveri Dell, 1963: 225 A. unifasciata A. unifasciata L. diemenensis A. unifasciata Litorina antipodum L. unifasciata A. unifasciata L. mauritiana Melarhaphe unifasciata M. zelandiae A. cincta M. oliveri L. unifasciata antipodum A. unifasciata A. antipodum The Subfamily Littorininae (Gastropoda: Littorinidae) in the Temperate Southern Hemisphere: The Genera Nodilittorina, Austrolittorina and Afrolittorina Reid, DG Williams, ST Records of the Australian Museum 2004 56 75 122 JXB7 (Philippi, 1847) Philippi 1847 [860,1355,1020,1044] Gastropoda Littorinidae Austrolittorina Animalia Littorinimorpha 17 92 Mollusca species antipodum   Figs. 4E,F, 9–11   Littorina diemenensisQuoy & Gaimard, 1833: 479–480(in part; not  L. diemenensisQuoy & Gaimard, 1833=  A. unifasciata). Reeve, 1858:  Littorinasp.pl.17, fig. 94 (not Quoy & Gaimard, 1833). Hutton, 1878: 27(not Quoy & Gaimard, 1833).   Litorina diemensis.—Philippi, 1847:195,  Litorinapl. 4, fig. 1 (not  L. diemensisPhilippi, 1847, emendation of  Littorina diemenensisQuoy & Gaimard, 1833=  A. unifasciata; in part; includes  A. unifasciata). Weinkauff, 1883:218(in part;includes  A.unifasciata, A. cincta, Echinolittorina novaezelandiae).   Litorina antipodumPhilippi, 1847: 195–196,  Litorinapl. 4, fig. 2 ( New Zealand; lectotype, here designated,Philippi, 1847:  Litorinapl. 4, fig. 2; fig. 9D herein). Weinkauff, 1882: 61–62, pl. 8, fig. 4.   Littorina (Melarhaphe) antipodum.—Adams & Adams, 1854: 314 (as  Melaraphe).     Littorina antipodum.—  Nevill, 1885: 143.    Littorina (Austrolittorina) unifasciata antipodum.—  Rosewater,1970: 423(as antipoda), 470–471, pl. 359, figs. 6–8, pl. 360, fig. 5, pl. 362 (distribution). Powell, 1976: 87, pl. 16, fig. 12. Powell, 1979: 87, pl. 23, fig. 9.    Nodilittorina (Nodilittorina) antipodum.—  Reid, 1989: 99.   Nodilittorina antipodum.—Spencer & Willan, 1996: 18.    Nodilittorina (Austrolittorina) antipodum.—  Reid, 2002 a: 154.   Austrolittorina antipodum.— Williams et al., 2003.   Littorina caerulescens.— Tenison Woods, 1879: 65–72(as coerulescens; not  Turbo caerulescens Lamarck, 1822=  Melarhaphe neritoides; in part; includes  A. unifasciata, Afrolittorina praetermissa, A. africana, Littoraria mauritiana, M. neritoides). Hutton, 1880: 79(not Lamarck, 1822; in part; includes  A. unifasciata). Hutton, 1882: 164, pl. 7, fig. E (radula; not Lamarck, 1822).   Littorina mauritiana.—Smith, 1884: 60–61 (not  Phasianella mauritiana Lamarck, 1822=  Littoraria mauritiana; in part; includes  A. unifasciata). Suter, 1901: 214(not Lamarck, 1822). Iredale, 1910: 70–71(not Lamarck, 1822).    Littorina (Melarhaphe) mauritiana.—  Tryon, 1887: 247(as  Melaraphe; not Lamarck, 1822; includes  A. unifasciata, A. cincta, L. mauritiana, Echinolittorina ziczac).    Litorina (Melarhaphe) mauritiana.—  Suter, 1913: 188, pl. 38, fig. 28 (not Lamarck, 1822; includes  A. unifasciata, L. mauritiana; as  Melaraphe; as  Littorina mauritianain Atlas, 1915).    Litorina mauritiana.—  Bucknill, 1924: 37, pl. 7, fig. 22.    Melarhaphe unifasciata.—  Iredale, 1915: 447(not  Littorina unifasciata Gray, 1826=  A. unifasciata; in part; includes  A. unifasciata).  Oliver, 1915: 518(not Gray, 1826).    Melarhaphe zelandiae.—  Finlay, 1928: 241(not Finlay, 1927=  A. cincta).     Melarhaphe oliveri Finlay, 1930: 224–225(Hampden, east coast of South Island, New Zealand; holotype AucklandMuseum AK 70456 (formerly TM456) plus 1 paratypeAK 72574 (formerly TM456), not seen).  Powell, 1937: 67, pl. 9, fig. 12.  Powell, 1962: 83, pl. 9, fig. 12 (as Melarhapha). Morton, 1975: fig. 6a–c (radula), fig. 7a,b (stomach).    Melarapha oliveri.—  Dell, 1963: 225.  Taxonomic history. There has been a long history of inclusion of this species as a single taxon with the Australian  A. unifasciata, and sometimes also with other superficially similar white-shelled littorinids (see Taxonomic History of  A. unifasciata). Quoy & Gaimard (1833) listed the locality New Zealandfor their species  L. diemenensisand therefore evidently considered the Australian and New Zealandspecies to be the same (although the name has now been restricted to  A. unifasciata). This concept was followed by Philippi (1847), although in addition he introduced  Litorina antipodumfor smaller, taller-spired, less markedly striated shells with a dark peripheral band, found in New Zealand. This name was almost entirely neglected for 120 years (exceptions were Weinkauff, 1882; Nevill, 1885), until used as a subspecies of  L. unifasciataby Rosewater (1970). Instead, the species was generally combined with  A. unifasciataunder the name  L. mauritiana(Smith, 1884; Suter, 1901, 1913; Iredale, 1910; Bucknill, 1924). Following Iredale (1915), the name  Melarhaphe unifasciatawas briefly used in New Zealand( Oliver, 1915). Iredale (1915)had also suggested that “the Neozelanic shell may prove subspecifically separable”, which induced Finlay (1927)to introduce the new name  M. zelandiae. However, as Finlay (1930)later confessed: “Owing to a curious and unpardonable misconception of species I described the wrong shell” (the description applied to the other New Zealandlittorinid,  A. cincta). He therefore introduced yet another name,  M. oliveri, for the present species, which was widely used in New Zealand(e.g., Powell, 1937, 1962; Dell, 1963). This was superseded by the older name  L. unifasciata antipodumfollowing the revision by Rosewater (1970). The taxon was given full specific status in the worldwide family list of Reid (1989), although the anatomical evidence justifying this decision was not presented. Rudman (1996)noted that the differences between the egg capsules of  A. unifasciataand  A. antipodumsupported separate species status for both.  No type specimens of  Litorina antipodumare known to exist, and to obviate any further confusion as to the identity of the taxon Philippi’s (1847) figure is here designated lectotype. The shape and coloration of the shell leave no doubt as to its identity ( Fig. 9D). The specific epithet is a noun in the genitive case, not an adjective that should agree in gender with the feminine generic name (e.g., Rudman, 1996). It may be noted that  Littorina novaezelandiae Reeve, 1857was mistakenly said to come from New Zealandin its original description; it is a valid species of  Echinolittorinathat is endemic to Sri Lanka( Reid, 2001).   Diagnosis. Shell small, usually tall-spired, often smooth; white with broad band of blue-grey above periphery. Penis with smooth, slightly swollen filament; glandular disc and mamilliform gland on base. Pallial oviduct with two consecutive loops of egg groove, in albumen and capsule glands.   Material examined. 76 lots (15 AMS, 18 USNM, 14 BMNH, 4 IRSNB, 4 ZMA, 1 MNHNP, 1 NMW, 19 MNZ), including 11 penes, 5 sperm samples, 7 pallial oviducts, 3 radulae.  Shell( Fig. 9). Mature shell height 5.4–13.7 mm. Shape highturbinate to tall-spired (H/B = 1.36–2.20; SH = 1.43–2.63, exceptionally 3.22); spire outline straight to slightly convex; whorls gently rounded, suture slightly impressed, periphery slightly angled; solid. Columella pillar short, slightly concave; columella slightly excavated; small eroded parietal area sometimes present. Sculpture of 9–12 approximately equidistant primary spiral grooves (incised lines) may be present above periphery, continuing faintly on base, but frequently surface is almost smooth; microstriae absent; surface, especially spire, often eroded; growth lines weak. Protoconch rarely preserved, about 3 whorls, 375–425 µm length, sculptured by spiral rows of minute tubercles ( Pilkington, 1971). Colour white, with broad spiral bluegrey band above periphery; rarely grey-black with white base, or almost entirely white with faint grey band; small juveniles entirely black or brown, with spiral white line on base; aperture dark brown with basal white band.  Animal. Head, tentacles and sides of foot black. Opercular ratio 0.47–0.61. Penis ( Fig. 10A–E): filament 0.5 total length of penis, smooth, bluntly pointed, slightly swollen, sperm groove opens slightly subterminally; single large, often narrow, mamilliform gland and adjacent large flap of penial glandular disc borne together on short lateral branch of base; penial base often with black pigment. Euspermatozoa 75– 82 µm; paraspermatozoa ( Fig. 10H,I) spherical, 11–14 µm diameter, containing large spherical granules and a curved, twisted or torus-shaped rod body. Pallial oviduct ( Fig. 10F) with simple loop of albumen gland, followed by large, almost circular loop of capsule gland, within which portion adjacent to egg groove (translucent capsule gland) is differentiated as a ring; copulatory bursa confined to straight section of pallial oviduct, opening near anterior end.  Spawn and development. Spawn ( Fig. 10G) a transparent pelagic capsule 200–250 µm diameter, containing a single ovum 80 µm diameter, capsule with hemispherical dome sculptured by 3 concentric rings, peripheral ridge and a basal circumferential flange; development planktotrophic, inferred planktonic life of 1–2 months; spawning season November to March (at Otago; Pilkington, 1971).  Radula( Fig. 4E,F). Relative radular length 2.0–2.5. Rachidian: length/width 1.13–1.33; major cusp elongate, rounded or slightly pointed at tip. Lateral and inner marginal: major cusps large, elongate, bluntly rounded at tip. Outer marginal: 6–7 cusps.  Fig. 9.  Austrolittorina antipodum: ( A) Auckland Harbour, New Zealand (BMNH 20030405). ( B,F) Muriwai, New Zealand (BMNH 20030406). ( C,E) Island Bay, Cook Strait, New Zealand (USNM 671202). ( D) lectotype figure of  Litorina antipodumPhilippi, 1847(Philippi, 1847:  Litorinapl. 4, fig. 2). ( G) Long Beach, Russell, New Zealand (BMNH 20030407). ( H) Great Island, Three Kings Islands, New Zealand (MNZ M049237). ( I) Raoul Island, Kermadec Islands (MNZ M214380). ( J) Waitangi, Chatham Islands, New Zealand (MNZ M110492).  Fig. 10.  Austrolittorina antipodum: ( A–E) Penes. ( F) Pallial oviduct. ( G) Egg capsule (after Pilkington, 1971). ( H,I) Paraspermatozoa from two specimens. ( A–C,F,H) Wellington Harbour, New Zealand (BMNH 20030394). ( D,E,I) Kaikoura, New Zealand (BMNH 20030408). ( G) Portobello, Otago Harbour, New Zealand. Shell heights: ( A) 7.5 mm, ( B) 7.4 mm, ( C) 7.4 mm, ( D) 7.4 mm, ( E) 8.9 mm, ( F) 9.8 mm. Abbreviation: ( os) subterminal opening of penial sperm groove. Shading conventions as in Fig. 3.  Habitat. Throughout much of New Zealand, especially in the north, this species is abundant on rocky shores from the top of the littoral fringe to the upper part of the barnacle zone ( EHWSTto MTL). Juveniles occur below adults, often among barnacles. The upper limit depends upon shade and spray, so that on the most strongly exposed, shaded vertical cliffs, it can reach 12 mabove EHWST. It is less common on sheltered shores, scarce in extreme shelter, and absent where water is turbid. Shaded rock faces are favoured; on sunny faces, it occupies cracks and crevices and it may occur in shallow, high-level rock pools ( Dellow, 1950; Knox, 1953; Foster, 1966; Morton & Miller, 1968). In the south of South Island  A. antipodumis less common and occupies a more restricted vertical range, confined to the littoral fringe, and its distribution is shifted towards more sheltered shores ( Batham, 1956, 1958; Morton & Miller, 1968; Pilkington, 1971). At Rangitoto Island (near Auckland), it has been recorded among  Salicorniaand lava blocks on stiff mud ( Powell, 1933a). The diet included black lichens, diatoms,  Ulvaand  Enteromorpha( Powell, 1933a; Morton, 1975). It is frequently sympatric with  A. cincta(see Habitat of  A. cinctafor comparison).  Range( Fig. 11). New Zealand, Chatham Islands, Kermadec Islands. This species occurs around the entire coastline of North and South Islands, and is also recorded from Stewart Island (Ringaringa, 46°44'S 168°09'E, MNZM019600), Snares Islands(North Island, 48°02'S 166°36'E, MNZM015474), Chatham Islands(Waitangi Bay, 43°57'S 176°33'W, MNZM110471; Powell, 1933), Three Kings Islands(Great Island, 34°09'S 172°09'E, MNZ049237) and Kermadec Islands(Raoul Island, 29°16'S 177°55'W, MNZM 214380, AMS). The species is reported to be rare in the Kermadec Islands( Iredale, 1910; Oliver, 1915; Brook, 1998); it is likely that only chance migrants reach these islands and that the population is not self-sustaining.   Remarks. Although superficially similar to the Australian  A. unifasciata, these two species should not be confused.  Austrolittorina antipodumis smaller, reaching only 13.7 mmin height (  A. unifasciataattains 24.9 mm), more tall-spired, the spiral striae are less marked and often obsolete, microstriae are absent, the peripheral blue-grey band is more conspicuous, and the penial filament is shorter and not wrinkled. In addition, their ranges are allopatric. Throughout most of its range  A. antipodumis sympatric with  A. cincta, although it has a more northern distribution. The latter is readily separated by the brown spiral lines of its shell and larger size (to 20.2 mm); anatomically the two are similar, the penial filament of  A. cinctabeing only slightly more pointed. Occasionally shells of  A. antipodumare found that are extremely elongate (H/B up to 2.21, SH up to 3.22; Fig. 9J); by analogy with other species (see Remarks on  Afrolittorina knysnaensis) these probably represent specimens infected with trematode parasites, although this has not been confirmed by dissection. Molecular data have shown that  A. antipodumand  A. cinctaare sister species, with an estimated divergence time of more than 10 million years. Possible causes of this speciation are unknown; it is too recent to be ascribed to the Oligocene transgression that greatly reduced the land area of New Zealand(Cooper & Millener, 1992). In turn, this pair is sister to the pair  A. unifasciataplus  A. fernandezensis, with a separation estimated as 23–44 million years ( Williams et al., 2003). This is younger than the attainment of the present width of the TasmanSea, implying that dispersal took place from Australiato New Zealand.