Novocrania turbinata (Poli, 1795)

Jeffrey H Robinson, 2017, A review of all Recent species in the genus Novocrania (Craniata, Brachiopoda), Zootaxa 4329 (6), pp. 501-559 : 513-518

Novocrania turbinata (Poli, 1795)

Stratigraphic range: Early Miocene–Recent (Robinson in press)

1795: Anomia turbinata Poli. Test. ut. Sicil. eoru. hist. et anat., vol. 2, p. 189, vol. 4, pl. 30, figs. 15a–e, 21–24. 1818: Crania personata Blainville, Dict. des Scien, nat., vol. 11, p. 312.

1828: Crania ringens Höninghaus, Beit. zur Mono. der Gatt. Crania., p. 4, fig. 2a–b. 1828: Crania rostrata Höninghaus, Beit. zur Mono. der Gatt. Crania., p. 5, fig. 3a–b. 1836: Orbicula turbinata Deshayes & Edwards, Hist. nat. anim. sans vert., 7, p. 313. 1871: Crania anomala (Müller), Dall, Bull. Harv. Mus. Comp. Zoo., 3 (1), p. 34.

1888: Crania turbinata (Poli), Davidson, Trans. Lin. Soc. Lon., p. 188, pl. 27, figs. 14–23; pl. 28, fig. 8. 1977: Crania pourtalesi Dall, Cooper, Stud. Trop. Ocean, p. 53, pl. 3, fig. 18.

1979: Crania anomala (Müller), Logan, Bull. Inst. Ocean, 72, p. 29.

1981: Crania sp. Cooper, Smith. Cont. Palio., p. 11, pl. 13, fig. 2–4.

1986: Neocrania turbinata (Poli), Lee & Brunton, Bull. Nat. Hist. Mus. (Geol.), 40 (4), p 152. 2001: Novocrania turbinata (Poli), Lee & Brunton, Bull. Nat. Hist. Mus. (Geol.), 57 (1), p. 5. 2008: Novocrania sp. Bitner, Zoosystema, 30 (2), p. 424, fig. 3A–C.

2014: Novocrania anomala (Müller), Emig, Carnets de Géology, 14 (8), 159.

1862: Crania suessi Reeve, Conchologica Iconica, 13, 1 pl., 3 pp.

1986: Neocrania reevei (Reeve), Lee & Brunton, Bull. Br. Mus. Nat. Hist. (Geol.), 40, p. 152. 2001: Novocrania reevei (Reeve), Lee & Brunton, Bull. Br. Mus. Nat. His. (Geol.), 57 (1), p. 5. 2010: Novocrania reevei (Reeve), Bitner, Scient. Mar., 74 (4), p. 645, fig. 2D–K.

1863: Crania japonica Adams, Annals of Natural History, 11, p. 100.

1920: Craniscus japonicus (Adams), Dall, Proc. U. S. Nat. Mus., 57, p. 274.

1981: Craniscus japonica (Adams), Zezina, Galathea Report, Ƒolume 15, p. 9.

2002: Craniscus cf japonicus (Adams), Motchurova et al., Paleontological Research, p. 305.

2011: Novocrania japonica (Adams), Robinson & Lee, Mem. Assoc. Aust. Pal., 41, p. 27.

2014 a: Novocrania japonica (Adams). Robinson, Jour. Nat. Hist., 21–22, 1233, 1245, 1248, 1249, fig. 4F–H.

2014 b: Novocrania japonica (Adams). Robinson, Zool. Sci., 8, 542, 544.

Synonymy. I suggest the species turbinata is currently in the literature under two different craniid genera and three different species names. There are distinctive features in both the dorsal and ventral valves that, allowing for morphological variation, indicate this is a single very widespread species.

Poli (1795) first named this species as Anomia turbinata and illustrated it in colour, including specimens attached to a coral (reproduced in Emig 2014, fig. 1). Höninghaus (1828) proposed two new species, Crania ringens and C. rostrata , both of which were synonymised under C. turbinata by Dall (1871). Dall (1871, p. 34) listed a synonymy under Crania turbinata that included 5 genera and 5 species by 8 authors. However, Dall (1871, p. 32) noted that “I myself consider it a strict synonym of anomala .” Logan (1979) synonymised C. turbinata with C. anomala , but Lee & Brunton (1986) reinstated the species turbinata under the new genus name Neocrania . Lee & Brunton (2001) renamed their new genus Novocrania as Neocrania was preoccupied. Specimens from New Caledonia (NMNZ BR.001525) and Norfolk Ridge (NIWA 3248) were referred to as Novocrania turbinata in MacFarlan et al. (2009) and were listed under N. japonica in Robinson & Lee (2011) .

Reeve (1862) described and named specimens from Sydney, Australia, as Crania suessi , and noted that they resembled specimens of C. rostrata [= N. turbinata ] from West Africa. Lee & Brunton (1986) renamed the species reevei as suessi was preoccupied and placed the species in their new genus Neocrania . Bitner (2008, 2010) described and figured specimens from Fiji and New Caledonia. Robinson & Lee (2011) figured specimens from Australia and included Novocrania reevei in a short list of species synonymous with Novocrania japonica but did not formally synonymise N. reevei .

Adams (1863) described, but did not figure, a specimen from the Gotto Islands, Japan, naming it Crania japonica . Davidson (1871; 1888) redescribed and figured Adam’s type specimen. Dall (1920, p. 274) transferred this species to the genus Craniscus Dall, 1871 because of its raised anterior muscle scars and anterior septum in the dorsal valve, but noted that these features were “less emphatic” than in fossil Craniscus species (all other species in this genus are Jurassic). Hatai (1936a; 1936b; 1936c; 1939a; 1939b) listed localities around Japan where Recent Craniscus japonicus was collected during scientific dredging voyages, however, as I found five species of craniid in these collections, the identifications of Hatai must be considered unreliable (for craniids). Williams & Wright (1970) used material identified as Craniscus japonicus Dall, 1920 from Korea to represent the genus Craniscus in their study on the ventral valve structure of craniid genera and introduced the term ‘canal’ for the radiating tubes in the ventral valve. Brunton (1988) and Logan & Long (2001) noted that specimens of C. japonicus from Japan and Malaysia resembled N. turbinata from the Mediterranean. I have examined specimens of the type species of Cranicus, Jurassic Craniscus tripartitus (Münster, 1840) , from two European Institutions and photographs of specimens from another three European Institutions (for an unpublished study). Jurassic Craniscus specimens, known only from dorsal valves, have several very distinctive features not found in Recent Craniscus japonicus and the Recent species does not belong in the Jurassic genus, as suggested previously by Logan & Long (2001) and Cohen et al. (2008). Cohen et al. (2008) suggested that the species japonicus belonged in the Neoancistrocrania clade, however, Cohen et al. (2014, fig. 4) included material from Japan in the Tethys clade ( N. turbinata ) of their time-tree figure. Robinson & Lee (2011) and Robinson (2014a, 2014b) used the name Novocrania japonica without formally moving this species from the genus Craniscus . Recent material of Craniscus japonicus is here transferred to the genus Novocrania .

Cooper (1977, pl. 3, fig. 18) described and figured a specimen from the Caribbean as Crania pourtalesi . The specimen was examined (figured below) and is identified here as N. turbinata . The molecular analyses of Cohen et al. (2014) placed specimens identified as N. pourtalesi from off Jamaica and Belize into their Tethys clade ( N. turbinata ).

I propose that material assigned to Craniscus japonicus from Japan (Hatai 1940; Lee & Brunton 1986, Motchurova-Dekova et al. 2002), from off Korea (Williams & Wright 1970), one specimen from the Caribbean described and figured as N. pourtalesi (Cooper 1977) , a specimen described and figured as Crania sp. from off Madagascar (Cooper 1981), a specimen from off Madagascar identified as N. roseoradiata Jackson, 1952 (Bitner and Logan 2016, fig. 2C), material from the Mediterranean and North Atlantic described and figured as N. turbinata (Logan & Long 2001) , material from Fiji and Wallace Islands described and figured as Novocrania sp. (Bitner 2008), material from Australia and from New Caledonia described and figured as N. reevei (Bitner 2010) and from Australia as N. reevei / N. japonica (Robinson & Lee 2011) all belong to a single species: Novocrania turbinata . This species is also present off Tonga and Tahiti (specimens labelled Craniscus japonicus in the collection of University Museum, the University of Tokyo) and Bali and Sulawesi (Eric Simon pers. comm. 2013). The species names japonicus / japonica and reevei are placed in synonymy under N. turbinata .

All scale bars 2 mm unless indicated otherwise. C—Australia Museum, Sydney; NIWA—National Institite of Water and Atmospheric Research, Wellington, New Zealand ; OU—Geology Museum, University of Otago, Dunedin, New Zealand ; UMUT—Tokyo University Museum, Tokyo, Japan ; USNM—National Museum of Natural History , Smithsonian Institution, Washington, USA .

Abbreviations: aaq —anterior adductor quick-muscle scar, aas —anterior adductor slow-muscle scar, com —commissure, dmc —dorsal mantle canal, dv —dorsal valve, mp —median process, oi —oblique internal muscle scar, pa —posterior adductor muscle scar, sam —small anterior muscle scar, ss —support structure scar, sut —suture, ts —tiny anterior septum, tu —tubercles, vv —ventral valve.

All scale bars 2 mm unless indicated otherwise. NHMUK—Natural History Museum, London; NIWA—National Institite of Water and Atmospheric Research, Wellington, New Zealand ; OU—Geology Museum, University of Otago, Dunedin, New Zealand ; UMUT—Tokyo University Museum, Tokyo, Japan .

Abbreviations: aa —anterior adductor muscle scar, aaq —anterior adductor quick-muscle scar, aas —anterior adductor slowmuscle scar, ca —radiating canals, dv —dorsal valve, gl —growth layers, ir —irregular openings, laa —latent anterior adductor muscle scars, mp —median process, my —myotest, pa —posterior adductor muscle scar, pu—punctae, raa —raised anterior adductor muscle scars, ro —rostellum, vmc —ventral mantle canals, vs —valve surface, vsp —ventral spike.

Localities. N. turbinata has a very wide geographical range, from the Caribbean to the North Atlantic Ocean, the Mediterranean Sea, the Indian Ocean and the eastern and central Pacific Ocean and maybe in the Red Sea (Logan et al. 2008 as N. cf anomala ; Zuschin & Mayrhofer 2009 as Novocrania sp.). This species appears to be restricted to the lower latitudes between 45° North and 45° South. Localities are shown in Figure 2 which includes localities of material examined ( Table 5) and published localities (Appendix 3).

Stratigraphic range. Robinson (in press) gave this species a provisional stratigraphic range of Early Miocene (Otaian, 21.7–18.7 Ma) to Recent based on fossil specimens from New Zealand. Publications including figured material that may be N. turbinata ranging in age from Oligocene to Pleistocene are listed and discussed in Robinson (in press). The calibration ( Table 3) of the time-tree (App. Fig. 1 ) gives an age of ~ 23 Ma for the split within the Tethys clade between Recent specimens from the eastern and western sides of the Atlantic Ocean (node H, see below).

Type material and type locality. The location of the type specimen of N. turbinata is unknown, but the type locality is probably off the coast of Sicily at bathyal depths (Emig 2014).

Material examined. Recent specimens of N. turbinata were examined from localities off Gibraltar, Croatia, Lebanon, Tunisia, the Caribbean, Japan, the Philippines, Borneo, New Caledonia, the Kermadec Ridge (north-east of New Zealand) and Australia ( Table 5). The type material of N. reevei ( NHMUK ZB 1522) was borrowed from the Natural History Museum, London.

Description. The dorsal valve outline varies from sub-oval to sub-hexagonal to sub-quadrate, the apex is between the valve centre and the posterior margin ( Figs. 7N, 8A–E). The dorsal valve is sub-conical ( Fig. 8D), the exterior has a hummocky surface with an ornament of concentric growth rings and, in many populations, specimens may have minute pustules or spines. Specimens from Japan ( Fig. 8A), Australia ( Fig. 8B), the Kermadec Ridge ( Fig. 8C, D), France ( Fig. 8E), Lebanon ( Fig. 8F), and New Caledonia (Robinson & Lee 2011, fig. 3A, B D–H) have minute pustules or spines up to 70 µm wide at the base and 100 µm in length, adult valves may have more than 100 spines (Robinson & Lee 2011). The largest whole specimen examined was from New Caledonia, 9.5 mm long, 11.2 mm wide and 5.0 mm deep (Robinson & Lee 2011, fig. 2C, NMNZ BR.001525).

In the dorsal valve interior the posterior adductor muscle scars are sub-round to sub-oval, usually slightly convex and have punctae scattered across the surface ( Fig. 8G–J). The oblique internal muscle scars are small, suboval, lateral to the posterior adductor muscle scarsand often tilted so they are directed medially ( Fig. 8G, I). The anterior adductor muscle scars and support structure scars occur on raised pedestals ( Fig. 8G–O). These vary from low ( Fig. 8G) to high ( Fig. 8H–O). These muscle scars may face ventrally ( Fig. 8H) or be tilted towards the posterior, from gently ( Fig. 8G, I) to steeply ( Fig. 8J–L). There is a single ventrally-pointing median process at the anterior apex of the two pedestals and the small anterior muscle scars are placed on either side, facing posteriolaterally ( Fig. 8L). There is often a tiny septum running a short way anteriorly from the anterior base of the median process ( Fig. 8I, J). Anteriorly of the pedestals are the dorsal mantle canals (vascula media), often incised in larger specimens ( Fig. 8G). These branch from a single stem that emerges between the median process and the pedestals of the anterior muscles. The dorsal valve surface is densely punctate. The marginal rim is concave with tubercles on the inner edge ( Fig. 8H, I).

The ventral valve varies from being nearly flat to, more commonly, a shallow to deep bowl ( Fig. 9A–F). The posterior adductor muscle scars are large and sub-oval. The rostellum has a slim, round-topped spike ( Fig. 9A–H), the oblique internal muscles attach ventrally to the flattened sides of the spike’s rounded tip. The anterior adductor muscles attach ventrally to the rostellum, on either side of the spike ( Fig. 9F–I). The anterior adductor muscle scars may be flush with the rostellum surface ( Fig. 9A, B, E), may be convex ( Fig. 9F), or may be latent ( Fig. 9C, D, G). This variation appears to occur in all populations. The ventral mantle canals may be incised in the valve floor ( Fig. 9C, F). The marginal rim is usually convex and usually has sub-round canals showing at the surface, they may be large and very obvious ( Fig. 9A, C, D, F) or smaller and less obvious ( Fig. 9B, E), sometimes the canals open onto the valve floor where they commonly have an irregular shape ( Fig. 9C–E). There are no punctae on the ventral valve floor, but they sometimes occur on the ventral muscle scars ( Fig. 9I).

The calcitic ventral valves of craniids are composed of many thin layers, about 50 µm thick ( Fig. 9D, J). In N. turbinata the ventral valve growth layers form radiating tube-shaped canals 100–400 µm in diameter ( Fig. 9K–N) and the growth layers can be observed in the walls of the canals ( Fig. 9K). The canals begin at the valve edge as part of the rim, becoming longer as each successive layer is added to the valve ( Fig. 9K–M). Canals begun in the juvenile stages of valve growth are covered over as they become part of the valve bowl ( Fig. 9K). The 3D image ( Fig. 9N) shows the raised pedestals and process of the dorsal valve hanging above and surrounding the rostellum and ventral spike.

Ecology. In the Mediterranean N. turbinata is found in sea caves and attached to boulders and rock walls at depths ranging from 7–150 m (Logan & Long 2001). This species occurs in sea caves around the Ryuku Islands, Japan, with a depth range of shallow water down to 40 m (Saito et al. 2000). Specimens from New Caledonia attached to coral branches were collected at 24 m ( Fig. 7N, Table 5) and have been collected at 105–110 m (Bitner 2010). Specimens have been collected off Fiji and Wallace Islands between 200–420 m (Bitner 2008), and between 31–128 m around the Australian coast ( Table 5).

Remarks. Williams & Wright (1970, text fig. 14) proposed that the canals in the ventral valves of Craniscus japonicus [= N. turbinata ] were “storage centres” and illustrated the ventral mantle extending down into the canals. Lee & Brunton (1986, p. 156) suggested that the canals in the ventral valves of some species of Crania and Ancistrocrania were enlarged endopuncta and were a way of the animals building up their ventral valves without having to produce as much calcite as a solid valve would require.

Figures

FIGURE 1. Localities of Novocrania anomala (red dots) and N. altivertex (orange square). Yellow rectangles may be N. anomala or N. turbinata.

FIGURE 2. Localities of Novocrania turbinata, yellow rectangles may be N. anomala or N. turbinata.