BEU (19$) Taxonomy of Gastropods of the Families Ranellidae (= Gymatiidae) and Bursidae Part 5. Early History of the Families, W

Home , Argobuccinum, Bursa (gastropod), Flowerpot Bay, Kern River Beds, Marsupina, Personella

BEU (19$)

H •«!» Kai Spec. Pub. (Prof. T. Kotaka Commem. Vol.) - j.ls. 1 August 25, 1988

Taxonomy of Gastropods of the Families Ranellidae (= Gymatiidae) and Bursidae Part 5. Early History of the Families, with Four New Genera and Recognition of the Family Personidae

< Alan G. Beu*

ABSTRACT

Thf origins and phylogeny of the Bursidae are obscure; the only recorded Eocene species are •i.^f .ahia washingtoniana (Weaver, 1912) (Late Eocene, western USA), probacy 0. domenginica i Vokvs, 1939) (Middle Eocene, western USA), and Marsupina yasila (Olsson, 1930), Late Eocene, iVru. The oldest Ranellidae are six species of Sassia in Late Cretaceous rocks; the earliest is S. biiHibcnsis (Stantoi, 1893) (Turonian, western interior, USA); most others are Maastrichtian. I'mrhytriton Meek, 1864 is transferred to the neogastropod family Buccinidae, and Sassia i; r-x'arded as the stem group of all Ranellidae. The earliest, recorded undoubted member o' lUnollinae is Gyrineum (?) judithae Zinsmeister, 1983 (Danian, Early Paleocene, California) ii|'l>arently a representative of the stem group of an Eocene radiation of Ranellinae in the north Pacific. A new species of Ranella is described from the late Paleocene Lodo Formation of ('iilifornia Western USA Eocene Ranellinae are briefly reviewed, and the new genus Ameranella in proposed for Nyctilockus kewi Dickerson, 1915 (Late Eocene, Washington and California), Fusitriton" terrysmithae Hickman, 1980 (latest Eocene, Oregon) and Triton verruculosum Sowerby, 1846 (Miocene, Chile). The earliest recorded Cymatium species are C. (Monoplex) janetae Squires, 1983 (lower Middle Eocene, California) and C. (Monoplex) washingtonianum (Weaver, 1912) (Late Eocene, Washington). The new genus Eocymatium is proposed for Murex pyraster Lamarck, 1803 (Lutetian, Paris Basin) and an unnamed Late Paleocene species from the uppermost Ranikot beds of Pakistan. The oldest recorded Distorsio species is D. ^raegranosa (Cottreau, 1922), a large, typical species from the early Campanian (Late Cretaceous) of Madagascar. Because of the appearance of Distorsw parallel to Sassia (so its inclusion in Ranellidae would make that a polyphyletic taxon) and because of its distinctive anatomy and radula, the &mall group of genera related to ftistorsio is regarded as a sixth family of Tonnoidea, Family Personidae Gray, 1854. Personella Conrad, 1865 is an earlier name for Sassia (Byramia) MacNeil, 1984 (Ranellidae) and the new genus Personopsis is proposed for the small, weakly distorted Personidae that have previously been referred to Personella. The new genus and species Kotakaia simplex is proposed for a small personid from the Mid-Late Paleocene of the Chatham Island, New Zealand.

INTRODUCTION For several years I have been working because of their very long-lived plank- inwards understanding the classification totrophic larvae, and so most species and phylogeny of the tonnoidean gas- have received separate names in each of tropod families Ranellidae ( = Cyma- the different regions they occupy. Ear- tiidae) and Bursidae. Species in these lier taxonomists included a broad group families have very wide distributions, of gastropods in Ranellidae, in particu-

• New Zealand Geological-Survey, DSIR, PO Box 30368, Lower Hutt, New Zealand. 70 A.G. Brn lar, because of the previously genera that are incorrectly placed in the unrecognised convergent development of Ranellidae. The major conclusion from prominent varices and coarse sculpture this study is that Ranellidae, as treated in several unrelated gastropod groups. until now, is a polyphyletic taxon; the This is the fifth in a series of papers personid genus Distorsio evolved at intended to revise the taxonomy of these about the same time as Sassia (the stem families and to determine their group of all other taxa included here in phylogeny (Part 4: Beu & Kay, 1988). Ranellidae) and so Personidae is regard- The present paper is a brief summary ed as a family distinct from Ranellidae. of ideas developed during 20 years of It is with great pleasure that I dedi- considering the fossil record and possible cate this paper, and the new genus phylogenies of these families. I do not Kotakaia proposed below, to my good attempt to document full phylogenies, friend and former professor (during his but nse the record of first appearances as "New Zealand years"), Professor Tamio evidence to begin sorting out Kotaka, on the occasion of his retire- phylogenies that, combined with mor- ment, and in commemoration of his asso- phology, can be used to recognize some ciation with New Zealand paleontology.

ABBREVIATIONS In lists of material, and figure captions Museum of Natural History. r and acknowledgements, institutions are NZGS — New Zealand Geological Sur- abbreviated as follows: vey, Lower Hutt. * BM(NH) — British Museum (Natural UCMP — Museum of Paleontology, Uni- History), London. versity of California, Berkeley. CAS — California Academy of Sciences, USGS —United States Geological Sur- San Francisco. vey (in Menlo Park, California • and LACM — Los Angeles County Museum Washington, D.C.) of Natural History. USNM — United States National LACMIP — Department of Invertebrate Museum of Natural History, Washin- Paleontology, Los Angeles County gton D.C.

TAXONOMY its varices are closely spaced and nearly Superfamily Tonnoidea aligned up the whorls (i.e., little more Family Bursidae Thiele, 1929 than 18(^a|jjart around the spiral), the Remarks : The origins of the family sculpture is of hoarse nodes, and the short Bursidae are obscure. No fossils of spire more nearly resembles that of Bur- Eocene (or earlier) age are recorded from sidae than that of Ranellidae. How- the European region or in Asia. The ever, the specimen clearly lacks all sign earliest taxon that has been referred to of the most distinctive character of Bur- the family is Bursa saundersi Adegoke sidae, the posterior siphonal canal at the (1977, p. 209, pi. 31, fig. 27, 28), a small top of the outer lip, and so has more the (22.0 mm x 14.4 mm), wide, generalised appearance of Ranellidae Ranellinae West African shell that is difficult to than of Bursidae. The significance of assign with certainty to either Ranel- the specimen is therefore uncertain ; it is lidae or Bursidae. Adegoke (1977) pre- earlier than all recorded Bursidae, and so sumably assigned it to Bursidae because suggests the possibility that Bursidae •i^jeu i iftfypD

Ranellidae and Bursidae 71

evolved from Ranellidae Ranellinae, but nana (Broderip and Sowerby, 1829) have is likely to be a ranelline convergent on a calcareous intritacalx and a terminal the bursid shell form, as the very opercular nucleus, the many Indo-West different radulae and fore-gut anatomies Pacific species of Bufonaria (most of the Ranellidae and Bursidae (Beu, revised by Beu, 1987) have a finely 'X 1981) make a close phylogenetic relation- bristled conchiolin periostracum and 1 ship between them unlikely. The speci- have the opercular nucleus mid-way men is from the late Paleocene Ewekoro down the columellar margin. Mar- Formation of Nigeria, West Africa. supina and Bufonaria are probably not The only Eocene records of Bursidae I closely related. am aware of are ^he western Amer- Species of Olequahia are far from typi- icas : Olequahia Ste^arE, 1926 from the cal Bursidae, having a long straight ante- late Middle Eocene to Oligocene of west- rior siphonal canal with the columella ern North America, and Marsupina continuing undeflected down its left yasila (Olsson, 1930) and M. chira (Ols- edge ; they more closely resemble cassine son, 1930) from the Late Eocene (Stone, cassids, particularly Galeodea, than other 1949) Talara Formation and Chira For- Bursidae. The possibility therefore mation, respectively, of Peru (Olsson, needs considering that the present con- 1930, p. 62). The absence of Bursidae cept of Bursidae is polyphyletic. Fig- from Tethyan warm-water faunas, in ures are provided here of the type species which Ranellidae (notably Sassia) are so of Olequahia to enable evaluation of its diverse, suggests that the Bursidae relationships. appeared only during Eocene time, and probably appeared first in the eastern Pacific during the Middle Eocene. Genus Olequahia Stewart, 1926 Marsupina yasila (Olsson) and M. Olequahia Stewart, 1926, p. 382. Type species (by chira (Olsson) have been well illustrated original designation) : Cassidaria washing- (Olsson, 1930, pi. 10, fig. 3-7, 12, 13) and toniana Weaver, 1912, Late Eocene, western clearly have the aligned, widely USA. extended varices, distinctive posterior Remarks: Important characters of siphonal canal, and evenly granulous the genus are its inflated, Galeodea-\ike sculpture typical of modern species of form, its moderately long, straight Marsupina and Bufonaria. The earlier columella and anterior siphonal canal, its species, M. yasila, is a uniformly prominent terminal varix, its narrow granulous species without prominent inner lip collar closely similar to that of sculpture, but with a moderately low many modern species of Bufonaria, and spire, with weak varices down the entire its relatively wide, moderately deep pos- teleoconch, and with short anterior and terior canal cut into the outer lip exten- posterior canals ; it could well have been sion, beyond the terminal varix. The a stem taxon for all Miocene to modern oldest species referrable to Olequahia Bursidae, with the possible exception of seems to be "Ranella" domenginica the western American endemic genus Vokes (1939, p. 147, pi. 19, fig. 6, 20), Crossata Jousseaume, 1881. Referral to which has prominent varices over the Marsupina Dall, 1904 seems reasonable. entire teleoconch, and is taller and nar- I have previously regarded Marsupina as rower than all younger species. The a subgenus of Bufonaria Schumacher, type species and the probably synony- 1817 (Beu, 1981). However, further mous O. hornii (Gabb), from the Tejon study has shown that, whereas Mar- Formation of California, have varices on supina bufo (Bruguiere, 1792) and M. the earliest 2-3 spire whorls, but none on A.G. Beu Ranellidae and Bursidae 73

the adult shell, other than the terminal could have evolved from a tall-spired, one; Oligocene species have only a ter- cymatiid-like ancestor. minal varix. 0. domenginica (Vokes, 1939) is recorded from the early Middle Eocene (Squires, 1983) Llajas Formation Olequahia washingtoniana in Simi Valley, California, and from the (Weaver, 1912). late Middle to early Late Eocene Domen- Fig. IF ; PI. 1, fig. 1-9.

gine Formation of California. Exami- ? Tritonium hornii Gabb, 1864, p. 94, pi. 28, fig. 208 nation of the two syntypes of 0. domen- (in part); Gabb, 1869, p. 218 (in part). ginica (UCMP-672/15803, 15804, Domen- ? Tritonium (Trachytriton) tejonensis Gabb, 1869, p. gine Formation, north of Coalinga, 154, pi. 26, fig. 34. California) shows, that the larger (40.3 x Cassidaria washingtonianaWe&veT, 1912, p. 38, pi. 3, fig. 28. 22.5 mm) is simBar^in shape to 0. Nyctilochus washingtoniana, Dickerson, 1915, p. 90, washingtoniana (Weaver) ; Givens pi. 7, fig. 9. (1974, p. 79, pi. 9, fig. 4, 5) also included ? Bursa hornii, Anderson and Hanna, 1925, p. 54, pi. 0. domenginica in Olequahia. Other 13, fig. 3, 4, 8. species referrable to Olequahia are: 0. ? Siphonalia tularensis Anderson and Hanna, 1925, pi. 10, fig. 1 only (not p. 45). washingtoniana (Weaver) from the Late Olequahia washingtoniana, Stewart, 1926, p. 383; Eocene Cowlitz Formation of Washin- Weaver, 1943, p. 425, pi. 84, fig. 6, 10, 11. gton and Oregon, and its probable syno- ? Olequahia hornii, Stewart, 1926, p. 382, pi. 29, fig. nym 0. hornii (Gabb, 1864) from the 1, 4, 18. Late Eocene Tejon Formation in Califor- Remarks : Stewart's (1926, pi. 29, fig. nia ; 0. lincolnensis (Weaver, 1916) from 1,4, 18) figures of Olequahia hornii the Oligocene Lincoln Formation in (Gabb) show no significant differences Washington ; 0. schencki Durham, 1944, from the specimens of 0. washingtoniana from the latest Eocene Keasey and I have examined ; the two nominal taxa Oligocene Eugene formations of Oregon ; have the same shape, spiral sculpture, and 0. lorenzana (Wagner and Schilling, and varices on the top few spire whorls . 1923), from the Middle Oligocene of only. I show below that both Ranella California (Hickman, 1969, p. 89,90 ; washingtoniana (Weaver) (= Gyrineum Hickman, 1980, p. 48). The earliest uvasilis Anderson and Hanna) and species is therefore the relatively narrow, Ameranella kewi (Dickerson) are present strongly varicate 0. domenginica in the Late Eocene Tejon Formation of (Vokes), which suggests that the genus California and the at least partly coeval

Fig. 1. Protoconchs of Ranellidae and Bursidae, at two different scales: Fig. A, C-F enlarged X22 ; Fig. B, G, enlarged x33. A. Ameranella kewi (Dickerson, 1915), type species of Ameranella n.gen., Cowlitz Formation, Late Eocene, Cowlitz River, Washington, USA; WM13325, NZGS. B. Sassia delafossei (Rouault, 1850), Gan, near Pau, southern France, Cuisian, late Early Eocene ; Mississippi Bureau of Geology. C. Cymatium (Monoplex) cowlitzense (Weaver, 1912), Cowlitz Formation, Late Eocene, Cowlitz River, Washington, USA ; LACMIP loc. 5654. D. Ranella washingtoniana Weaver, 1912, same locality and collection as Fig. C. E. Sassia formosa (Deshayes, 1865), Calcaire Grossier, Lutetian, early Middle Eocene, Chaus- sy, Oise, Paris Basin, France, ex Cloez Colin., UCMP-B5350. F. Olequahia washingtoniana (Weaver, 1912), type species of Olequahia Stewart, 1926; Cowlitz Formation; Late Eocene, Cowlitz River, Washington, USA; UCMP-D8044/ 16071. G. Eocymatium pyraster (Lamarck, 1803), type species of Eocymatium n.gen.; Calcaire Grossier, Lutetian, early Middle Eocene, Amblainville, Oise, Paris Basin, France, ex Cloez Colin.; UCMP-B5349. 74 A.G. Brn

Cowlitz Formation of Washington and ch (Fig. IF) is small, turbiniform, of Oregon; 0. hornii may well be another about 3.5 apparently smooth, inflated species in common. As I am not sure of whorls (the tip is abraded in most speci- the synonymy in this case and as 0. mens), and similar in most characters to washingtoniana is the type species of those of Cassidae (e.g., Abbott, 1968, pi. Olequahia, the illustrated specimens from 5, 6) and all other Bursidae. All Bur- the Cowlitz River type locality of 0. sidae have closely similar turbiniform washingtoniana are identified by that protoconchs, differing only at the species name at present. level. The specimens of 0. washingtoniana The posterior canal of Olequahia is examined show considerable variation in closest to that of Crossata californica spire height, whorl inflation, prominence (Hinds, 1843). Immature specimens of of spiral cords, width of the outer lip C. californica (particularly of the thin- extension beyond the terminal varix, shelled form or geographic subspecies and width and depth of the posterior Bursa californica sonorana Berry, 1970, siphonal canal. The outer lip is p. 118) are closely similar to 0. washing- crenulated into a series of rounded nodes toniana in the following characters : the and sinuses; as the lip margin grows position of the major spiral coyds forward after varix formation, it pro- (although only the upper two bear nodes duces a complexly frilled outer lip not in Crossata), the straight, continuous unlike that of some Muricidae. This columella and left side of the "anterior complexly sculptured lip extension is canal, the form of the outer lip, and the notched by the posterior canal, which is widely open, shallow posterior canal. widely, thickly and smoothly callused, The complex frills on the outer surface of with a wide, evenly rounded external the outer lip flare have been observed flare. The canal is constricted by a large also on some specimens of Crossata nodule on the top of the inner edge of the californica. Crossata differs clearly outer lip, and another on the parietal from all Olequahia species younger than region. The two nodules seem to 0. domenginica in having varices on all become progressively more callused as whorls. It seems feasible that, despite the outer lip extension grows forwards; the lack of a fossil record in Miocene of the specimens seen, the one with the rocks of western America, Olequahia was widest outer lip is also that with the either the direct ancestor of Crossata, or narrowest and deepest posterior canal (to judge from\the lack of varices in (PI. 1, fig. 5, 6, 9). Most specimens Oligocene Ole^UaMia species), it bran- examined, however, have a widely open ched from a lineage that led to modern posterior canal. Crossata species. The widely open ante- The anterior siphonal canal is unusual rior and posterior canals, the large aper- among Bursidae in being straight; the ture, and the subdued teleoconch sculpture certainly make Crossata a distinc- very straight columella continues with- v out deviation down the left edge of the tive, endemic western American genus canal. The complete length of the un- that appears to have had a long history broken canal is not demonstrated by any of isolation in the area. specimens I have examined. All speci- An important aspect of the morphol- mens examined have three prominently ogy of Olequahia is the outer lip exten- noded major spiral cords, low varices sion, a thin flange extending beyond the restricted to the first 2-3 teleoconch terminal varix. I am not aware of the whorls, and a clearly raised collar on the existence of such a lip flange in any left edge of the inner lip. The protocon- Cassidae, and its presence in Olequahia Ranellidae and Bursidae 75 suggests strongly (along with the poste- Oligocene and Neogene Ranellinae (par- rior apertural canal) that 0. washing- ticularly in genera Argobuccinum and toniana is convergent on the cassid shell Fusitriton) in the North Pacific, but form. Indeed, the morphology of Ole- Smith (1970) pointed out that the same quahia washingtoniana seems closest to treatment was needed for the Eocene >hat of Bufonaria in its elongate but taxa of western North America. A brief inflated form, the presence of only three review is undertaken here of most widely spaced, noded, primary spiral Paleocene and Eocene ranelline taxa of cords, and the straight columella. western USA. liufonaria differs in being more dorso- ventrally compresse^m^|aving evenly granulous interstitial sculpture, varices Genus Ranella Lamarck, 1816 on all whorls, and a longer, gutter-or Ranella Lamarck, 1816, pi. 412-414, "Liste" p. 4. tube-like posterior canal, but a close Type species (by subsequent designation, Chil- relationship appears feasible. It dren, 1823): Ranella gigantea Lamarck, 1816 remains enigmatic that two such marked- ( = Murex olearium Linne, 1758), (Oligocene?) Miocene to Pliocene, Europe; Recent, Mediter- ly different bursids as Olequahia and ranean, eastern and western Atlantic, South Marsupina should have appeared, appar- Africa, St Paul & Amsterdam Islands, Reunion, ently nearly simultaneously in North New Zealand, and Kermadec Islands. and South America respectively; the Gyrina Schumacher, 1817, p. 77, 253. Type species (by monotypy): Gyrina maculata Schumacher, first appearance of Bursa occurred else- 1817 ( = Murex olearium Linne, 1758) (not where, probably in the tropical Indo- Gyrina Linne, 1767, Insecta). West Pacific. Eugyrina Dall, 1904, p. 132. Type species (by original designation): Ranella gigantea Lamarck, Dimensions: Most complete speci- 1816 (= Murex olearium Linne, 1758). men seen (PI. 1, fig. 1-3): height 55.1 Mayena Iredale, 1917, p. 324. Type species (by mm, diameter 33.7 mm. original designation) : Biplex australasia Localities : All examined well- Perry, 1811, late Miocene-Recent, New Zealand preserved specimens are from the Cowlitz and southern Australia. | Gyrinopsis Dall, 1925, p. 18. Type species (by River near Vader, Lewis County, Wa- monotypy): Gyrinopsis cowlitzi Dall, 1925 shington, from the Cowlitz Formation (= Ranella washingtoniana Weaver, 1912), (Late Eocene). Several specimens from Mid-Late Eocene, western USA. this locality have been examined in I'SNM, UCMP, CAS, and LACMIP. The probably synonymous 0. hornii Ranella washingtoniana (Gabb) occurs in the coeval Tejon For- Weaver, 1912. mation near Tejon, Kern County, Fig. ID ; PI. 2, fig. 1-3. California. 0. hornii has also been Ranella washingtoniana Weaver, 1912, p. 41, pi. 2, recorded from the far eastern USSR by fig. 14. Schmidt" (Stewart, 1926, p. 382, Bursa washingtoniana, Dickerson, 1915, p. 64, pi. 4, synonymy list) so this species seems to be fig. 4, 6. widespread in Late Eocene rocks of east- Gyrineum uvasilis Anderson and Hanna, 1925, p. 57, pi. 6, fig. 1; pi. 10, fig. 5 ; pi. 13, fig. 13. ern Asia and western North America. Gyrinopsis cowlitzi Dall, 1925, p. 18, pi. 18, fig. 4, 6. Cymatium cf. washingtoniana, Turner, 1938, p. 90, pi. 16, fig. 17. Family Ranellidae Gray, 1854 Cymatium washingtonianum, Weaver, 1943, p. 412, Subfamily Ranellinae Gray, 1854 pi. 81, fig. 9, 11, 12 (in part). Remarks : The excellent monograph Remarks : Beu (1976, p. 424), Hick- by Smith (1970) has greatly clarified the man (1980, p. 48) and Squires (1983, p. taxonomy and phylogeny of the 358) have previously referred this species 76 A.G. Brn to Ranella sensu stricto; Weaver (1912) ships suggested here. presumably intended to place the species The identification of material of R. in what is now called Bursa. washingtoniana and other Ranellidae The shells of large, typical Ranella figured by Weaver (1943, pi. 81, 82) is. washingtoniana illustrated by Weaver muddled. Ranella washingtoniana actu- (1943, pi. 81, fig. 9, 11, 12) are closely ally appears on pi. 81, fig. 6, 9,11,12, and similar to both living species, R. olearia 13; fig. 13 shows the distinctive large (Linne, 1758) (PI. 4, fig. 4, 5) and R. protoconch of R. washingtoniana, australasia (Perry, 1811) (PI. 4, fig. 7), in although it is identified as Cymatium spire proportions, in sculpture, in having cowlitzense (Weaver, 1912). Weaver's the varices aligned — or nearly so — (1943) pi. 81, fig. 8, 10 and pi. 82, fig. 2, 3, down the entire teleoconch, and in aper- and 10 show a single species of tural characters. These include a rela- Cymatium (Monoplex) identified on pi. tively long, straight, narrow canal 81 as C. cowlitzense, and on pi. 82 as C. deflected slightly to the left, the presence etheriftgtoni Weaver, 1943; as the speci- of several low nodules of transverse men illustrated by Weaver (1943) in pi. ridges on the columellar base (apparently 81, fig. 10 is the holotype of C. cowlitzenr a consistent character or Ranellinae), se I conclude that this is the valid name and the presence of a shallow posterior for a species later also named C. ethering- apertural notch. The interior of the toni by Weaver (1943). Examination of p outer lip is almost smooth and curves many collections of Cowlitz River fossils gently out over the terminal varix in R. showed that only one species of washingtoniana. The type material of Cymatium (Monoplex) occurs there* A Gyrinopsis cowlitzi Dall, 1925 third taxon, identified below by the (examined, in USNM) consists of several name Ameranella kewi (Dickerson), was large, excellent specimens of R. washing- also identified as C. cowlitzense by toniana. The recently named R. kath- Weaver (1943, pi. 81, fig. 7 ; pi. 82, fig. 1, erinae Squires (1983, p. 358, fig. 2E-G) 4) who seems to have regarded this as the from the Llajas Formation (early Middle adult form of his C. cowlitzense ; the two Eocene) of the Simi Valley, California, is are readily distinguished by the varices still more like the living R. olearia, as it being nearly aligned in A. kewi but situ- has a flat-faced terminal varix with ated at each 0.66 whorls in C. cowlitzen- many small nodules along the inner edge se. of the outer lip. Several Cowlitz For- Dimensions : height 96.5 mm, diame- mation specimens of R. washingtoniana ter 55.8 mm (USNM 333358, lectotype of bear a well preserved protoconch (Fig. Gyrinopsis cowlitzi Dall, 1925) ; largest ID); it is closely similar to that of R. specimen seen. olearia and R.-australasia. The Localities : Examination of the types protoconch of R. washingtoniana is un- and of a series of topotypes of Gyrineum usually large for a member of Ranellinae uvasilis Anderson and Hanna, 1925 (4.5 mm high), rather narrowly tur- showed that this taxon was based on biniform, with 5 almost smooth, weakly specimens of R. washingtoniana from the inflated whorls, faint signs of spiral Tejon Formation at Grapevine Canyon threads and — unlike that of R..olearia and nearby localities in Kern County, — it develops three low, wide spiral California. Turner (1938, p. 91) record- cords over the last half-whorl. The ed what appears to be typical Ranella much smaller protoconchs of modern washingtoniana from the lower Coaledo Argobuccinum species and of Ameranella Formation (which he thought coeval kewi (Fig. 1A) help confirm the relation- with the Tejon and Cowlitz formations) Ranellidae and Bursidae 77

.it the east side of the lighthouse reefs, Ranella tuomeyi Aldrich, 1886. ('ape Arago, Oregon (UCMP loc. A715). Pi. 2, fig. 4-6. S.jtiires (1983, p. 358) identified a specimen from the lower Middle Eocene Ar- Ranella (Argobuccinum) tuomeyi Aldrich, 1886, p. 20, pi. 3, fig. 3. 'lath Shale in San Diego County, Califor- Triton tuomeyi, Harris, 1899, p. 64, pi. 8, fig. 12. nia. as R. washingtoniana. Numerous Ranella tuomeyi, Palmer and Brann, 1966, p. 873 "{H'cimens (below) have been examined (with further synonymy). from the Cowlitz River, Washington, Remarks: Figures are provided of from the Cowlitz Formation (Late the holotype of Ranella tuomeyi, which Kocene, approximately coeval with is among the earliest species I am aware Tejon Formation). The species is there- of in Ranella (sensu stricto). The fore widespread in. western North Amer- holotype is small (39.0 mm X 24.1 mm) ica, and occurs in rlc^of lower Middle and superficially resembles a large, very to Late Eocene age. finely sculptured form of the living Material examined : USGS loc. 4026, Western Pacific Gyrineum bituberculare I'SNM 333358, Dall's figured specimen (Lamarck, 1816), but its apex bears the (here designated lectotype) of Gyrinopsis last fragments of a formerly large, cowlitzi Dall, 1925, from Little Falls, smooth protoconch, much larger than Cowlitz River, Washington, with 10 that of Gyrineum species. The aligned, other specimens (all syntypes ?-not narrow varices, the long anterior canal, labelled as to status); USGS 18510, the form of the outer lip (with 8 interior "type locality of Cowlitz Formation", 2 nodules along an inner ridge, forming an (USGS, Menlo Park, California, and 2 almost flat outer face) and the simple with no locality data); two syntypes of spiral sculpture of narrow cords are all Ranella washingtoniana Weaver, 1912, characters in common with Ranella. R. Cowlitz River, Washington (CAS 488, tuomeyi and the following new species 188a).j holotype (CAS 829) and 2 differ from Ameranella n. gen. in their paratypes (CAS 830, CAS 872) of much larger protoconch and weaker Gyrineum uvasilis Anderson and Hanna, sculpture, and particularly in the lack of 1925, from Tejon, California; Cowlitz prominent nodules on the varices. Al- River below mouth of Drew Ck., 1.5 miles from Olequa, Washington, 9 (CAS drich (1886) and Harris (1899, p. 64) 61656.01); Tejon Formation, Liveoak mentioned several specimens, and Harris Canyon, Kern Co., California, 1 (CAS (1899, pi. 8, fig. 12) figured one that is 61661.01); UCMP7162/33863, Weaver's more complete than the holotype, bear- (1943, pi. 81, fig. 12) figured specimen, ing a large protoconch similar to that of Cowlitz River, Washington; Cowlitz R. olearia. River Eocene general collections, Locality: Woods Bluff, Tombigbee Museum of Paleontology, University of River, Clarks County, Alabama, USA; California, Berkeley (many); UCMP Bashi Member, Hatchetigbee Formation, loc. 7200, Tejon Formation, Tejon Quad- upper Wilcox Group, Early Eocene rangle, Kern Co., California, 5; UCMP (Palmer and Brann, 1966, p. 873) ; A452, Tejon, 3; UCMP A4522, Cowlitz USNM registered no. 638755. River, Washington, several, including Remarks: The only other Eocene specimen with complete protoconch; species I am aware of that seems refer- LACMIP loc. 5654, north bank Cowlitz able to Ranella is Triton eogassinense River 1.5 miles E of Vader, Lewis Sacco (1904, pi. 10, fig. 12), from Gassina, County, Washington, 5. Italy, Bartonian (Late Eocene), based on a poor internal mould resembling R. 78 A.G. Brn olearia (Linne). length unknown (broken off below junction of outer lip). Spire apex and protoconch missing. Ranella louellae n. sp. Dimensions: height (very incomplete) 34.1mm, (estimated) 52 mm, PI. 4, fig. 1-3. diameter (slightly incomplete) 27.0 mm. ?"Fusitriton" sp. aff. Murex (Argobuccinum) Repository : holotype (LACMIP mansfieldi Gardner ; Smith, 1975, p. 469. 7689) in LACMIP. Description: Shell very small for Locality : University of California at genus, very prominently sculptured, Los Angeles (UCLA) locality no. 6456, with prominent varices aligned down at basal Lodo Formation shellbed, abrupt least last few whorls of teleoconch (spire hill slope on south side of road, south missing). Sutural ramp wide, steeply side of Panoche Creek, 0.75 miles east of descending at first, then strongly con- junction with Silver Creek (3450'S, 500' cave. Sculpture of narrowly crested, E of NW corner section 21, T15S, R12E), widely spaced spiral cords, five on penul- Tumey Hills, Fresno County, California, timate whorl (lowest immediately above coll. L.R. Saul, 10 June 1977; holotype suture), nine on last whorl, and at least only ; Late Paleocene (see below). five further weak ones on anterior canal Remarks: Ranella louellae differs (which is very incomplete); interspaces clearly from all other Ranella species crowded with fine spiral threads of sev- other than R. tuomeyi Aldrich in its eral orders, at least four secondary small size, and from all other species,, in threads in each interspace; crossing its very prominent sculpture. R. prominent, closely spaced, rounded axial tuomeyi is closely similar in size and costae that rise on centre of sutural ramp, general appearance, differing from R. become very prominent around periph- louellae in its weaker sculpture (particu- ery (and are raised into low nodules larly the much less prominent axial cos- where crossed by spiral cords), and fade tae), its much lower varices, and its thin- out over lower half of last whorl; five ner but more complexly sculptured inner costae on each of the last three intervar- lip. The concave sutural ramp, the iceal intervals. Varices high and nar- abaperturally hollowed and buttressed row, hollowed abaperturally and buttres- varices, the nodules on the inr>er angula- sed by spiral cords, but spiral cords only tion of the outer lip, and the squarely slightly raised where they cross outer raised, thickened posterior apertural surfaces (i.e., varices lack nodules). margin and left edge of the inner lip are Aperture subcircular; outer lip flared characters that R^mellae shares with over inner two-thirds of terminal varix, the living type speciesf^Jlanella olearia weakly digitate around outer rim, bear- (Linne) (PI. 4, fig. 4, 5). Apart from the ing eight prominent, narrowly rounded much smaller size, the main differences nodules on angulation at inner edge of from R. olearia are the slightly more outer lip ; inner lip thick, with squarely prominent spiral cords, the markedly elevated rim along posterior margin of more prominent axial costae, and the aperture and alongside columella, fewer, more prominent nodules on the smooth except for a single prominent outer lip of R. louellae. Thus, although parietal nodule and five low, narrow, the lack of knowledge of the protoconch transverse ridges on base of columella. and spire proportions make a position in Anterior canal only narrowly open, Ranella tentative, I am reasonably deflected to left, with very low fasciole confident that is nevertheless the correct and narrow pseudumbilical groove, but position for R. louellae. Ranellidae and Bursidae 79

Illustrations of specimens from the Foraminifera of zone P4 (late Montian- basal Lodo Formation shellbed shown to early Thanetian). According to Zins- me by Dr J.Terry Smith (Palo Alto; meister (1983, p. 1282-3) and Saul (1983, during August 1987), identified by her as fig. 2) the Simi Conglomerate (type for- "Fusitriton" close to Murex (Argobuc- mation of Gyrineum judithae Zinsmeis- cinum) mansefieldi Gardner (Smith, ter, 1983) partly underlies and is partly a 1975, p. 469), are too fragmentary to be lateral facies equivalent of the Las Vir- identified with certainty, but have var- genes Sandstone, which in turn underlies ices situated at each 0.66 whorls and (in ascending order) the "Martinez appear to belong in Sassia. A single marine member" and the Santa Susana well preserved spire of a tall, narrow Formation. Saul (1983, p. 71) regarded Sassia species in LACMIP collection the unnamed molluscan stage (including 7044 (road cut oa east bank of Silver both the Las Virgenes Sandstone and the Creek, 0.25 miles^sQfffeh of Panoche Santa Susana Formation) below the Creek, Tumey Hills, Fresno County, Martinez stage as Danian in age, so it is California, coll. C.E. Weaver, 1949; not likely that the still lower Simi Conglom- now exposed) from the basal Lodo For- erate it also Danian (Early Paleocene). mation shellbed resembles juvenile speci- Age assignment is complicated by the mens of Charonia lampas (Linne) in fact that LouElla Saul (LACM; letter sculpture and proportions, and also 21 Sept. 1987), who has investigated the resembles Sassia mansfieldi (Gardner) location of Zinsmeister's fossils, reports (1935, p. 258, pi. 23, fig. 3-6) in sculpture. that his locality appears to be in a It is likely the specimen belongs in the tongue of Santa Susana Formation, but species referred to by Smith (1975, p. that it is nevertheless of late Danian or 469). perhaps early Montian age. Although Etymology : I have great pleasure in refinement of the ages of the earliest naming the new species in honour of Dr Cenozoic formations of California is LouElla R.Saul (LACMIP), for her clearly needed, there is little doubt that many valuable contributions to Califor- Gyrineum judithae is significantly older nian Cretaceous and early Cenozoic than Ranella louellae. paleontology. Relative ages of Lodo Formation and Simi Conglomerate: The authors of Genus Ameranella n. gen. faun'al revisions of the Lodo Formation (Smith, 1975) and the Simi Conglomer- Type species: Nyctilochus kewi Dickerson, 1915, late Eocene, western USA. ate (Zinsmeister, 1983) both considered the faunas they revised to be Thanetian Diagnosis: Shell small for the sub- (early Late Paleocene) in age. For family (about 40-60 mm high), with understanding the apparent first appear- small protoconch similar to that of mod- ances of ranelline genera it is desirable to ern Argobuccinum species. Varices refine the relative ages of these forma- aligned, or nearly so, down initial 3-4 tions as closely as possible. Saul (1983) teleoconch whorls, becoming separated has reviewed the micropaleontological by 5°-10° on later whorls of some species, evidence for age and the turritellid and bearing 3 moderately to very prominent, Venericardia biostratigraphy of several rounded nodules where crossed by the important Californian Paleogene major spiral cords. Intervariceal sculp- sequences, concluding that the basal ture of 3 or 4 rows of small to large, Lodo Formation shellbed (type forma- rounded nodules on the major spiral tion of Ranella louellae n. sp.) contains cords, and of interstitial spiral threads. T

80 A.G. Brn

Aperture similar to that of Ranella, but be considered an early species of the MY REC with outer lip interior gently rounded Argobuccinum group of lineages (which 0" PLEIS and weakly sculptured or with low nod- include Mediargo Terry, 1968) and in- Pic Z ules along inner edge, lacking the promi- cluded in Argobuccinum. G. judithae cer- nent angulation of Ranella species; a tainly is a small (30.5 mm high), moder- row of small nodules on base of columel- ately proportioned, evenly granulous 10- la ; parietal tubercle constricts a shallow species that seems to be a suitable mem- posterior sinus; anterior canal rather ber of a stem group for all the known short, moderately narrow, straight or later ranelline Ranellidae. weakly deflected dorsally and to the left. The next youngest named ranelline Remarks: The definitive generic taxa in western North America that 20- characters are: a rather small, Ranella- resemble Argobuccinum, rather than like teleoconch, an aperture close to that Ranella, are Argobuccinum (?) califor- 25- of Ranella, but lacking the angulation nicum (Gabb) and the species here along the inner edge of the outer lip that identified as Ameranella kewi (Dicker- 30- is present in most Ranellai species, sculp- son). The identity and taxonomic posi- ture dominated by prominent rounded tion of Argobuccinum (?) californicum 35- nodules (in particular, strongly protrud- (Gabb) (1866, p. 154 ; Gabb, 1869, pi. 26, La ing on the varices), and a small protocon- fig. 33) are obscure. Gabb's holotype w ch (much smaller than ,ihat of Ranella) (Stewart, 1926, pi. 30, fig. 6 ; Smith, * 40- resembling that of Argobuccinum. Com- 1970, pi. 49, fig. 11) is a small, immature, bined, these characters define a species slightly incomplete specimen with finely 45- group that does not fit into any ranellid and evenly granulous sculpture, from the genus I am aware of. This was also the Late Eocene Tejon Formation at Tejon, 50- opinion of Hickman (1980, p. 47). At California. The available material does not make it clear whether Argobuccinum present I know of only three species that 55- belong here: the western North Amer- (?) californicum (Gabb) is a distinct ican Eocene Nyctilochus kewi Dickerson, taxon occurring in the Tejon Formation 1915, "Fusitriton" terrysmithae Hickman, with A. kewi (the opinion of Dickerson, 60- 1980, from the latest Eocene Keasey 1915 and of Anderson and Hanna, 1925) Formation in Oregon, and the Chilean or whether only one variable taxon 55" Miocene Triton verruculosum G.B. Sower- occurs there. Argobuccinum (?) califor- by I, 1846. Although the two Eocene nicum is tentatively accepted as the 70- species are similar to and presumably name for the finely and evenly congeneric with the Miocene one, it granulous, Argobuccinum-like shell illus-

RECENT A W ^ > k > » ^ i ^ > WW PLEISTOCENE 11 ?l Riacenzian Zanclean Messinian

Tortonian

Serravalian

Langhian

Burdigalian

Aquitanian

Chattian

"o Rupelian

Tongrian Lattorfian Pnabonian

Bartonian ,« / 1/ Wemmelian

Lutetian Cv*san i Ypresian w / J Ml Landenian \ Thanetian / Se«landian / Montian SUBFAMILY CYMATIINAE SUBFAMILY _ J Danian RANELLINAE

Maastrichtian 1 111 •a LU 1 < • to LU Q U- Z O ; o CO CC | o • * < < 1- a CO | Santonian 1 1 RANELLIDAE Coniacian I I Turonian

Canomanian ? ancestral tonnoidean ?

R»>corded time ranges of the family Bursidae and of the genera of Ranellidae and Per- «'fii(lae. with a tentative phylogeny. Time ranges are shown against the European "standard" .vquence and an approximate time scale in years (at left), correlated largely from Stevens (I'.HI). Heavy vertical bars show recorded time ranges; narrow dashed lines suggest ancestries hm not necessarily times of origin. Only part of Late Cretaceous time is included. 82 A.G. Brn meister was a member of an Early spiral cords (particularly the interstitial Paleocene species group ancestral to all ones), and in the prominence of the younger Ranellinae. Ranella had sparse hemispherical nodules on the var- evolved by Late Paleocene time and ices and on the major spiral cords. All occurred from western North America to specimens have the nodules arranged in the Tethys by Late Eocene time ; it has axial rows, continuing further down onto remained little-changed since that time. the base than on A. verruculosa. All Argobuccinum seems likely to have specimens bear the row of small nodules evolved directly from small, generalised, on the base of the columella that char- early species resembling Argobuccinum acterises the subfamily Ranellinae. The (?) californicum (Gabb). Priene similar- inner surface of the outer lip" is curved ly is little different from Oligocene North gently out over the terminal varix, and Pacific Argobuccinum species, has only a bears nodules that range from small, Pleistocene fossil record in South Amer- rounded denticles to short, narrow, trans- ica, and perhaps represents a stock of verse ridges. All specimens have a low Argobuccinum that crossed the tropics to spiral cord on the sutural ramp raised the eastern South Pacific early in Cen- into a fourth low variceal nodule that is ozoic time. Fusitriton also seems likely absent from A. verruculosa. The best- to have evolved frbm an early evenly preserved Tejon Formation topotype granulous species group similar to examined (UCMP 7200 ; PI. 2, fig. 8) has Gyrineum judithae and Argobuccinum (?) relatively coarse interstitial spiral cords, californicum, by lengthening of the spire but these can be matched in the Cowlitz and anterior canal. Another minor line sample; it also has relatively low nod- here named Ameranella n. gen., had ules on the cords and varices, but these branched off the finely granulous stem are little different from those on the more group similar to Gyrineum judithae by finely sculptured shells examined from Late Eocene time and developed large, the Cowlitz Formation, including the rounded variceal nodules, and ultimately finely sculptured specimens illustrated gave rise to the Chilean Miocene Triton by Weaver (1943, pi. 81, fig. 7 ; pi. 82, verruculosum Sowerby. fig. 1, 4). There seems little doubt that populations from the Tejon and Cowlitz Formations are conspecific. The coarse- Ameranella kewi (Dickerson, 1915) ly sculptured specimens illustrated here Fig. 1A ; PI. 2, fig. 7-13. (PI. 2, fig. 9, 10) are uncommon in the Cowlitz Formation population, and are Nyctilochus kewi Dickerson, 1915, p. 64, pi. 7, fig. 5a, illustrated to demonstrate the similarity 5b. Gyrineum kewi, Anderson and Hanna, 1925, p. 56, of this species to Ameranella verruculosa pi. 10, fig. 4 ; pi. 13, fig. 12. (Sowerby). ^ k* i Cymatium washingtonianum, Weaver, 1943, p. 412, pi. 81, fig. 7 (only), pi. 82, fig. 1, 4 (only) (in Dimensions : height 37.8 mm, diame- part). ter 23.3 mm (WM13325, NZGS; PI. 2, "Mayena" kewi, Smith, 1970, p. 523, pi. 49, fig. 3, 4. fig. 11-13); height 41.6 mm, diameter 25. ? NOT Gyrineum kewi, Clark, 1938, p. 717, pi. 2, fig. 0 mm (not including attached polychaete 32 (Argobuccinum ? californicum (Gabb)). tube) (LACMIP 5654 ; PI. 2, fig. 9); Remarks : Examination of a series of height 39.3 mm (apex incomplete), dia- topotypes from the Tejon Formation of meter 26.7 mm (UCMP-A3787, largest California and of a large series of well seen ; PI. 2, fig. 10). preserved Cowlitz River specimens shows Material examined: UCMP 7200, that this is a highly variable species. Tejon, Kern County, California, Tejon Specimens vary in the prominence of Formation, Late Eocene, 1; UCMP- Ranellidae and Bursidae \ 89

A971, Tejon (as above), 2 ; UCMP-A452, coarsely sculptured specimens of A. kewi, Tejon (as above), 3; USGS loc. 4026, although it does not appear to have the Little Falls, Cowlitz River, Washington, same prominent variceal tubercles as are Cowlitz Formation, 1 (USGS, Menlo present on A. kewi and A. verruculosa Park, California); USGS loc. 18511, (Sowerby); more material is needed to Cowlitz Formation, Cowlitz River, 1 clarify this character. The holotype of (USGS, Menlo Park, California) ; A. terrysmithae differs from all specimens "plesiotypes" of Anderson and Hanna of A. kewi I have seen in its more numer- (1925), Tejon, California (CAS 827, CAS ous rows of nodules on whorl surfaces; 828); Tejon Formation, Canada de las the mid-sutural ramp cord in raised into Uvas, 1 mile E of Grapevine pumping a row of nodules as large as those below station, Kern Co., California 1 (CAS (it is weaker in all A. kewi) and there are »»1065.01); Tejon Formation, east side five lower rows of nodules in A. terrys- (irapevine Canyon, Kern Co., California, mithae, rather than the three rows and 1 (CAS 61663.01) ; holotype of weak fourth row in A. kewi. The par- (li/rineum kewi Dickerson, 1915, Tejon, atype of "Fusitriton" terrysmithae (Hick- California, UCMP 458/11052 ; Weaver's man, 1980, PI. 6, fig. 4) has widely separ- (1913) illustrated specimens ident. as ated varices and appears to be a speci- "Cymatium cowlitzense (Weaver)", Cow- men of a Sassia species close to S. litz River, Washington, UCMP-7163/ bilineata (Dickerson, 1916, p. 493, pi. 37, :i:lS68; UCMP-B5838, type section Cow- fig. 6a, b). litz Formation, Cowlitz River, several; Dimensions : height 36.5 mm, diame- rS0S loc. 4026, Cowlitz Formation, ter 20.0 mm (holotype ; Hickman, 1980, Cowlitz River, Washington, with p. 47). *vntypes of Gyrinopsis cowlitzi Dall, 1 Locality: USGS loc. 15268, upper (t\SNM 333358); LACMIP loc. 5654, part of Middle Member of Keasey Forma- north bank of Cowlitz River 1.5 miles tion (latest Eocene; Hickman, 1980, p. «\tst of Vader, Lewis County, Washin- 6), in the upper Nehalem River Basin, gton, Cowlitz Formation, 3. northwestern Oregon; holotype only (USNM 251369). Ameranella terrysmithae (Hickman, 1980) Ameranella verruculosa yusitriton" terrysmithae Hickman, 1980, p. 47, pi. (». fig. 4-6. (Sowerby, 1846) PI. 3, fig. 1-4, 6, 7 Remarks : The description and clear figures by Hickman (1980) make it clear Triton verruculosus G.RjSowerby I, 1846, p. 260, pi. that "Fusitriton" terrysmithae is a second 4, fig. 23. ? Tritonium exiguum Philippi, 1887, p. 57, pi. 3, fig. western North American Late Eocene 23 (juvenile ?). • juries of Ameranella. The holotype of "Tritonium" (Ranella 1) verruculosum, Tavera ' Fusitriton" terrysmithae has varices Jerez, 1979, p. 90, pi. 19, fig. 67a, b. .tinned to form two continuous ridges NOT Tritonium verruculosum, Philippi, 1887, p, 57, pi. 4, fig. 10 [ = Sassia armata (Hupe, 1854)}..** «lo\vn early teleoconch whorls, although iliw become a little separated over the Remarks : The illustration by Sower- U>t whorl in the same manner as most by (1846, pi. 4, fig. 23) and Sowerby's • jwrimens of A. kewi (e.g., PI. 2, fig. 12). type (BM(NH) Department of The holotype is similar in size, propor- Palaeontology, no. G26396) show that tions, apertural characters, anterior si- this is an unusual Bursa-\ike species j'honal canal, and sculpture to the more with aligned lateral varices, each varix 84 A.G. Bm bearing 3 large, hemispherical nodules; Miocene, -collected by Charles Darwin the intervariceal surface bears a few (but note that Darwin's locality large nodules on the major spiral cords, "Navidad" is neither at the present site similar to those on the varices. It lacks of Navidad nor in Topocalma Bay, a bursid posterior siphonal canal, and where the locality is said to be by mod- clearly belongs in Ranellidae Ranellinae. ern Chilean geologists (e.g., . Cecioni, Examination of W.J. Zinsmeister's col- 1978, p. 3); Darwin's locality is that lections of Chilean Miocene fossils now known as Matanzas, about 15 km showed that this is a reasonably common south of the mouth of Rio Rapel; see species in the Middle Miocene Navidad Cecioni, 1978, fig. 1; pers. comm. W.J. Formation in the Navidad-Matanzas Zinsmeister, letter 1987). Department area, near the mouth of Rio Rapel, Chile. of Earth and Atmospheric Sciences, Pur- Few complete, well preserved shells have due University, Indiana, collections from been seen, but many are complete the Chilean Middle Miocene Navidad enough to demonstrate that Triton ver- Formation (Group ?) (Punta Perro-La ruculosum resembles Ameranella kewi Era Formation of Cecioni, 1978) by W.J. closely in critical characters, and is refer- Zinsmeister : loc. 145, float material from able to Ameranella. Rio Rapel-Navidad area, 1; loc. 264, Ameranella verruculosa differs from A. sandstone 10 m above beach, Punta kewi and A. terrysmithae in its slightly Perro, southern point of mouth of Rio larger size (complete large shells prob- Rapel, (33°55'10"S, 71°50'20"W), 1; loc. ably reached about 60 mm high), its 286, base of seacliff 1 km north qf mouth considerably narrower form and taller of Rio Rapel (33°52'50"S, 7r49'40"W), 1 spire, its weaker interstitial spiral cords, (PI. 3, fig. 1, 2); loc. 326, base of sea cliff its lack of a raised cord on the sutural 200 m south of loc. 286, North Rio Rapel ramp and its much more prominent nod- (33°52'59"S, 71°49'40"W), 4; loc. 327, ules, particularly on the varices. base of sea cliff 100 m south of loc. 326, Philippi (1887, pi. 3, fig. 23) figured as North Rio Rapel (33°53'00"S, 71°49'49" Tritonium exiguum a very small, un- W), 2 ; loc. 331, 3 m above beach, seacliff diagnostic specimen of a taxon with 150 m north of loc. 286, North Rio Rapel aligned varices and large nodules. The (33°52'40", 71°49'40"W), 2 ; loc. 332, sea- only Chilean ranelline taxon this shell cliff 3 m higher than loc. 331, 9 (PI. 3, fig.. 4, 5, 6, 7); loc. 335, north side of mouth can be referred to is Ameranella ver- ,/ ruculosa. As Philippi (1887, pi. 4, fig. of Rio Rapel (33°52'55"S, 71°50'26 N), 1. 10) misidentified the common Chilean Remarks: It may be added that a Sassia armata (Hupe) (1854, p. 22) by species of Ranella (sensu stricto), closely the name Tritonium verruculosum, it is resembling both short-spired specimens feasible that he renamed a juvenile speci- of R. washmgtoniana (Weaver) and the men of A. verruculosa. New Ze|tah4 early Miocene R. kaipar- Dimensions : Height (incomplete) 46. aensis (Finlay), occurs in the Chilean 9 mm, diameter (incomplete) 26.3 mm Miocene Navidad Formation ; W.J. Zins- (estimated originally ca. 32 mm) (most meister's locality 254, fossiliferous lens at complete large specimen seen, WJZ loc. low tide ca. 100 m north of Matanzas 286); height 34.3 mm, diameter 25.2 mm township (33°57'30"S, 71°52'20"W)^3 km (Sowerby's very incomplete holotype, south of Navidad, Chile (Darwin's drigi- BM(NH), G26396). nal "Navidad" locality ; pers. comm. W. Material examined : holotype, J. Zinsmeister), 2 specimens; neither BM(NH) Palaeontology Department, no. Sassia armata (Hupe) nor Ameranella G26396, from Navidad, Chile, Middle verruculosa are present in this collection. Ranellidae and Bursidae 85

Subfamily Cymatiinae Iredale, (Dall, 1904, p. 130 ; later usages listed by 1913 (1891) MacNeil and Dockery, 1984, p. 120) in Genus Sassia Bellardi, 1872 regarding Personella septemdentata (Gabb, 1860) (type species of Personella Sassia Bellardi, 1872, p. 219. Type species (by subsequent designation, Cossmann 1903, p. 93): Conrad, 1865, p. 21) as a member of the Triton apenninicum Sassi, 1823, Miocene and group of small Personidae with only Pliocene, Europe. weakly distorted coiling and with the basal columellar nodules situated direct Synonymy : Beu (1987, p. 307). ly on the columella, rather than elevated Remarks : Sassia is very much the on a prominent callous ridge as in Distor- most diverse, the most widespread, and sio and in Kotakaia n.gen. MacNeil and the longest-ranging^nus of Ranellidae ; Dockery (1984, p. 118) and I previously it is the only undoubtedfranellid genus I thought there were no taxa resembling am aware of that occurs in Cretaceous Byramia in the American Middle rocks (Distorsio, formerly placed in the Eocene, but comparison of figures and Ranellidae, is removed below to the sepa- specimens recently brought to light the rate Family Personidae). More than real relationship of Personella septem- half of the named species of Ranellidae dentata : it is the earliest recorded belong in Sassia. species of the subgenus previously I have previously (Beu, 1987, p. 308) known as S. (Byramia). Relatively suggested that three groups of Sassia small (immature ?), complete specimens species were sufficiently distinctive to be of P. septemdentata (PL 3, fig. 9) share recognized as subgenera, but I have with Byramia species a small size, regu- recently recognized that there is confu- lar coiling, similar low, cancellate sculp- sion about one of these subgenera, and it ture with relatively widely spaced, flat- is desirable to rectify current usage, as topped spiral cords, a moderately long, one of 'the names has been incorrectly straight, narrow, open anterior canal, 4 or applied to a genus (or subgenus) of 5 low basal columellar ridges, a slightly Personidae (discussed below). Two sub- enlarged second nodule from the top genera are distinctive enough to be easily inside the outer lip, and a prominent recognized, and are not discussed fur- angulation at the top of the outer lip and ther: Sassia (Austrotriton) Cossmann, terminal varix, formed at the junction of 1903 (p. 98), containing southern Aus- near-horizontal and near-vertical seg- tralian fossil and living species with ments of the lip. They differ from the caricelloid apices or no protoconch at all, genus of small Personidae previously reflecting direct development; and Sas- confused with Personella (renamed Per- sia (Cymatiella) Iredale, 1924 (p. 253), sonopsis n.gen. below) by the following containing Paris Basin Eocene, North characters: the second, rather than American Oligocene, Caribbean Miocene, third, outer-lip nodule is enlarged, coi- and southern Australian Miocene to ling is almost completely regular, spiral Recent species of small size (few over 20 cords are more distinct, the anterior mm high), with tall spires, short anterior canal is long and narrow, rather than an canals, and proportionally small anterior spout at the end of a gradually protoconchs. tapered last whorl as in Personopsis, and the characteristically personid, widely It is the third subgenus, identified spread inner lip of Personopsis is absent. previously as Sassia (Byramia) MacNeil I regard Sassia (Personella) as an earlier in MacNeil and Dockery (1984, p. 118), name for S. (Byramia), and S. caseyi that needs reconsideration. I have MacNeil in MacNeil and Dockery (1984, previously followed earlier workers 86 A.G. Brn p. 120) as a second North American was the stem group of Ranellidae, as Oligocene species of S. (Cymatiella) (see noted above. Two typical European also S.fuscicava MacNeil in MacNeil and Eocene species are illustrated as exam- Dockery, 1984, p. 118). The following is ples of Sassia teleoconch (PI. 3, fig. 5, 8) a single, very uniform American lineage and protoconch (Fig. IB, E) morphol- of Sassia (Personella) species, apart from ogy : the very finely sculptured, elongate an apparent speciation event that S. formosa (Deshayes) (to which the New produced two Oligocene members: S. Zealand and southern Australian living septemdentata (Gabb), Middle Eocene; species S. parkinsonia (Perry, 1811) is S. jacksonensis (Meyer) (see Harris and closely convergent) and the coarsely can- Palmer, 1947, p. 336, p. 44, fig. 7-9), Late cellate aS. delafossei (Rouault). Eocene; and S. abbreviata (Conrad, Several other genera that occur in 1848) and S. mississippiensis (Conrad, Cretaceous rocks have traditionally been 1848), both Oligocene. Whether any placed in the Ranellidae, but in my European fossil species are referable to opinion belong in several other families. Sassia (Personella) requires reevaluation. Beu and Maxwell (1987) have removed The following discussion refers entirely the Plesiotriton group of genera, along to species of Sassia (Sassia). with Semitriton and Tatara, to the fam- Sassia (Sassia) species are mostly of ily Cancellariidae; it is now clear that small size (20-50 mm high), although no Ranellidae have columellar plaits. several lineages have given rise to much Trachytriton Meek, 1864 (type species: larger species. They liave a small tur- Buccinum ? vinculum Hall and Meek, biniform protoconch, although it varies 1856; Pierre Shale, Late Cretaceous, from vary small (about 2 mm in diame- USA) was discussed by Sohl (1967, p. 28, ter) and smooth, in the majority of pi. 6, fig. 10,14-17) who considered that species (e.g., the European Eocene S. T. vinculus is the sole species of the formosa (Deshayes, 1865); Fig. IE) to a genus; his illustrations show clearly little larger and coarsely cancellate, in that this is a thin-shelled, fusiform, species closely related to S. apenninica finely cancellate taxon without any var- (Sassi) (Beu, 1978, fig. 6,9), to 4 mm ices, but with internal dentate thicken- wide, with fine, close, conspicuously can- ings at growth pauses. It is quite unlike cellate sculpture (in the European all Ranellidae, and appears to belong in Eocene S. delafossei (Rouault, 1850); the Buccinidae. "Charonia ?" univar- Fig. IB). S. delafossei is the earliest icosum (Wade, 1926) may be a buccinid species I am aware of with cancellate neogastropod related to Ranellina protoconch sculpture, and seems likely to Conrad, 1865, as the holotype have given rise to the lineage of S. (USNM32916, ôm the Ripley Forma- apenninica. The Sassia teleoconch is a tion, Maastrichtianât Coon Creek, Ten- very generalised ranellid one, of moder- nessee) has a subsutural fold and concave ately elongate shape, with an equally sutural ramp, a shallow shoulder sinus, long spire and anterior canal, a relatively and only a terminal varix; but it has small muriciform aperture with small, much more prominent axial sculpture, even denticles inside the outer lip, and than Ranellina and possibly belongs in vV evenly convex whorls with sculpture of Sassia. The holotype of Tintorium similar spiral cords and axial costae. To pagodaeforme Sohl, 1960 (type species of judge from morphology, it is the most Tintorium Sohl, 1960), also from the likely stem group from which all other Maastrichtian Ripley Formation at Coon Ranellidae were derived. Stratigraphic Creek, Tennessee (USNM 128570), is a evidence also strongly suggests that it small shell (9.6 mm high) with a tall Ranellidae and Bursidae \ 89 spire, short last whorl, no varices, and a stromboidean fa. Vul- coronate shoulder; it clearly belongs !> SI , <*> species of Colwt <*> near Cerithioderma in the mesogastropod (Cossman, 1904, p. family Trichotropidae. With the ly resemble Ranelli ^moval, below, of Distorsio to the dis- the ancestry of th tinct family Personidae, Sassia remains noidea. as the only Cretaceous ranellid. All earliest Paleoc The earliest undoubted species of Sas- am aware of also belc cas- sia I am aware of is Tritonium kanabense v sia). The Paleocene- , Jcxie radiation Stanton (1893, p. 15j, dI. 31, fig. 12) of the subfamily Ranellinae has been (holotype examined, USN5122919), from discussed above; by Late Paleocene- the upper Kanab Valley, Utah. The Early Eocene time other genera of holotype was collected from the Inocer- Cymatiinae were beginning to radiate amus fragilis zone of the Colorado from Sassia. The earliest species refer- Group, of Turonian age (Cobban and rable to Cymatium is C. (Monoplex) Reeside, 1952, table 10b). Most other janetae Squires (1983, p. 355, fig. 2A-D) Cretaceous Sassia species are of Maas- from the lower Middle Eocene Llajas trichtian age : Triton konincki Binckhor- Formation in the Simi Valley, Califor- st (1861, p. 4, pi. 1, fig. lOa-c), Europe ; nia. (Although Cymatium janetae Tritonium tuberculosum Kaunhowen Squires and C. amnicretum MacNeil (in (1898, p. 77, pi. 9, fig. 3, 3a-d), Europe ; MacNeil and Dockery, 1984, p. 116, pi. Triton sauryi Basse (1933, p. 74, pi. 10, 18, fig. 4, 5) were placed in Cymatium fig. 1, la), Madagascar; Tritonium (Sas- (Septa) at my suggestion, I have since sia) sp. of Douville (1904, pi. 40, fig. 25), (Beu, 1987, p. 274) advocated subdivi- Iran; and' possibly Tritonium univar- sion of the broad subgenus C. (Septa) icosum Wade (1926, p. 147, pi. 51, fig. 9, into several subgenera aîd now refer 10; see also Sohl, 1960, pi. 18, fig. 44), these fossil species to C. (Monoplex)). USA; this last species more closely The next earliest species is C. (Mono- resembles the group of Buccinidae plex) cowlitzense (Weaver, 1912) related to Ranellina (see above). (Weaver, 1943, pi. 81, fig. 8,10 ; pi. 82, Abdel-Gawad (1986, p. 115-166) recently fig. 2, 3, 10) [ = C. etheringtoni Weaver, recorded S. tuberculosa (Kaunhowen) 1943; distinguished above from Amer- from early Late Campanian and Maas- anella kewi (Dickerson)] from the Late trichtian localities in the Vistula Valley, Eocene Cowlitz Formation at the Cow- Poland. The specimen illustrated as litz River, Washington. These seem rea- "Charonia" multicostata (Favre, 1869) sonably typical of C. (Monoplex) in all by Abdel-Gawad (1986, pi. 14, fig. 11, 12) characters; the protoconch (Fig. 1C) is is in my opinion not a tonnoidean and well preserved on many Cowlitz speci- the taxonomic position of the poorly mens, and confirms that it was already preserved material is enigmatic. "Bi- taller than those of any known Sassia flex" cretaceus Abdel-Gawad (1986, p. species. This species group, then, is a 115, pi. 15, fig. 1) is similarly enigmatic, suitable stem group for the large variety possibly a muricacean neogastropod, but of later Cenozoic to Recent Cymatiinae. certainly not tonnoidean. Several other This long preamble on the apparent first very poorly known Maastrichtian species appearances of early Cymatiinae is neces- named in "Triton " could possibly belong sary to make clear my opinion on the in Sassia, but cannot be evaluated from phylogeny and taxonomic positions of published figures. Most other Mesozoic the named taxa, and to reveal the dis- taxa assigned to "Triton " belong in the tinctive characters of the following new 88 A.G. Brn genus. phology to Oligocene to Recent Cymatium (Ranularia) species results from convergence. The protoconch Genus Eocymatium n. gen. characters and the teleoconch form jus- Type species: Triton pyraster Lamarck, 1803, tify^ the proposing of a new genus Lutetian (lower Middle Eocene), France and Eocymatium for Triton pyraster Lamar- Italy. ck, 1803; the only other taxon that Diagnosis: Teleoconch shape and seems to belong here is "Ranularia ?" of sculpture much as in Cymatium Cossmann and Pissarro (1909, p. 37, pi. 3, (Ranularia) Schumacher, 1817, with fig. 35, 36), a more finely sculptured but only a terminal varix or in addition one as yet unnamed species from the Late earlier varix to left of aperture on last Paleocene uppermost Ranikot beds at whorl; spire short; anterior canal long, Jhirak, Pakistan (Vredenberg in Coss- narrow, deflected slightly dorsally and to mann and Pissarro, 1909, p. xi, xii). the left. Whorls evenly inflated. The early Middle Eocene age and the Sculpture dominantly of prominent, nar- unique protoconch morphology of row, flat-topped, spiral cords crossed by Eocymatium pyraster indicate that narrow-crested, widely spaced axial Eocymatium represents an early, brief folds. Aperture subcircular, lips thick- lineage with no apparent descendants; ened ; inner edge bf outer lip prominent- it is unlikely to have descended directly ly denticulate, uppermost nodule largest, from Sassia. constricting a shallow posterior sinus. Protoconch very small, low-turbiniform, slightly inclined to the axis of the Eocymatium pyraster teleoconch on most specimens, of 1.8 (Lamarck, 1803). inflated whorls, with a relatively large, Fig. 1G ; PI. 3, fig. 10,12,13 in-rolled apex; the first half-whorl is Murex pyraster Lamarck, 1803, p. 225; Lamarck, microscopically reticulate, merging im- 1822, p. 575 ; Palmer, 1977, p. 15, pi. 4, fig. 9a, perceptibly into a succeeding whorl with b. about 20 very fine, well raised spiral Triton pyraster, Deshayes, 1835, p. 616; Deshayes, threads, in turn gradually developing 1837, pi. 80, fig. 36-38. fine, very weak axial costellae (initially Triton piraster [sic], Deshayes, 1865, p. 303; Coss- mann, 1899, p. 121. only below the upper suture) that Triton (Gutturnium) pyraster var. bilineatum become more prominent over the last Gregorio, 1880, p. 102, pi. 4, fig. 18. quarter-whorl; demarcated only weakly Lampusia (Gutturnium) piraster, Cossmann and from the teleoconch, by increase in spac- Pissarro, 1900, PUî.P1- 14, fig. 12. ing of axial riblets and the development Tritonium (Ranulariaj'pirmter, Cossmann, 1903, p. 97, pi. 3, fig. 21. ' ^ of more prominent, more widely spaced Eutritonium (Ranularia) piraster, Cossmann and spiral cords on the early teleoconch. Pissarro, 1913, pi. 35, fig. 167-22. Remarks : I know of no other ranel- Eutritonium piraster, Furon and Soyer, 1947, p. 119. lid with a short spirally lirate protoconch Remarks : The important taxonomic resembling that of Eocymatium pyraster criteria are stated in the diagnosis above. (Lamarck); its form and lack of obvious The Paris Basin Lutetian specimens I demarcation from the teleoconch suggest have examined are variable in size, that this species had lecithotropic devel- width, spire height, length of anterior opment, a rare mode in this family of canal, and prominence of the axial folds ; "ultradispersalists". The unique the two illustrated specimens indicate protoconch demonstrates conclusively the range of variation. Gregorio's (1880, that the resemblance in teleoconch mor- p. 102, pi. 4, fig. 18) description and illus- Ranellidae and Bursidae 89

t ration of his "bellissima varieta del col southwest of Lokia), close to local- pyraster" show no significant differences ities where specimens of an Inoceramus from Paris Basin shells other than hav- species were collected that were later ing only a terminal varix, an individual identified by Sornay (1968, p. 10) as I. character. pseudoregularis Sornay, of lower to lower Dimensions : height 26.7 mm, diame- middle Campanian (Late Cretaceous) '•'^x ter 16.7 mm ; height 24.0 mm, diameter age. 12.6 mm (the 2 illustrated specimens). As Distorsio praegranosa (Cottreau) Localities : Lamarck (1803) and most had already evolved typical Distorsio subsequent authors seem to have record- morphology by lower Campanian time, ed the species only from the Calcaire older than all known Undoubted Ranel- Grossier (Lutetian^paijg Middle Eocene) lidae of the genus Sassia other than the at Grignon and nearby localities in the Turonian S. kanabensis (Stanton) (see Paris Basin, where it seems to be reason- above), it is clear that Distorsio and ably common. Cossmann and Pissarro related genera (Distorsionella Beu, 1978 ; (1900, p. 131) recorded it from Fresville, Personopsis n.gen.; Kotakaia n.gen.) rep- near Yalognes, Normandy, presumably resent a distinct monophyletic group also Lutetian. Gregorio (1880, p. 102-3) that evolved independently of Ranel- did not state a specific locality for his lidae ; inclusion of this group (even as Italian specimen, but stated in the pref- subfamily Personinae) would make ace (p. XV) that the four fossil localities Ranellidae a polyphyletic taxon. at S. Giovanni Ilarione are Ciupio, Poz- The concept of Personidae as a distinct zani, Bosco del Prete, and Croce Grande, family of Tonnoidea is strengthened by and most specimens came from the first the distinctive fore-gut anatomy of Dis- or last of these. The age is given only as torsio (Lewis, 1972), resembling that of "Parisian". Ficus (Beu, 1981, p. 251) more than the f Ranellidae. Beu (1981, p. 251) also pointed out that the small, black, subcir- Family Personidae Gray, 1854 cular operculum of the ficid Thalassocyon Phylogeny : The earliest species I am Barnard, 1960 more closely resembles aware of that is referrable to Distorsio that of Distorsio than of any other ton- Roding, 1798 or to closely related genera noidean. 1 is Eutritonium (Sassia) praegranosum The anatomical evidence therefore sug- Cottreau (1922, p. 66, pi. 9, fig. 4-7). gests that Personidae could be more This is a moderately large (to 34 mm nearly related to Ficidae than to Ranel- high; but the two illustrated internal lidae, but the very different shell form of moulds lack anterior canals and spire Ficidae (elongate and fig-shaped, with apices and probably represent shells 45- weak sculpture and no varices) from that 50 mm high), highly distorted, coarsely of Personidae suggests that the relation- cancellate species, with coiling distortion ship is more distant than at the family and sculpture similar to those of Neogene level. Another distinctive character of and Recent Distorsio species. It particu- Distorsio and Distorsionella is the unique larly resembles the Caribbean Distorsio radula with a crescentic rachidian tooth clathrata (Lamarck, 1816), and I am in (Clench and Turner, 1957, pi. 132 ; Beu, no doubt that it belongs in Distorsio. 1978, fig. 30) that is not closely similar to The specimens are from two localities in the almost uniform radulae of Ranel- Madagascar (Province of Mananjaray: lidae, all easily derived from the range of hills west- south-west of Antani- equidimensional rachidian tooth of Sas- hody; Province of Vatomandry: third sia. (The radula of Personopsis (?) 90 A.G. Brn pusilla (Pease) is unknown, and must be low parietal ridges, and four or five low examined to confirm its placement.) transverse ridges (the uppermost the lar- The combination of a first appearance gest) on the basal columellar area and parallel with that of Sassia with a dis- left edge of anterior canal; interior of tinctive, distorted shell form and an outer lip bearing a row of seven of eight anatomy and radula more like those of short radial ridges, all low and narrow Ficidae than of Ranellidae necessitate except for the third from top, which is recognizing Personidae as a sixth family elevated into a large, rounded nodule. of the Tonnoidea. Evolutionary stasis Remarks : The new genus Personopsis is exemplified by the remarkably low has formerly been confused with Sassia generic diversity of the family and the (Personella), from which it differs in long time range of unchanged shell form having slightly distorted (rather than in the most diverse genus, Distorsio, regular) coiling, more strongly cancellate which has existed from at least lower sculpture with more closely spaced spiral Campanian to Recent. cords, a shorter anterior canal and more The genera now included in Family gradually tapered last whorl and, in Personidae are: (1) Genus Distorsio particular, a much more widely spread Roding, 1798 (synonymy: Beu, 1987, p. inner lip shield and a more enlarged 310); (2) Genus Distorsionella Beu, third (rather than second) outer lip nod- 1978 (p. 38 ; type species : Distorsio ule. These apertural characters give (Distorsionella) lewisi Beu, 1978, Recent, Personopsis a close resemblance to Dis- Reinga Ridge, Norfolk Ridge and Ker- torsio, and are assumed to express a close madec Islands, north of New Zealand); phylogenetic relationship betweert the (3) Genus Personopsis n. gen.; (4) two genera. Personopsis differs from Genus Kotakaia n. gen. Distorsio in its markedly smaller size, its less distorted coiling, its less widely flared inner lip, its less prominently Genus Personopsis n. gen. armed aperture, and in bearing the basal Type species: Triton grasi Bellardi, 1872, columellar ridges directly on the Pliocene, Italy. columella, rather than on the highly Diagnosis: Shell small (to 25 mm elevated basal columellar callus-ridge high in P. grasi ; not over 20 mm high in that is characteristic of Distorsio. Per- other species), with slightly irregular sonopsis differs from Distorsionella in its whorl coiling; sculpture of regularly smaller size (D. lewisi (Beu) reaches 38 cancellate spiral cords and axial costae, mm high), its shorter and wider form and all relatively low and closely spaced, less gradually tapered last whorl, its sx with several orders of closely spaced shorter and mu$k |iarrower anterior interstitial spiral threads; terminal canal, and its more.. widely flared inner varix low and narrow but obvious, ear- lip ; the two genera have similar apertur- lier varices low and indistinct, situated al armature, except that the second regularly at each 0.66 whorls ; last whorl (rather than third) inner lip nodule is tapering gradually to a moderately short, enlarged in Distorsionella. open anterior canal that curves gently to Species referred to Personopsis at present left and dorsally ; inner lip spread wide- are: ly (but much less so than in Distorsio) to 1. Personopsis rutoti (Vincent, 1930), form a thin, adherent shield over parietal Montian (Mid-Paleocene), Belgium and area, over lower third to half of last Poland. See Krach (1963, p. 102, pi. 23, whorl to left of aperture, and over fig. 6) for reference and apertural charac- columella, smooth except for one or two ters. Ranellidae and Bursidae 91

2. Personopsis ? alvaradoi (Villalta, outer lip. It is assumed tentatively that 1956, p. 182, pi. 7, fig. 4a, b), late the small size has necessitated the loss of Lutetian-early Bartonian (Middle-Late the uppermost two outer lip nodules and Eocene), blue marls at Isun, Spanish adapical movement of the columellar Pyrenees , (Villalta, 1956, p. 120). De- nodules in P. pusilla, but another new scribed in the genus Distorsio, and genus might prove necessary for this although the weakly distorted coiling distinctive species. suggests a position in Personopsis, the tall spire makes a position in Sassia just as likely. Apertural characters (not Genus Kotakaia n. gen. visible on the broken holotype) must be Type species: Kotakaia simplex n. sp., mid-Late determined before tm> ^f^tiily position is Paleocene, Pitt Island, Chatham Islands, New- clear. Zealand. 3. Personopsis beui (Maxwell, 1968, Diagnosis: Shell small (to 20 mm p. 135), Late Eocene, New Zealand. Max- high), with regularly coiled whorls; well (1968) provided a useful review of sculpture of simple low spiral cords (no the distribution of Personopsis, under the axial sculpture visible); low, wide var- name Distorsio (Personella). ices every 0.66 whorls ; a short, straight, 4. Personopsis interposita (Tate, widely open anterior canal deflected 1894, p. 172, pi. 10, fig. 3), Oligocene, strongly to the left; and a heavy aper- Torquay, Port Phillip, Victoria. P. tural armature of high, narrowly interposita has a very, weakly rounded ridges, four inside the outer lip, differentiated, elevated basal columellar one on the parietal area, and two (and, ridge (bearing the basal columellar nod- on some specimens, a very weak third) on ules) and slightly more distorted coiling a large, protruding callus-pad at the top than other Personopsis species, but agrees of the anterior siphonal canal. in size, sculpture, and most apertural Remarks : The new genus Kotakaia characters with Personopsis. bears the large basal columellar-upper 5. Personopsis grasi (Bellardi, 1872, siphonal canal callus-pad, coarsely p. 262, pi. 14, fig. 18), Pliocene, Italy (PI. nodulous and protruding strongly into 3, fig. 11); a relatively large species the aperture, that is the primary charac- which, however, agrees with Paleocene to ter of Distorsio Roding, 1798, and is Oligocene species in all other significant absent from its close relatives Personop- characters. sis n. geri. and Distorsionella Beu, 1978, 6. Personopsis (?) pusilla (Pease, in which the small basal columellar 1861, p. 397), Recent, western Pacific ridges are situated directly on the (Amami Islands to New Caledonia). P. columella. Kotakaia differs from all other pusilla resembles other Personopsis Personidae in having completely regular species in sculpture and general appear- coiling (not strongly distorted, with a ance but is still smaller (rarely over 9 large bulge opposite the aperture in each mm high; largest seen 12.5 mm high) intervariceal interval, as in Distorsio). and differs in its much narrower shape, It has sculpture of simple spiral cords its smaller and narrower aperture, its (prominent axial costae cancellate the longer and narrower anterior canal, its spiral cords in all other Personidae); a very narrow inner lip, in having the short, wide teleoconch, with evenly "basal" columellar nodules enlarged and rounded whorls and a short, widely open situated on the middle to basal columel- anterior siphonal canal; and a much lar area, and in having only five nodules more coarsely and simply armed aperture (the uppermost the largest) inside the than all other Personidae, strongly con- 92 A.G. Brn stricted by similar large, simple ridges another rounded "channel" only slightly protruding from both lips. The apertur- larger than the upper one. The al ridges extend as far into the aperture protoconch is missing from all specimens. of the three prepared specimens as the Dimensions: height (spire incom- hard matrix has been removed (at least 5 plete) 18.2 mm, diameter 12.9 mm mm). (holotype); height (incomplete) 19.5 Etymology: I have great pleasure mm, diameter (incomplete) 12.4 mm naming the new genus Kotakaia in hon- (paratype, TM6787); height (incom- our of Professor Tamio Kotaka ("Tam" plete) 15.7 mm, diameter (incomplete) to his many New Zealand friends) of 10.5 mm (paratype, TM6786). Tohoku University, Sendai. Repository : Holotype (TM6785) and four paratypes (TM6786-TM6789) in Kotakaia simplex n.sp. NZGS. Locality. NZGS loc. 12173, N.Z. PI. 3, fig. 14-17. National Fossil Record no. CH/f478, cal- Description : Most important charac- careous Red Bluff Tuff beneath Matan- ters are stated above. There are six ginui Limestone in northern cliff of main spiral cords, all low, wide and with Rocky Side Bay, on west side of neck of convex surfaces, on the spire whorls, 12 Tarawhenua Peninsula, Pitt Island, on the last whorl and base; low, wide Chatham Islands; collected A.G. Beu? secondary cords fill all spiral interspaces P.A. Maxwell, H.J. Campbell, February on well preserved specimens (the 1977 ; age at this locality mid-late Teur- holotype is slightly abraded). The var- ian (Montian-Landenian), Mid-Late ices are very low, but have an abruptly Paleocene (Fasciculithus tympaniformis raised abapertural margin; the adaper- total range zone; pers. comm. A.R. tural margin is low, wide, and descends Edwards, NZGS). gradually. The inner lip is narrow and Remarks: The volcanogenic matrix strongly thickened, with a squarely at Rocky Side Bay and nearby localities raised, adherent left edge. The outer lip on northern Pitt Island contains an un- also is strongly thickened inside, and usual fauna of epifaunal gastropods, a tapers rapidly outwards to a smooth few bivalves and brachiopods, common margin. The uppermost and largest barnacles in a few places, and diverse ridge on the basal columellar callus-pad ahermatypic corals, helping to fill what is a truly spiral ridge, extending as far is otherwise a partial faunal vacuum in into the aperture as the matrix has been the extensive sequences of early Cen- removed. The parietal ridge and the ozoic rocks exposed in New Zealand. uppermost ridge inside the outer lip con- Several other strffinjltaxa remain to be strict an obvious, narrowly rounded pos- described from this fauna, which terior siphonal canal, and the edge of the includes diverse Cyraeacea for a New outer lip is shallowly embayed at this Zealand fauna (four taxa); this is the point on the holotype; it appears that source of Bernaya chathamensis Cerno- the embayment is largely due to break- horsky (1971, p. 117), from Flowerpot age of the lip, which is thinnest at this Bay, nearby on northern Pitt Island point. The parietal ridge and the upper- (P.A. Maxwell, pers. comm.); Red Bluff most basal columellar ridge constrict Tuff crops out also in Flowerpot Bay.

ACKNOWLEDGEMENTS This paper relied particularly on much help from many friends in California: Ranellidae and Bursidae 93

Judy Terry Smith, Palo Alto; Wyatt Polar Studies, Ohio State University); Durham, Carole Hickman, Liz Nesbitt, and David Dockery, Mississippi Bureau and Joseph Peck, UCMP; Louie Marin- of Geology. Tony Edwards, NZGS, pro- covich and Warren Addicott, USGS, vided data on calcareous nannofossils. Menlo Park; Dick Squires, California Phillip Maxwell, NZGS, and Judy Terry State University, Northridge ; Jim Smith commented on the manuscript. McLean, LouElla Saul, and Dr E.C. Wendy St George, NZGS, took the photo- Wilson, LACM; and Barry Roth, then graphs ; Ron Brazier, NZGS, drew the of CAS. I am grateful, also, for access protoconchs; and Alison Lee typed the to collections, loan of types and helpful manuscript. The - Department of discussion, to ^jprnj^n Sohl, Warren Scientific and Industrial Research pro- Blow, Druid Wils6n and Tom Waller, vided partial fares for me to visit the USNM; Ron Cleevely, Noel Morris and listed museums during 1979,1981,1984 Pat Nuttall, BMNH; Bill Zinsmeister, and 1987. Purdue University (then of Institute of

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