A Taxonomic Review of California Holocene Callianax (Olivellidae: Gastropoda: Mollusca) Based on Shell Characters

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A taxonomic review of California Holocene Callianax (Olivellidae: Gastropoda: Mollusca) based on shell characters

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Charles L. Powell II, Fred Vervaet and David Berschauer

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FRONT COVER: Callianax biplicata photographed by Robyn Waayers, in situ, on July 6, 2016, Clam Beach, Humboldt County, California. Photo used with permission. All rights reserved. (Cover artistic credit: Publication date: March 1, 2020 Rex Stilwill). © San Diego Shell Club, Inc. ISSN 0738-9388 2

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A taxonomic review of California Holocene Callianax (Olivellidae: Gastropoda: Mollusca) based on shell characters

Charles L. Powell, II 1, Fred Vervaet 2 and David Berschauer 3 1 U.S. Geological Survey, 345 Middlefield Road, Menlo Park, CA 95111; [email protected] 2 Boekenburglaan 12, 2215 AD Voorhout, the Netherlands; [email protected] 3 25461 Barents Street, Laguna Hills, CA 92653; [email protected]

ABSTRACT Type specimens and (or) photographs of the 21 named taxa used for California Holocene olivellids were examined to recognize four modern species now assigned to the genus Callianax and one species presumed extinct. The California Holocene species include C. alectona, C. biplicata, C. diegensis, and C. strigata, while the extinct species is Olivella pedroana. Other olivellid names previously used in California are also discussed and these include O. baetica, O. biplicata alba, O. biplicata angelana, O. biplicata brunnea, O. biplicata fucana, O. biplicata lapillus, O. biplicata parva, O. boetica, O. b. diegensis, O. b. mexicana, O. dama, O. glandinaria, O. intorta, O. minuta, O. nota, O. porteri, and O. pycna. Olivella dama and O. intorta are valid species that occur further south in Mexico and are not present in California today despite reports. Olivella nota is also a valid name but for a west Pacific species.

KEY WORDS Olivellidae, Callianax, Olivella, California, Holocene, morphology

McLean (2007) assigned the northeastern examination of these specimens is beyond the Pacific species of Olivella to the genus scope of this paper, so we report their Callianax H. and A. Adams (1853), which he occurrences with little taxonomic verification. elevated from a subgenus of the tropical to subtropical genus Olivella Swainson (1831). He This report is based on shell morphological also designated three Callianax species from the characters of type and supplementary specimens Holocene1 of California and the northeastern attributed to the modern molluscan fauna of the Pacific, C. alectonica (as O. baetica), C. northeast Pacific. Charles Powell contributed biplicata, and C. strigata (as O. pycna). In most of the writing, Fred Vervaet aided in California, the names for many species and revising species descriptions and taxonomic subspecies have been used over the years and discussions, while David Berschauer provided the aim here is to resolve the confusion most of the photographs used in the plates. surrounding these names by discussing and illustrating type specimens (where possible) for each name. Fossil occurrences from the literature have also been cited. However, the

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1 The Holocene ( /ˈhɒləˌsiːn, ˈhoʊ-/) is the current geological epoch. It began approximately 11,650 calendar years before present, after the last glacial maximum, and continues today (Wikipedia, https://en.wikipedia.org/wiki/Holocene, retrieved 4/2019). ISSN 0738-9388

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Abbreviations used and collections examined. ANSP — Academy of Natural Sciences of Drexel University, Philadelphia, Pennsylvania. CASIZ — California Academy of Sciences, Invertebrate Zoology and Geology, San Francisco, California. LACM — Natural History Museum of Los Angeles County, Malacology Department, Los Angeles, California. LACMIP — Natural History Museum of Los Angeles County, Invertebrate Paleontology Department, Los Angeles, California. LIVCM — National Museum Liverpool, Liverpool, England MNHN — National Natural History Museum, Paris, France. NHMUK — British Museum of Natural History, London, England. SBMNH — Santa Barbara Museum of Natural History, Santa Barbara, California.

METHODS tested using gross anatomy (including radula), ecology, biogeography, or the fossil record. All olivellid taxonomic names used over nearly 200 years for Holocene California species are Genus Callianax H. and A. Adams, 1853 evaluated by examining their original descriptions, type photographs where possible, The genus Callianax is presumably named for as well as specimens in some museums and the Greek physican Calli’anax (Καλλιάναξ) who private collections. Modern northeastern Pacific probably lived in the third century B.C. He was Callianax species are described here using the one of the followers of Herophilus, and was features illustrated in Figure 1. In addition, known for the roughness and brutality of his literature related to California Quaternary manners towards his patients. Callianax/olivellids are evaluated and some fossil occurrences are evaluated. Taxa are listed Olsson (1956) places Callianax as a subgenus alphabetically in the systematic section with within Olivella. He defined Callianax as a large valid species listed first and in bold typeface, to medium sized shell, variously colored, with a while non-valid or non-native species names are pillar structure of a simple fold at the base of the listed secondary, are discussed, and are either columella, which is sometimes lirate. Recently excluded from the California malacofauna or McLean (2007) raised the subgenus Callianax put into synonymy with the species we consider to genus level for its smooth columella, with valid. one fold anteriorly; the callus in advance of the aperture extending slightly above the lip of SYSTEMATICS mature shells, but not to the suture above and with an operculum. Callianax is a cooler water Phylum Mollusca Linnaeus, 1758 genus as opposed to Olivella which is Class Gastropoda Cuvier, 1795 subtropical to tropical. As a fossil the genus has Order Neogastropoda Wenz, 1938 been reported from the USA (California, Family Olivellidae Troschel, 1869 Oregon and Florida), Mexico and Colombia Subfamily Olivellinae Troschel, 1869 (http://fossilworks.org/bridge.pl?a=taxonInfo&t axon_no=11488, visited 5/2018) in rocks from We do not follow Kantor and others (2017) the Eocene (Clark and Durham, 1946; Richards because their molecular phylogeny has not been ISSN 0738-9388

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and Palmer, 1953; Puri and Vernon, 1964; Scarlato, 1967]) and the subtropical western Palmer and Brann, 1965) to Quaternary.2 Atlantic (C. moorei [Abbott, 1951] and C. Modern occurrences of the genus are from the thompsoni [Olsson, 1956]; fide Olsson, 1956). northeastern Pacific (herein), the northwest The type species of Callianax is Oliva biplicata. Pacific (C. borealis [Golikov in Golikov &

Figure 1. Apertural view of Callianax showing the shell features used to describe shells.

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2 The Quaternary ( /kwəˈtɜːrnəri/) is the current and most recent of the three periods of the Cenozoic Era in the geologic time scale. It follows the Neogene Period and spans from 2.588 ± 0.005 million years ago to the present. The Quaternary Period is divided into two epochs. the Pleistocene (2.588 million years ago to 11.7 thousand years ago) and the Holocene (11.7 thousand years ago to today) (Wikipedia, https://en.wikipedia.org/wiki/Quaternary, retrieved 4/2019). ISSN 0738-9388

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NORTHEASTERN PACIFIC although becoming broader with age and a CALLIANAX SPECIES white to purplish protoconchs.

Species considered valid are listed first and Original description.“"Histoire naturelle have their names bolded, while those not générale et particulière de tous les genres de considered part of the modern California coquilles univalves marines à l’état vivant et olivellid fauna are listed secondary and are fossile, publiée par monographie. Genre Oliva." either invalid (not bolded) or do not occur in the Duclos’s 1835 illustrations are reprinted below Holocene of California. Many of these invalid (Figure 2). names are put into synonymy, while O. dama, O. intorta, and O. nota are valid but not found living in California and O. pedroana is believed to be extinct.

CALIFORNIA CALLIANAX

Callianax alectona (Duclos, 1835) (Figures 2, 3)

Oliva alectona Duclos, 1835, [vi] Genre Olive. Pl. 4, fig. 15, 16. Olivella baetica Carpenter, 1864, p. 537, 541, 590, 661; Reprint 1872, p. 23, 27, 76, 100, 147, nomen nudum. Figure 2. Type illustration of Oliva alectona Duclos, 1835, pl. 4, Oliva baetica Marrat in Sowerby, 1871, p. 35, fig. 15, 16. From http://olivirv.myspecies.info/en/taxonomy/ pl. 350, figs. 409. term/610, visited 6/2018. Used with permission. Olivella boetica is a misspelling of O. baetica [J.L. Baily, Jr., in lit., Burch and Burch (1959, p. 8)]. Olivella (anazora Ducl. var.?) porteri Dall, 1910, p. 133-134 (unfigured). Olivella boetica mexicana Oldroyd, 1921, p. 118, pl. 1, fig. 3k.

Types. Holotype — unknown. Syntype — MNHN-IM-2000-1230, 4 syntypes; MNHN- IM-2000-1445, 1 syntype.

Type locality. Unknown.

Diagnosis. A narrow shell with a moderate to high spire, a weak to strong suture at the bottom of its whorls and a siphonal fasciole with a Figure 3. Syntype of Oliva alectona Duclos. Height 16.2 mm. single strong plait, pointed at both ends Courtesy of National Natural History Museum, Paris, France. ISSN 0738-9388

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Revised description. The shell is slender to Discussion. This species is reported from moderately wide with respect to height, spire California by Sally Kaicher (Kaicher, 1987, moderate to high, channeled, commonly with a card OL2-5001; date of publication fide white or, seldom, purplish protoconch. A typical Rosenberg and Petit, 2003) and the following parietal callus is often weakly developed and web sites: EOL (http://eol.org/pages/580180 commonly extends past the anal canal. The anal /overview, visited 5/2018), Gastropods.com canal points upwards but may have a slight twist (http://www.gastropods.com/4/Shell_71574.sht either right or left at its end. Siphonal callus ml, visited 5/2018), and http://olivirv.myspecies. generally ends somewhat past the middle of the info/en/taxonomy/term/610, visited 5/2018). On aperture and may blend or be distinct from the initial inspection of Duclos’s (1835) illustration parietal callus. The top margin of the siphonal it does not appear to match well with C. baetica, callus points moderately downward towards the as the former appears to have a callus that goes abaxial side and has one sulcus across it. The up the outer lip on the adapercular side and the siphonal fasciole has a moderate to strong offset siphonal callus goes further up the aperture of from the siphonal callus, is pointed and curved the shell, features that do not match any at both ends with a strong to moderate plait. California Callianax. However, Vervaet (2018) Coloration varies from tan to light yellow with a illustrates syntypes of C. alectona, specimens variable pattern of darker brown wavy lines attributed to C. baetica, along with the neotype sometimes present at the top of a whorl, of O. pedroana, and concluded they are all the sometimes on the lower part of a whorl, and same variable species. We believe O. pedroana sometimes absent. Specimens can be up to is a separate extinct species based on its distinct about 23 mm in length. siphonal fasciole that is not seen in modern specimens. Given Vervaet (2018) syntypes Geographic range. Prince William Sound which are typical of C. alectona, we accept the (60.7°N; CASIZ collections), Alaska, south to remainder of his findings. Estero Beach, Ensenada, Baja California (31.9°N; CASIZ collection) at water depths Olivella porteri, described by Dall (1910), was between the intertidal zone and 150 m (CASIZ considered as synonymous with Callianax collections). South of Point Conception, Santa baetica (=C. alectona) by Burch and Burch Barbara County Callianax alectona is partially (1959) and that is accepted here. Olivella replaced by C. diegoensis (McLean, in lit. 2016; boetica is an incorrect spelling of the species CASIZ collections). In CASIZ collections and name because the edition of Carpenter’s work in as pointed out by Oldroyd (1921, p. 117) which the name was first proposed used a type- specimens from Washington to Alaska face in which the combinations of letters (ae) commonly grow larger and can be beautifully appear to be a single character which looks like colored in cream with brown zigzag stripes; an ‘o’ [J.L. Baily, Jr., in lit., Burch and Burch they occasionally also show erosion of the spire (1959, p. 8)]. The species name boetica has no as seen in the type illustration of Oliva baetica taxonomic significance. Marrat in Sowerby, 1871, pl. 23, fig. 409. The type of Callianax diegensis is distinct from Fossil occurrences. There are no fossil typical forms of this species mainly in lacking occurrences under this name. See Olivella channels on the spire. However, C. alectona is baetica. highly variable and some lots at CASIZ appear similar to C. diegensis. We feel that C. ISSN 0738-9388

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alectona-C. diegensis are a species complex that Original description. O. testâ ovali, griseo- needs further study as the former shows fulvescente, longitudinaliter substriatâ, laevi; remarkable variability over a broad geographic spirâ subacuminatâ, suturâ subfuscâ; columellâ range. For the time being, they are considered laevi, supernè callosâ, ad basim biplicatâ; separate species until such time as integrated apertura, columellae basi, cinguloque basali morphometric, internal anatomy, DNA, or violaceo tinctis; long. 1 lat 6/10 unc. ecological studies show otherwise.

Callianax biplicata (G.B. Sowerby I, 1825) (Figure 4)

Oliva biplicata G.B. Sowerby, I, 1825, p. 87, xxiii-xxxiv. (Reeve, 1850 [1843-1878], 6. Oliva, pl. 20, f. 3). “Olivella glandinaria (Nutt.) MS.” Carpenter, 1856, p. 209-229. Olivella biplicata var. alba Williamson, 1892, p. 212 [nomen nudum]. Olivella biplicata var. brunnea Williamson, 1892, p. 212 [nomen nudum]. Olivella biplicata lapillus Vanatta, 1915, p. 71. Olivella biplicata angelena Oldroyd, 1918, p. 34. [misspelled as O. b. angelina by Oldroyd, 1921; 1927]. Olivella biplicata fucana Oldroyd, 1921, p. 118, pl. 5, fig. 4. Olivella biplicata parva Oldroyd, 1921, p. 119, pl. 5, fig. 7. Figure 4. Callianax biplicata (G.B. Sowerby I, 1825). LACM hypotype 114745. Shelikof Bay, Kruzof Island, Sitka County, Alaska. Height 27.8 mm. Photograph by J.H. McLean, provided Types. Of Oliva biplicata — unknown. No by LACM Malacology. neotype is designated because the authors do not consider a name-bearing type is necessary to Revised description. The shell is relatively define this species (ICZN Code, article 75). large (to about 30 mm), broad, robust, with moderate to poorly channeled whorls. Color is Type locality. “West coast of North America” variable, commonly from light gray to purplish (G.B. Sowerby I, 1825). gray, occasionally all white, brownish gray, tan, or even dark purple, without a pattern. The Diagnosis. Shell large (to 30 mm), broad and columellar callus is relatively strong, extending robust, with a broad parietal callus and small, from aperture to fasciole, 2-3 small folds at the narrow, siphonal fasciole pointed at both ends base often with several incised spiral lines. The separating this species from other northwest siphonal fasciole is pointed at both ends, Pacific Callianax. ISSN 0738-9388

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somewhat broadly at the base commonly with a Pliocene/Pleistocene deposits from central and siphonal plait. southern California by Cooper (1888), Arnold and Hannibal (1913), Oldroyd (1921), Grant Geographic Range and Ecology. In and Gale (1931), and Powell and Stevens (2000). CASIZ/LACM collections this species is It is common in the Middle and Late Pleistocene represented by specimens from the Gulf of fossil record through much of coastal southern Alaska (LACM) south to Cabo San Lucas, Baja California (Cooper, 1888; Arnold, 1903; Chace California Sur, Mexico (22.9°N; CASIZ) and and Chace, 1919; Grant and Gale, 1931; into the Gulf of California to La Paz (24.1°N; Stephens, 1929; Woodring and others, 1946; CASIZ), at water depths from the intertidal zone Vedder and Norris, 1963; Addicott, 1966; to 65 m (35 ftms; CASIZ). It is not as common Russell, 1991; Powell and Stevens, 2000; in collections from north of Washington State Powell and others, 2005), Baja California, than south. Ecologically it is common at lagoon Mexico (Jordan, 1924; 1926; DeLong, 1941; entrances and protected sandy areas of the open Chace, 1956; Valentine, 1956; 1957; Addicott, coast at low tide, sometimes abundant in 1959) and Oregon (Arnold and Hannibal, 1913). shallow water offshore along exposed sandy beaches. Hickman and Lipps (1983) found that Discussion. Callianax biplicata is easily it feeds on foraminifera. distinguished from other west coast Callianax by it large size and being wider at similar Fossil occurrences. Likely late, possibly lengths than other species. In addition, it has a middle Miocene to Holocene. The earliest fossil very short and wide spire for its size. occurrence of middle Miocene age is reported from the Empire Formation from Cape Blanco, Reproduction is detailed by Edwards (1968). Curry County, Oregon (Arnold and Hannibal, Commonly brought into the rocky intertidal 1913), but this occurrence and others cannot be zone by hermit crabs (Valentine, 1961; Onuf, confirmed until specimens from these various 1972). Shells thus inhabited by hermit crab can formations are examined. Other Miocene sometimes be recognized by the trace fossil occurrences are from the late Miocene Santa Helicotaphrichnus (Walker, 1992). This species Margarita Sandstone (“Margaritan” California was used by native Californians and traded provincial molluscan stage [CPMS]) in central among various tribes (Hartzell, 1991; Eerkens California (Addicott and Vedder, 1963; and others, 2005; 2009). Addicott and others, 1978; Powell, 2008). Occurrences in formations assigned to the Miocene/Pliocene include the Etchegoin (Grant and Gale, 1931), Purisima (Nomland, 1917, Grant and Gale, 1931, as lower Merced Formation; also Powell, 1998), and Wilson Grove (Osmont, 1905; Dickerson, 1922; Powell and others, 2005) formations. From Pliocene deposits it is reported from California (Woodring & Bramlette, 1950 [1951], and Addicott, 1969), Baja California, Mexico (Ashby and Minch, 1984), and Oregon (Arnold and Hannibal, 1913). It has been reported from ISSN 0738-9388

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Callianax diegensis (Oldroyd, 1921) Diagnosis. A narrow shell with an unchanneled (Figure 5) to slightly channeled spire, purplish protoconch, and a broad siphonal fasciole. However, these Olivella boetica diegensis Oldroyd, 1921, p. characteristics are plastic and may overlap with 118, pl. 5, fig. 2. some specimens of C. alectona when diverse samples are examined. Types. Holotype — CASIZ 064353. Paratype — CASIZ 067156 (23), 067157 (77), Original description. “This differs from the SBMNH 34841 (1) (paratypes, from the modern typical shells of British Columbia, in the color fauna), CASIZ 66181.01 (many) (paratypes, being a light drab, sometimes mottled; not as from the Late Pleistocene at Santa Monica, Los oval in outline, spire longer and running more Angeles County, California), CASIZ 66183.01 sharply to a point. Length, 19; breadth, 8 mm.” (100) (paratypes from the Late Pleistocene at (Oldroyd, 1921). San Pedro, Los Angeles County, California). Revised description. Shell of moderate height (to 22 mm fide P. Recourt, in litt., 11/2019), narrow, robust, variously colored but tending towards off white, tan, and brown, without a pattern and lacking a channel at the base of the whorls. Protoconch and nuclear whorls bluish- purple to purple. Parietal callus relatively weak, extending from above the aperture to past the siphonal fasciole with a single fold at the base of the upper part of the siphonal fasciole. The siphonal callus shows a distinct band at its upper end ending at the siphonal fasciole. A single plait is present, situated low on the top margin of the siphonal fasciole.

Modern occurrence. The distribution of this species is not well known but it has been reported from San Pedro (CASIZ) to San Diego, southern California (Oldroyd, 1927). McLean (written communication, 2016) suggests that it overlaps and may eventually mostly replaces C. alectona in the Southern California Bight. Depth data is not sufficiently recorded. Figure 5. Calllianax diegensis (Oldroyd, 1921). CASIZ holotype 064353. San Diego, San Diego County, California. Height 19 mm. Photograph by J.H. McLean, provided by Fossil occurrences. Reported by Oldroyd (1925) LACM Malacology. from the upper San Pedro (=Palos Verdes Sands) in San Pedro, Los Angeles County. This Type locality. San Diego, San Diego County, occurrence needs to be verified. California. ISSN 0738-9388

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Discussion. This species was placed in synonymy with Callianax baetica by Burch and Burch (1959, p. 9). However we feel that the lack of a channel at the base of each whorl, the shape of the strong siphonal fasciole, and the bluish-purple to purple protoconch and nuclear whorls sets it apart from C. alectona.

Callianax strigata (Reeve, 1850) (Figures 6, 7, 8)

Oliva strigata Reeve, 1850, pl. 25, fig. 72 a-b. Oliva baetica Marrat in Sowerby, 1871, p. 35, pl. 350, fig. 410. Olivella pycna Berry, 1935, 262-265.

Type. Nine syntypes at BMNH [Natural History Museum (2014). Dataset: Collection specimens. Resource: Specimens. Natural History Museum Figure 6. Type illustration of Olivella strigata Reeve, 1850, pl. 25, fig. 72. From http://olivirv.myspecies.info/en/taxonomy/ Data Portal (data.nhm.ac.uk) https://doi.org/ term/609, used with permission. 10.5519/0002965]. Revised description. Shell of moderate height 3 Type locality. West Indies, in error. (< 16 mm), moderately wide, robust, with a zigzag pattern, and a distinct channel at the base of Diagnosis. The shell is small, broad (but not as the whorls. Color is variable, with the zig-zag much as Callianax biplicata), robust, and lines always darker. Also the entire shell has variously colored in shades of tan and brown minute spiral sculpture. The parietal callus is with very faint evenly spaces radial lines and a wide, distinct except against the siphonal callus, zig zag pattern that sets this species apart from where it becomes obscure and ends at or about other west coast species. Additionally, the the anal canal. The siphonal callus shows a siphonal callus shows a distinct band at its distinct band at its upper end, a rounded upper upper end and the siphonal fasciole is long and margin and is without plaits while the siphonal narrow with a subdued plait. It also has fine fasciole is sharply rounded at its posterior end spiral sculpture (Figure 7) setting it apart from and the upper margin shows a single plait. all other California Callianax.

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3 The term "West Indies" was used by British Europeans to describe the British West Indies their acquired territories in the Americas, which included much of the Caribbean but also Guyana and Belize. ISSN 0738-9388

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Figure 7. Shows fine radial sculpture and concentric growth lines of Callianax strigata. The field is about 4 x 2.5 mm. Photograph by Bob Abela.

Modern occurrence. Specimens in the collections at CASIZ occur from Morro Bay, San Luis Obispo County, California (35.4°N) to Banks Island, Skeena-Queen Charlotte C, Figure 8. A-H =Oliva strigata Reeve, 1850. Eight (of 10) British Columbia, Canada (53.4°N) (44 lots; syntypes. A = syntypes registration number NHMUK CASIZ collections) in water depths from 15 to 1872.9.24.17. B-H = Seven syntypes registration number 25 ftms (27-46 m; CASIZ collections). There is NHMUK 1987003/2-9. ‘West Indies’. NHMUK 1892.9.24.17 from Steere collection, remaining shells from the Hugh Cuming also a lot (CASIZ lot 29903 [2 specimens]) collection. Scale is in millimeters. Copyright of the Natural reported from La Paz, Baja California Sur, History Museum of London, Images by Kevin Webb, NHMUK Mexico, which is assumed to be mislocated. Photographic Unit, provided by Andreia Salvador, NHMUK.

Fossil occurrence. There is no fossil Discussion. Castro Oliveira (2011) considers occurrence under the name Callianax strigata. Callianax strigata as a synonym of Porphyria minuta. Olivella minuta differs from the type illustration of Oliva strigata in having abundant columnar folds and abundant teeth on the inside of the outer lip. These features easily separate the type illustration of C. strigata from O. ISSN 0738-9388

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minuta. Once C. strigiata is removed from as “Quam O. alectona graciliusculior, coloribus synonymy with O. minuta, the type illustration magis nebulatis variegata; balteo casteneo, of O. strigata is nearly identical to the type of O. fasciato.—The colours with which most of the pycna leading us to determine that Oliva specimens are variegated are more clouded, and strigata is an earlier name for O. pycna and the lines more spread into patches than in therefore have to assume that the type locality is Duclos’ figure of O. alectona, but the difference in error. seems hardly essential. ED.”

The geographic ranges of extant California Callianax is shown in Figure 9, below on p. 13.

OLIVELLID TAXA EXCLUDED FROM THE MODERN CALIFORNIA MALACOFAUNA

Olivella baetica (Marrat in Sowerby, 1871) [=Callianax alectona (Duclos, 1835)] (Figure 10)

Olivella baetica Carpenter, 1864, p. 537, 541, 590, 661; Reprint 1872, p. 23, 27, 76, 100, 147, nomen nudum. Oliva baetica Marrat in Sowerby, 1871, p. 35, pl. 350, figs. 409. Figure 10. Type illustration of Oliva baetica Marrat in Sowerby, 1871, pl. 23, fig. 409, 410, however figure 410 is herein Types. Of Olivella baetica Carpenter (1864) — attributed to Callianax strigata. From http://olivirv.myspecies. no type given. Of Oliva baetica Marrat in info/en/taxon-pages/oliva-baetica, visited 1/2019. Used with permission. Sowerby (1871) — no type given. Fossil occurrences. Pliocene to Holocene. Type locality. Of Olivella baetica Carpenter Pliocene/Pleistocene. Merced Formation (1864) — none give. Of Oliva baetica Marrat in (Cooper, 1888), San Diego Formation (Cooper, Sowerby (1871) — Vancouver Island, British 1888) and “San Pedro” Formation (Powell and Columbia, Canada. Holocene. Stevens, 2000). Pleistocene. Lomita Marl (Woodring and others, 1946), Palos Verdes Original description. Carpenter’s (1864) Sand (=upper San Pedro horizon) (Oldroyd, description of “Narrow, dull, thin,” is 1921; Woodring and others, 1946), San Pedro inadequate to separate it from other eastern Formation (Oldroyd, 1925; Woodring and Pacific olivellids and Carpenter’s name is others, 1946), “San Pedro” Formation in Orange therefore a nomen nudum. A few years later and Los Angeles counties (Powell and Stevens, Marrat in Sowerby (1871) describes this species ISSN 0738-9388

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Figure 9. Map showing the geographic distribution of northern eastern Pacific Callianax. Province boundaries of Valentine (1966). ISSN 0738-9388

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2000), Timms Point Silt (Woodring and others, Olivella biplicata alba Williamson, 1892 1946), unnamed Late4 Pleistocene sediments nomen nudum [=Callianax biplicata from southern California (Stevens, 1929; (G.B. Sowerby I, 1825)] Valentine, 1956; Emerson and Chace, 1959; Powell and others, 2005) to Baja California, Olivella biplicata alba Williamson, 1892, p. Mexico (Jordan, 1926; Valentine, 1957). Most 212. of these occurrences are unverified. Type. None given. Discussion. We find the type illustration of Oliva baetica in Sowerby (1871) to represent Type locality. None given. two northeastern Pacific species; figure 409 is referred to Callianax alectona while figure 410 Original description. None given. is attributed to C. strigata. Sowerby’s (1871) illustration figure 409 strongly resembles C. Discussion. This name is found in a list of alectona in having a high-spire with a channel, molluscan species from San Pedro, Los Angeles and a nearly straight siphonal callus at its upper County without description or illustration and is margin. It also resembles C. alectona in having therefore considered a nomen nudum. White a spire that appears eroded, which is commonly specimens of Callianax biplicata are not observed in specimens from localities between particularly rare and are considered a color form Washington and Canada (CASIZ collections). of the nominal species. This color form was also Sowerby’s (1871) figure 410 is attributed here named O. biplicata lapillus by Vanatta (1915). to C. strigata with its lower spire and strongly curved upper margin of the siphonal callus. There are also differences in the size, shape, and Olivella biplicata angelena Oldroyd, 1918 pattern that support this conclusion. [=Callianax biplicata (G.B. Sowerby I, 1825)] (Figure 11) Palmer (1958) found Carpenter’s (1864, p. 661) description of this species inadequate to which Olivella biplicata angelena Oldroyd, 1918, p. we agree. Therefore the species is attributed to 34-35. Marrat in Sowerby (1871) who provided an illustration, a type, and a type locality for Types. Holotype — CASIZ 064312. Carpenter’s name making Marrat the author of Paratypes — CASIZ 064313 (1), CASIZ this species. However, it was first described as 065621 (12), SBMNH 34839 (1) (paratypes C. alectona many years earlier. from the modern fauna), CASIZ 66182.01 (22) (paratypes from the Late Pleistocene at Santa Monica, Los Angeles County, California). CASIZ 66182.01 (22) (paratypes from the Late Pleistocene at Signal Hill, Long Beach, Los Angeles County, California). ______

4 Late is capitalized here because it has been formally described as Late (chronostratigraphically) or Upper (lithologically), see http://www.stratigraphy.org/ICSchart/ChronostratChart2018-08.pdf, visited 1/2019. ISSN 0738-9388

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Type locality. San Pedro, Los Angeles County, Olivella biplicata brunnea Williamson, 1892 California. Holocene. nomen nudum [=Callianax biplicata (G.B. Sowerby I, 1825)]

Olivella biplicata brunnea Williamson, 1892, p. 212.

Type. None designated.

Type locality. None designated.

Original description. “The brown variety is rare, and not found living. Dr. Cooper says O. biplicata ‘varies in color from black through brown, purple, gray and pink to white.’”

Discussion. This name is found in a list of molluscan species from San Pedro, Los Angeles Co. The description is inadequate to distinguish this taxon from the nominal forms of C. biplicata and it is considered a nomen nudum.

Olivella biplicata fucana Oldroyd (1921) [=Callianax biplicata (G.B. Sowerby I, 1825)] Figure 11. Callianax biplicata (G.B. Sowerby I, 1825). CASIZ (Figure 12) holotype 064312, holotype of O. biplicata angelena. San Pedro, Los Angeles County, California. Height 27 mm. Photograph by J.H. McLean, provided by LACM Malacology. Olivella biplicata fucana Oldroyd, 1921, p. 118, pl. 5, fig. 4. Original description. “This variety differs from Sowerby’s type in being more delicate and Types. Holotype — CASIZ 064355. slender, with callous not so heavy, spire more Paratypes — CASIZ 067153(3). elevated, sloping more gradually from the middle of the shell to the apex. Variety Type locality. Near Cape Flattery, Strait of angelena is found fossil in both the upper and Juan de Fuca, Washington (Oldroyd, 1921). lower San Pedro beds [=Palos Verdes Sand and San Pedro Sand, respectively] of the middle to Original description. “Shell broader across the Late Pleistocene. “Length 27, width 13 mm.” middle and lower part of the aperture than var. angeliena; spire running more sharply to a point Discussion. Synonym of Callianax biplicata from the middle of the shell. Color more (Burch and Burch, 1959, p. 20) and we find no uniform, being a light drab. Length, 28; breadth, reason why it should be separated. Olivella 14 mm.” (Oldroyd, 1921, p. 118). biplicata angelina is a misspelling on the name. ISSN 0738-9388

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Figure 13. Callianax biplicata (G.B. Sowerby I, 1825). ANSP holotype 111977, holotype of O. b. lapillus. San Pedro, Los Figure 12. Callianax biplicata (G.B. Sowerby I, 1825). CASIZ Angeles County, California. Height 24 mm. Photograph by J.H. holotype 064355, holotype of O. b. fuscana. Cape Flattery, McLean, provided by LACM Malacology. Clallam County, Strait of Juan de Fuca, Washington. Height 28 mm. Photograph courtesy of Christine Garcia (CASIZ). Original description. “Shell globose, white, spire short, interior of the aperture cream- Discussion. Synonym of Olivella biplicata colored.” “Alt. 24, diam. 14 mm.” (Burch and Burch, 1959, p. 19-20). Reported as a fossil from the Timms Point Silt (Middle Discussion. Vanatta (1915) states that this is Pleistocene) of San Pedro, California (Coan and Olivella biplicata alba of Williamson (1892) Kellogg, 1990). which is synonymous here with Callianax biplicata. He also states that it was illustrated by Sowerby (1880). Olivella biplicata lapillus Vanatta, 1915 [=Callianax biplicata (G.B. Sowerby I, 1825)] (Figure 13) Olivella biplicata parva Oldroyd, 1921 [=Callianax biplicata (G.B. Sowerby I, 1825)] Olivella biplicata lapillus Vanatta, 1915, p. 71. (Figure 14) Type. ANSP holotype 111977. Olivella biplicata parva Oldroyd, 1921, p. 119, pl. 5, fig. 7. Type locality. San Pedro, Los Angeles County, California. Collected by Mrs. E.M. Gaylord. ISSN 0738-9388

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Types. Holotype — CASIZ 064314. the upper Pleistocene in San Pedro. Length, 14; Paratypes — CASIZ 067154 (27), SBMNH breadth, 8 mm.” 34840 (4) (from the modern fauna), CASIZ 66181.02 (8) (from the Late Pleistocene at Santa Discussion. Burch and Burch (1959, p. 19-20) Monica, Los Angeles County, California), placed it in synonymy with Callianax biplicata CASIZ 66182.03 (3) (from the Late Pleistocene and this designation is accepted herein. at Signal Hill, Long Beach, Los Angeles County, Reported as a fossil from Late Pleistocene of California). Santa Monica and Signal Hill, Long Beach, both Los Angeles County, California (Coan and Kellogg, 1990).

Olivella boetica Carpenter, 1864 [=Callianax baetica (Marrat in Sowerby, 1871)]

Discussion. Reported from the northern eastern Pacific by Dall (1878; 1921), Packard (1918), Ellis and Lee (1919), Oldroyd (1927), and Bourne (1984) under this misspelling of the species name. Also used in conjunction with the two subspecies described by Oldroyd (1921) (O. boetica diegensis and O. b. mexicana). Olivella boetica is a misspelling of the species name baetica because the edition of Carpenter’s work in which the name was first proposed used a font in which the combinations of letters in a single character made the ‘a’ look more like an ‘o’” [J.L. Baily, Jr., in lit., Burch and Burch (1959, p. 8)]. The species name boetica has no taxonomic significance. Figure 14. Callianax biplicata (G.B. Sowerby I, 1825). CASIZ holotype 064314, holotype of O. b. parva. Punta Abreojos, Mulegé Municipality, Baja California Sur, Mexico. Height 13 mm. Photograph courtesy of Christine Garcia (CASIZ). Olivella boetica mexicana Oldroyd, 1921 [=Callianax alectona (Duclos, 1835)] Type locality. Punta Abreojos, Baja California (Figure 15) Sur, Mexico. Holocene. (Oldroyd, 1921). Olivella boetica mexicana Oldroyd, 1921, p. Original description. Thomas Oldroyd (1921) 118, pl. 1, fig. 3. described this species as “This var. is nearest angelena, but much smaller, a little broader in Types. Holotype — CASIZ 064354, from proportion; outer lip more curved, shell more Laguna Scammon, Baja California Mexico. highly colored and variable in color; found in Paratypes — CASIZ 66181.03 (15) (from Late Pleistocene at Santa Monica, Los Angeles ISSN 0738-9388

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County, California), CASIZ 66182.02 (43) Los Angeles County, California (Oldroyd, (from Late Pleistocene at Signal Hill, Long 1921). Beach, Los Angeles County, California), CASIZ 66183.02 (160) (from Late Pleistocene Discussion. Synonym of Callianax baetica at San Pedro, Los Angeles County, California). according to Burch and Burch (1959, p. 9) which we follow.

Olivella dama (Wood, 1828)

Voluta dama Wood, 1828, p. 11, pl. 4 Voluta fig. 37.

Discussion. Reported from California by Yates (1886), Cooper (1894), and indirectly by Gardner (1948). Olivella dama has been reported from Holocene archaeological sites in southern California (Gamble and King, 2011). While a valid species, there are no modern representatives of O. dama from California and it is therefore excluded from the modern California Olivellid fauna.

“Olivella glandinaria (Nutt.) MS” Carpenter (1857) [=Callianax biplicata (G.B. Sowerby I, 1825)] (Figure 16) Figure 15. Olivella baetica mexicana Oldroyd, 1921. CASIZ holotype 064354. Laguna Ojo de Liebre (formerly Laguna Scammons), Baja California, Mexico. Height 10 mm. “Olivella glandinaria (Nutt.) MS.” Carpenter, Photograph courtesy of Christine Garcia (CASIZ). 1857, p. 209-229.

Type locality. Laguna Ojo de Liebre Type. NHMUK syntypes 20190057 (four (Scammon’s Lagoon), Baja California Sur, specimens figured below). Mexico. Type locality. “California Superiore” Original description. Reported by Oldroyd (Carpenter, 1857). “W. Coast N. America” fide (1921) as “This differs from var. pedroana in Andreia Salvador (in litt., 2/2018). being smaller, more slender, spire not running quite as sharply to a point. Length, 10; breadth, 4 mm.”

Fossil occurrences. Reported from the Late Pleistocene (=Palos Verdes Sand) of San Pedro, ISSN 0738-9388

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Figure 16 A-D. “Olivella glandinaria (Nutt.) MS” of Carpenter (1857), four syntypes, registration number NHMUK 20190057. ‘W. Coast N. America’. Tankerville collection no. 2332. Scale is 1 cm. Copyright of the Natural History Museum of London, Images by Kevin Webb, NHMUK Photographic Unit, provided by Andreia Salvador, NHMUK.

Original description. “O. t. bulbiformi, in basin biplicata; labio calloso, laevi; callositate medio inflata, utrinque regulariter constricta; basali haud lata.” “Long. 88, long. Spir. 29, lat. spira satis elevata, acuta; haud polita, purpureo- 47, div. 70” (Carpenter, 1857). fusca, in spira aurantia, circa basin violaceo tincta; apertura antice dilatata; columella ad ISSN 0738-9388

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Discussion. As discussed in Palmer (1958) Type locality. “San Juan, California. Dr. “Carpenter (1856) described an Olivella from Green.” According to Palmer (1958) Carpenter ‘upper California,’ which Nuttall had named in definitely put San Juan in the Golfo de manuscript Glandinaria Californica. Gould and California region, Mexico. Carpenter first used the name in 1856. Later Carpenter (1864) identified the species as Diagnosis. Shell small, moderately narrow with conspecific with C. biplicata. We illustrate the a moderately high, unchanneled spire sets this syntypes of O. glandinaria and also find it species apart from others in the northwest equivalent to C. biplicata. Pacific.

Original description. Described by Carpenter Olivella intorta Carpenter (1857) (1857) as “O. t. parva, ovoidea, subtumente; (Figure 17) sutura vix sulcata; albido-grisea, fascia indistincta subsuturali olivacea, flammulis et Oliva intorta Carpenter, 1857, p. 209-224, p. maculis purpureo-fuscis plus minusve ornata; 228-229. apertura antice aperta, postice angusta; callositate parietali ad suturam penultimam Type. Holotype—NHMUK 1950.11.9.3. producta; columella maxime intorta, plica ad basin acuta, in pariete duabus saepe indistinctis; extus, linea spirali antica unica.” “Long. 52, long. Spir. 17, lat. 26, div. 60°.”

Revised description. The shell has a smooth outline and is slightly wider with respect to length than similar-sized Callianax alectona. In aperical view the parietal callosity extending onto the penultimate whorl hiding the color of the spire. The parietal callus is narrow with a short narrow plait near the widest part of the shell and appears to be separated from the siphonal callus, the top of which is nearly straight. The siphonal callus is poorly defined and not easily separated from the parietal callus and appears to be without plaits.

Geographic range. “Magdalena Bay, Baja California, to the southern end of the Gulf of California” (Keen, 1971).

Fossil occurrences. Although reported from formations of late Miocene to the Late Figure 17. Olivella intorta Carpenter, 1857. LACM Hypotype Pleistocene in California, all these reports 1971-181.20. Punta PequeXa, Baja California Sur, Mexico (LACM 71-181.20). Height 11.8 mm. Photograph by J.H. should be questioned until such time as McLean, provided by LACM Malacology. specimens can be checked. Reported from the ISSN 0738-9388

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Miocene to Pliocene Purisima Formation by Discussion. The name Olivella minuta has been Arnold (1908) and Martin (1916) were referred used in California only indirectly as a synonym to Callianax strigata (as C. pycna) by Powell of Callianax strigata, which it is not. Given the (1998). Arnold (1908) and Martin (1916) also geographic range of this species reported by reported it from the Pliocene to Pleistocene Castro Oliveira (2011) it is not part of the Merced Formation. In addition, it has been eastern Pacific olivellid fauna. reported from the Pleistocene of Port Los Angeles, Los Angeles County by Arnold (1903). Olivella nota Marrat, 1868 Discussion. McLean (2007) defined the genus (Figure 18) Callianax by its smooth columella, with one fold anteriorly, a callus in advance of the Oliva nota Marrat, 1868, p. 213. aperture extending slightly above the lip of a mature shell, but not to the suture above, and Types. National Museum Liverpool syntype has an operculum. Olivella intorta has a smooth LIVCM.17.6.1875[512] https://gbmolluscatypes. columella with a single faint fold, which is well ac.uk/specimens/683, visited 6/2018). Natural posterior of other species in Callianax, and it History Museum, London syntype 1924.1.5. lacks an operculum. These characters make its 1657 - 1663 (http://olivirv.myspecies.info/en/ inclusion in the genus Callianax incorrect. node/5554, visited 5/2019).

Reported from California by Cooper (1888), Type locality. None given in the original Keep (1887), Williamson (1892), Arnold (1903; description. Reported as Vancouver island, 1906; 1908), Packard (1918), Baily (1935), Canada by Marrat in Sowerby (1871). Woodring and others (1946), and Smith and Gordon (1948). Packard (1918, pl. 37, fig. 7) illustrated a specimen referred to this species that we referred to C. strigata.

This species is not reported from the Holocene of California and is excluded from California’s modern molluscan fauna.

Olivella minuta (Link, 1807)

Porphyria minuta Link,1807, p. 98.

Types. Lost (Castro Oliveira, 2011).

Type locality. Caribbean (Castro Oliveira, 2011).

Geographic range. South Caribbean Sea to Figure 18. Oliva nota. Left. Oliva nota Marrat, 1871, pl. 351, fig. 428. Right. Olivella nota, Syntype LIVCM.17.6.1875[512] Paraná State, Brazil (Castro Oliveira, 2011). from https://gbmolluscatypes.ac.uk/specimens/683, retrieved 6/2018. ISSN 0738-9388

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Original description. "Shell ovate fusiform, rather inflated; spire considerably exserted, canaliculate; suture with large brown blotches; white, with broad wavy brown lines, which become darker about the belt; belt white; columella rounded, basal band white; a single deep-purple fold at the base; the pattern of the shell is seen through in the interior."

Discussion. The type illustration of Olivella nota (Figure 18) superficially resembles a mature Callianax alectona while the syntype resembles an immature specimen of that species. Burch and Burch (1959) determined Olivella nota to be a synonymy of C. baetica (= C. alectona), however Vervaet (2018) recently referred it to the Japanese species O. nota based on the slight purple staining on the columella fold that he states makes it distinct from other west Pacific species. The type locality as reported by Marrat in Sowerby (1871) is given as Vancouver Island, Canada. Also C. alectona Figure 19. Olivella pedroana Conrad, 1855. USNM neotype (along with C. biplicata and C. dama) are 498559. San Pedro, California. Palos Verdes Sand (Late Pleistocene). Height 12.4 mm. Photograph by J.H. McLean, known to have light purple staining on the provided by LACM Malacology. siphonal fasciole in some specimens. Even given these features we follow Vervaet (2018) Type locality. San Pedro, California. Post- and recognize O. nota as a valid species from Pliocene or recent formation. Conrad’s (1855) the western Pacific Ocean. post-Pliocene or recent formation was interpreted by Woodring and others (1946) as “near the mouth of the Los Angeles River ... ǂ Olivella pedroana (Conrad, 1855) before the lower course of the river ... from the (Figure 19) Palo Verdes Sand, Late Pleistocene in age.”

Strephona pedroana Conrad, 1855, p. 17. Diagnosis. The distinctive siphonal fasciole and Stephona pedroana, Conrad, 1857, p. 327, pl. small siphonal callus without plaits on either 6, fig. 51. make this species easily identifiable. In addition, the moderately high spire, parietal callus that is Types. Conrad’s (1855) type lost. Woodring more or less an even distance across and and others (1946, pl. 35, fig. 22; USNM 498559) terminating at the anal canal makes this species designated a neotype. distinguishable from other California Holocene species. ISSN 0738-9388

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Original description. Conrad (1855) 1917) formations [later (Powell, 1998) referred description of Olivella pedroana consisted of all O. pedroana from the Purisima to C. strigata “Small, elliptical; spire conical, about equal in (= C. pycna)]. Pliocene reports are from length to the aperture; base of columella with a unnamed deposits at Elsmere Canyon (Arnold, prominent fold.” Woodring and others (1946) 1907) [later updated as C. baetica from the Pico selected a neotype for Conrad’s species and Formation by Squires, 2012; now attributed to C. described it as “A small, moderately slender alectona] and Holser Canyon, Los Angeles species that as a moderately thick and definitely County (English, 1914). Pliocene to Pleistocene limited parietal callus is identified as O. reports of this species are from the Merced and pedroana.” San Diego formations, the “San Pedro” Formation in Orange and Los Angeles counties Revised description. Shell of moderate height (Hoskins, 1954; Yerkes, 1972). The latter (to about 20 mm), moderately narrow, robust, occurrence was attributed to C. baetica by without a pattern, and having a distinct channel Powell and Stevens (2000) and are herein at the base of the whorls. The parietal callus is referred to C. alectona. Reports from the distinct, with a relatively straight ventral margin Pleistocene include the Palos Verdes Sand ending at the anal canal. It is half-moon shaped (=upper San Pedro) and San Pedro Formation siphonal callus is small, indistinct, and without (=lower San Pedro; Grant and Gale, 1931) and plaits. unnamed Pleistocene terraces from California (Stephens, 1929; Addicott, 1966 [this latter Geographic range. Determining an accurate reference referred to C. pycna (= C. strigata) geographic range for this species is beyond the above]) to Baja California, Mexico (Jordan, scope of this paper and the use of the name O. 1924; 1926). pedroana is confused in the literature. Discussion. Reported living off California by Fossil occurrences. In the literature, this Dall (1921), Grant and Gale (1931), Woodring species is reported from late Miocene and early and others (1946), and Olsson (1956), this Pliocene to Holocene. However, previous species has been attributed to Callianax authors have not recognized O. pedroana in the alectona at various web sites (http:/ sense developed here so all fossil occurrences /olivirv.myspecies.info/en/taxonomy/term/610, need to be re-examined. Miocene reports visited 5/2018; http://www.marinespecies.org/ include the Empire (Dall, 1909; Arnold and aphia.php?p=taxdetails&id=448234, visited Hannibal, 1913; Howe, 1922), Montesano 5/2019) and by Vervaet (2018). However, it is (Weaver, 1912), San Pablo (Clark, 1915) determined here to be a distinct, extinct species formations, Santa Margarita Sandstone (Powell, that does not occur in the modern California 2008) and Temblor Formation (Anderson and malacofauna (hence the use of the symbol ǂ). Martin, 1914; Clark, 1915). These reports may be attributed to Callianax ischnon Keen (1945) The genetic placement of Olivella pedroana is and/or Olivella subpedroana Loel and Corey in question. It is similar to Callianax alectona (1932) upon further study (Addicott, 1970). in many respects but lacks the anterior fold on Miocene to Pliocene reports are from the the columella a feature characteristic of the Etchegoin (Arnold and Hannibal, 1913; Martin, genus Callianax. Given this we questionably 1916; Nomland, 1917) and Purisima (Arnold place pedroana in the genus Olivella until such and Hannibal, 1913; Martin, 1916; Nomland, time as it can be further evaluated. ISSN 0738-9388

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Olivella porteri Dall, 1910 of the whorls brilliantly polished, smooth, [=Callianax alectona (Duclos, 1835)] except for microscopic spiral close striation (Figure 20) which is present on all, but more uniform on particular individuals; color pale olive, yellow, Olivella (anazora Ducl. var. ?) porteri Dall, or whitish, with sharp angular axial brown lines 1910, p. 133-134. superposed between the suture and the basal fasciole, sometimes forming a tent-like pattern Types. Syntype USNM 209677, Syntype and sometimes reduced to fine, close, more or SBMNH 34782. less cloudy zigzags; a pale band in front of the suture, usually with vivid brown zigzag pointed forward, but sometimes plain, the brown lines when present broader and stronger than elsewhere; basal fasciole short, with a marginal and an adjacent narrow fold or plait anteriorly, lighter than the body, yellow or rich bluish purple; aperture narrow, simple, with a deep sutural sinus and a moderate parental callus when mature. Height of shell 15; of aperture 9, max. diam. 6 mm” (Dall, 1910, p. 133).

Fossil occurrences. Pleistocene of Baja California, Mexico (Jordan, 1924).

Discussion. The name was first used in the northeastern Pacific by Dall (1910, 1921). Grant and Gale (1931) suggested it might be a deeper water form of Olivella baetica [= Callianax alectona]. Burch and Burch (1959) state “A large set is in the Burch collection ex. the late Dr. Fred Baker and labelled, San Diego Bay, topotypes.” This set ranges from very small to Figure 20. Callianax alectona (Duclos, 1835). USNM syntype very large, near 30 mm. Two other sets from Dr. 209677, syntype of Olivella porteri Dall, 1910. Sandbar at the entrance to San Diego Harbor, San Diego County, California. Baker are labelled “...off South Coronado Island, Height 14.8 mm. Photograph by J.H. McLean, provided by Mexico, in 10 fathoms and the other in six LACM Malacology. fathoms. These make a remarkable series ranging from the semi-microscopic to the adult. Type locality. “Sandbars near the entrance to It is possible to select a series from these lots San Diego Harbor…” (Dall, 1910). that would match Dall’s description and be recognizable distinct, but the obvious presence Original description. “Shell of about the form of intergrades to the more typical [C. alectona] and size of the east American O. jaspidea, with indicates that the proper place for porteri is in very variable coloration; whorls about seven, synonymy.” He later cites a note by George spire acute with a very deep and narrow Willett saying “I think porteri is a variant of channeled suture not observed by callus; surface baetica of no taxonomic importance.” Given ISSN 0738-9388

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this and the strong channel at the bottom of the States National Museum, the British Museum, whorls of both C. porteri and C. aectona we and the San Diego Museum. also determine that they are synonymous. Type locality. None given by Berry (1935), however Burch and Burch (1959) gives it as Olivella pycna (Berry, 1935) “Bolinas Bay, California; taken by trawl in 3-4 [= Callianax strigata (Reeve, 1850)] fathoms. F.W. Weymouth, June 22, 1911” (Figure 21) (Berry collection catalogue no. 3316 cited in Burch and Burch, 1959) which is confirmed by Oliva baetica Marrat in Sowerby, 1871, p. 35, labels with the type material. pl. 350, fig. 410. Olivella pycna Berry, 1935, 262-265. Original description. Berry (1935) describes this species as “Shell small, heavy, elongate- nuciform, widest near the middle, the spire tapering almost straightly to an acute point; anterior extremity truncate. Whorls seven, their slopes nearly straight on the spire, the body- whorl large and strongly convex. Suture sharply, narrowly, and deeply channeled. Aperture about three-fifths the length of the shell, the outer lip sharp and distinctly arcuate, its obtuse anterior lobe slightly exceeding the columella anteriorly past the short notch-like canal. Parietal wall covered by a strong white callus, heaviest and of greatest extent posteriorly where it-rounds rather abruptly to pass under the free and overhanging outer lip just in front of the sutural channel, the parietal callus not passing the suture but confluent in the channel with a second low callus just posterior, developed as the termination of a low calloused band which bounds the channel posteriorly and covers about a third of the adjoining whorl before its gradual Figure 21. Olivella pycna Berry, 1935. Holotype CASIZ 64619. and final disappearance. Columellar fold Bolinas Bay, Marin County, California. Height 13.1 mm. Photograph by J.H. McLean, provided by LACM Malacology. moderately heavy, usually distinctly duplex but sometimes single, the columellar and extreme Types. Holotype—CASIZ 64609; Paratypes— anterior region covered by a third white callus CASIZ 62107, 65999. According to Berry which is overlain near the aperture by the (1935) other paratypes are deposited with parietal callus. Surface smooth, lustrous, under Stanford University (now with the California magnification seen to be very finely and closely Academy of Sciences), the Academy of Natural covered with minute wavy spirals on the Sciences of Drexel University, the United uncalloused portions which are again minutely decussated by the numerous fine lines of growth. Parietal callus minutely punctate. Colour light ISSN 0738-9388

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brownish buff (usually quite near Tilleul Buff), Allen, 1945; Touring, 1959; Cummings and clouded brownish, though usually with a rather others, 1962; Addicott and others, 1978; Perry, wide buff spiral band persisting below the 1993) formations and from Pleistocene terrace suture, the whole conspicuously ornamented by deposits at AXo Nuevo State Reserve, central numerous highly irregular vertical wavy stripes California (Addicott, 1966; as O. pedroana). of fawn colour or army brown, with occasional Until fossil specimens from central California suffusion of light vinaceous drab; interior, through Oregon are examined the fossil record except the buff lip-margin, walnut brown to of this species remains incomplete. However, cameo brown; callous portions white as noted.” given the occurrences cited here we determine a probable range of early Pliocene to Holocene Fossil record. The only fossil record of Olivella age range. pycna is Powell (1998) from the Miocene to Pliocene Purisima Formation on the San Discussion. Reported from California by Francisco Peninsula, central California. Kaicher (1987; card OL2-4944), Hemery and However, judging from published pictures and others (2017), on the Malacolog Version 4.1.1 specimens examined by the senior author in web site (http://www.malacolog.org/search.php? collections at CASIZ, LACMIP, UCMP, C. nameid=5424) and the WMSDB Worldwide strigata occurs in the Careaga Sandstone Mollusc Species Database (http://www. (Woodring and Bramlette, 1950[1951]), bagniliggia.it/WMSD/PDFFamily/OLIVELLID Carlotta(?), Elk River, Moonstone Beach, (Roth, AE.pdf, visited 6/2018). 1979), Purisima (Arnold, 1908; Martin, 1916;

1A Spire with large to moderate channeled at base of whorls 2 1B Spire with minimal to no channel at base of whorls Callianax diegensis 2A Spire high with both siphonal plait, and siphonal fasciole plait 3 2B Spire short, shell wide compared to height Callianax biplicata 2C Spire high, no siphonal or siphonal fasciole plaits Olivella pedroana 3A Shell wide compared to height, long relatively narrow siphonal Callianax strigata fasciole with obscure plait. This species also shows very fine, faint, radical sculpture 3B Shell narrower compared to height, long relatively narrow siphonal Callianax alectona fasciole with strong plait and strong plate in siphonal callus

Table 1. Key to distinguishing large, adult California Callianax, living and fossil. Many of these features are variable in juvenile specimens. ISSN 0738-9388

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DISCUSSION single plait, and commonly the posterior end of the plait is narrowly rounded. This species Four species of modern California Callianax are occurs commonly on sand between Gulf of recognized based on shell characteristics. These Alaska south and Cabo San Lucas, Baja are C. alectona (= Olivella. baetica, = O. California Sur, Mexico and around into the Gulf porteri), C. biplicata, C. diegensis, and C. of California to La Paz, at water depths from the strigata (= C. pycna). Olivella pedroana intertidal zone to 65 m. It is commonly brought commonly reported living off California into the rocky intertidal by hermit crabs appears to be strictly a fossil species only occupying the shell. known from the Pleistocene. Olivella dama and O. intorta are valid species but are not known Callianax diegensis can attain a height of 20 from the modern California malacological fauna mm and is similar to C. alectona. However, C. occurring further south in Mexico. Olivella nota diegensis has minimal to no channels on its is a valid western Pacific species also not whorls which are more rounded than other known from California. A key for California Callianax, and the high-spire is distinguishing the four California Callianax commonly topped by a bluish-purple to purple species and Olivella pedroana is presented in protoconch and nuclear whorls. Color is draber Table 1 and, in part, illustrated in Figures 22 than C. alectona and without a pattern. The and 23. siphonal callus is not well offset from the parietal callus and has a moderately broad The first California Callianax, C. alectona raised area on its anterior side. The siphonal (formerly Olivella baetica) is narrow compared fasciole plait is wide at both ends with a single to its height with moderate to strong channels at plait low on the upper margin. The distribution the base of its whorls, reaches a maximum of this species is not well documented as it can height of about 23 mm, and commonly has a be difficult to distinguish from C. alectona, white protoconch. Color can vary but this however it appears to be restricted to the species commonly has a pattern of wavy brown Southern California Bight between Santa lines on a cream background. The siphonal Barbara County, California, and northern Baja callus, broader towards the posterior, and California, Mexico. siphonal fasciole both have subdued plaits. This species occurs from Prince William Sound, The smallest of the California Callianax, C. Alaska, south to Estero Beach, Ensenada, Baja strigata, attains a maximum height of about 16 California at water depths between the intertidal mm with a moderately high, and a strongly zone and 150 m. channeled spire, similar to C. alectona. It is slightly wider, compared to its length, than C. The largest and easiest species to identify of the alectona, but less so than C. biplicata. The California Callianax, C. biplicata, is also the siphonal callus is small and set off from the largest and widest compared to height, comes in siphonal plait at its upper margin. The siphonal a wide variety of colors all without a pattern. It fasciole is smaller than in C. alectona with a attains a maximum height of about 30 mm with strong plait in the middle of its upper margin. It a short to very short spire. It has a single also differs in having very fine, subdued radial siphonal plait on the siphonal fasciole which is lines. This species occurs from Banks Island, offset from the parietal callus. In addition, the Skeena-Queen Charlotte C, British Columbia, siphonal fasciole is pointed at both ends, has a ISSN 0738-9388

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Canada to Morro Bay, San Luis Obispo County, modern specimens with these charactertics was California at depths of 27-46 m. observed. It is similar to C. alectona in having a moderate to strong channels on the bottom of its Olivella pedroana (which may prove to be a whorls but is smaller attaining a maximum Callianax species) is considered here to be height of about 15 mm. The occurrences of this strictly a fossil species distinguished by its large, presumably extinct species is beyond the scope half-moon shaped siphonal fasciole and small of this paper. siphonal callus, both without plaits as no

Figure 22. Comparison of California Callianax (not to size). A = Callianax biplicata (28.3 mm), B = Callianax strigata (15.1 mm) (LACM 1973-45.6), C = Callianax alectona (15.6 mm) (LACM 1968-173.11), D = Callianax diegensis (11.3 mm) (LACM 158258), E = Olivella intorta (10.4 mm). ISSN 0738-9388

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Figure 23. Comparison of the lower half of the apercular side of various northeast Pacific Callianax. A = Callianax biplicata, B = Callianax strigata (LACM 1973-45.6), C = Callianax alectona (LACM 1968-173.11), D = Callianax diegensis (LACM 158258), E = Olivella intorta. Not to scale. The same specimens illustrated in Figure 22 are illustrated here. ISSN 0738-9388

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ACKNOWLEDGEMENTS Addicott, W.O., R.Z. Poore, J.A. Barron, H.D. Gower, and K. McDougall. 1978. We thank Patrick LaFollette, Pierre Recourt, Neogene biostratigraphy of the Indian Creek- and Ron Voskuil for helpful discussions on Shell Creek area, northern La Panza Range, California Callianax. Bob Abela is thanked for California. In Addicott, W.O., ed., Neogene providing photographs of Callianax strigata, biostratigraphy of selected areas in the Kevin Webb (NHMUK) and Andreia Salvador California Coast Ranges. U.S. Geological (NHMUK) are thanked for providing Survey Open-File Report 78-446, p. 49-82. photographs of the syntypes of Olivella Addicott W.O. and J.G. Vedder. 1963. glandinaria and Oliva strigata. Christine Garcia Paleotemperature from late Miocene mollusks in the San Luis Obispo Bakersfield area (CASIZ) is thanked for photographs of several California. U.S. Geological Survey Professional of the Oldroyd types. Lindsey Groves (LACM), Paper 475C, p. C337-C339. Mary McGann (USGS), Edward J. Petuch Allen, J.E. 1945. Geology of the San Juan (Florida Atlantic University), and Pierre Bautista Quadrangle, California. Berkeley, CA. Recourt (MNHN) are thanked for their helpful University of California, Ph.D. dissertation, 94 reviews. Cheryl Millard is also thanked for her pp. help with editing. Anderson, F.M., and B. Martin. 1914. Neogene records in the Temblor basin, REFERENCES California, and Neogene deposits of the San Juan district, San Luis Obispo County. Abbott, R.T. 1951. New deep-water Olivellas Proceedings of the California Academy of from Florida, with notes on the O. jaspidea- Sciences, 4th series, 4(1), p. 14-112. nivea complex. The Nautilus 64(4), p. 110-116. Arnold, R. 1903. The paleontology and Adams, H. and A. Adams. 1853. The genera of stratigraphy of the marine Pliocene and recent Mollusca; arranged according to their Pleistocene of San Pedro, California. Memoir organization. London, J. van Voorst. 484 pp. of the California Academy of Sciences 3, 419 Addicott, W.O. 1959. Late Pleistocene pp. invertebrates from Punta Cabras, Baja Arnold, R. 1906. The Tertiary and Quaternary California, Mexico. American Museum Pectens of California. U.S. Geological Survey Novitates 1925, 33 pp. Professional Paper 47, 264 pp. Addicott, W.O. 1966. Late Pleistocene marine Arnold, R. 1907. New and characteristic paleoecology and zoogeography of central species of fossil mollusks from the oil-bearing California. U.S. Geological Survey Professional Tertiary formations of southern California. Paper 523C, 27 pp. Proceedings of the U.S. National Museum Addicott, W.O. 1969. Late Pliocene mollusks 32(1545), p. 525-546. from San Francisco Peninsula, California, and Arnold, R. 1908. New and characteristic their paleogeographic significance. Proceedings species of fossil mollusks from the oil-bearing of the California Academy of Sciences, 4th Tertiary formations of southern California. series, 37(3), p. 57-93. Proceedings of the U.S. National Museum Addicott, W.O. 1970. Miocene gastropods and 32(1545), p. 525-546. biostratigraphy of the Kern River area, Arnold, R., and H. Hannibal. 1913. The California. U.S. Geological Survey Professional marine Tertiary stratigraphy of the north Pacific Paper 642, 174 pp. coast of America. Proceedings of the American Philosophical Society 52(212), p. 559-605. ISSN 0738-9388

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report. Appendix. Article II. Washington. U. S. DeLong, J.H. 1941. The paleontology and 33d Congress, 2nd session, Ex. Doc. 78; House stratigraphy of the Pleistocene at Signal Hill, Ex. Doc. 91, p. 327, pl. 6, fig. 51. Long Beach, California. Transactions of the Cooper, J.G. 1888. Catalogue of California San Diego Society of Natural History 9(25), p. fossils, parts 2-5. California State 229-250. Mineralogists, 7th annual report. Pp. 21-308. Dickerson, R.E. 1922. Tertiary and Quaternary Cooper, J.G. 1894. Catalogue of west North history of the Petaluma, Point Reyes, and Santa American and many foreign shells, with their Rosa quadrangles. Proceedings of the geographic ranges. For labels, exchange and California Academy of Sciences, 4th Series checklists, with a supplement. Sacramento, 11(19), p. 527-601. Calif. State Office, A.J. Johnson, supt. State Duclos, P.L. 1835. Genre Olive. In. Histoire Printing. naturelle générale et particulière de tous les Cummings, J.C., R.M. Touring, and E.E. genres de coquilles univalves marines à l'état Brabb. 1962. Geology of the northern Santa vivant et fossile, publiée par monographie. [4] Cruz Mountains, California. In Bowen, O.E., Jr., plates 1-12. Paris. ed., Geologic guide to the gas and oil fields of Edwards, D.C. 1968. Reproduction in Olivella northern California. Bulletin of the California biplicata. The Veliger 10(4), p. 297-304. Division of Mines and Geology 181, p. 179-220. Eerkens, J.W., G.S. Herbert, J.S. Rosenthal, Cuvier, G. 1795. Second Mémoire sur and H.J. Spero. 2005. Provenance analysis of l'organisation et les rapports des animaux à Olivella biplicata shell beads from the sang blanc, dans lequel on traite de la structure California and Oregon coasts by stable isotope des Mollusques et de leur division en ordre, lu à fingerprinting. Journal of Archaeological la société d'Histoire Naturelle de Paris, le 11 Sciences 32, p. 1501-1514. prairial an troisième [30 May 1795]. Magazin Eerkens, J.W., J.S. Rosenthal, H.J. Spero, Encyclopédique, ou Journal des Sciences, des N.E. Stevens, R. Fitzgerald, and L. Brink. Lettres et des Arts, 1795 [1. année] 2, p. 433- 2009. The source of early horizon Olivella 449. beads. isotopic evidence from CCO-548. SCA Dall, W.H. 1878. Fossil mollusks from later Proceedings 23, 11 pp. Tertiaries of California. Proceedings of the U.S. Ellis, A.J., and C.H. Lee. 1919. Geology and National Museum 1(8), p. 10-16. ground waters of the western part of San Diego Dall, W.H. 1909. Contributions to the Tertiary County, California. U.S. Geological Survey paleontology of the Pacific Coast, I. The Water-Supply Paper 446, 321 pp. Miocene of Astoria and Coos Bay, Oregon. U.S. Emerson, W.K., and E.P. Chace. 1959. Geological Survey Professional Paper 59, 278 Pleistocene mollusks from Tecolote Creek, San pp. Diego, California. Transactions of the San Dall, W.H. 1910. New species of west Diego Society of Natural History 7(21), p. 335- American shells. The Nautilus 23(11), p. 133- 346. 134. English, W.A. 1914. The Fernando Group near Dall, W.H. 1921. Summary of the marine shell Newhall, California. University of California bearing mollusks of the northwest coast of Publications, Department of Geology Bulletin America, from San Diego, California to the 8(8), p. 203-218. Polar Sea, mostly contained in the collections Gamble, L.H., and C.D. King. 2011. Beads of the United States National Museum, with and ornaments from San Diego. Evidence for illustrations of hitherto unfigured species. U.S. exchange networks in southern California and National Museum Bulletin 112, 217 pp. the American southwest. California and Great ISSN 0738-9388

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Basin Journal of Anthropology 31(2), p. 155- Publications, Department of Geological 178. Sciences Bulletin 14(3), p. 85-114. Gardner, J. 1948. Mollusca from the Miocene Jordan, E.K. 1924. Quaternary and recent and lower Pliocene of Virginia and North molluscan faunas of the west coast of Lower Carolina. Part 2. Scaphopoda and Gastropoda. California. Bulletin of the Southern California U.S. Geological Survey Professional Paper 199 Academy of Sciences 23(5), p. 145-156. B, p. 179-279. Jordan, E.K. 1926. Expedition to Guadalupe Golikov, A.N., and O.A. Scarlato. 1967. Island, Mexico, in 1922. No. 4. Molluscan Моллюски залива Посьет (Японское море) и faunas of the Pleistocene of San Quintin Bay, их экология. Моллюски и их роль в Lower California. Proceedings of the California биоценозах и формировании фаун. [Molluscs Academy of Sciences, 4th series 15(7), p. 241- of the Possjet Bay (the Sea of Japan) and their 255. ecology. Molluscs and their role in biocenoses Kaicher, S.D, 1987. Card Catalogue of World- and formation of faunas]. Trudy wide Shells Pack #49—Olividae Part II. S.D. Zoologicheskogo Instituta Leningrad 42, p. 5- Kaicher, St. Petersburg, FL. Cards [i-ii], 4942- 154. 5046. Gould, A.A., and P.P. Carpenter. 1856. Kantor, Y.I., A.E. Fedosov, N. Puillandre, C. Descriptions of shells from the Gulf of Bonillo, and P. Bouchet. 2017. Returning to California and the Pacific Coast of Mexico and the roots: morphology, molec-ular phylogeny California, part 2. Proceedings of the and classification of the Olivoidea (Gastropoda: Zoological Society of London 24, p. 198-281. Neogastropoda). — Zoological Journal of the Grant, U.S., IV, and H.R. Gale. 1931. Linnean Society, 2017 180: 493-541. Catalogue of the marine Pliocene and Keen, A.M. 1945. New mollusks from the Pleistocene Mollusca of California. Memoir of Round Mountain Silt (Temblor) Miocene of the San Diego Society of Natural History 1, California. Transactions of the San Diego 1,036 pp. Society of Natural History 10(2), p. 25-60. Hartzell, L.L. 1991. Archaeological evidence Keen, A.M. 1971. Sea Shells of Tropical West for stages of manufacture of Olivella shell America. Marine Mollusks from Baja beads in California. Journal of California and California to Peru. Second edition. Stanford, Great Basin Archaeology 13(1), p. 29-39. Calif. Stanford University Press. 1,064 pp. Hemery, L.G., K.K. Politano, and S.K. Keep, J. 1887. West Coast Shells. A familiar Henkel. 2017. Assessing differences in description of the marine, freshwater, and land macrofaunal assemblages as a factor of sieve mollusks of the United States found west of the mesh size, distance between samples, and time Rocky Mountains. San Francisco, Calif. of sampling. Environ Monit Assess 189, p. 413- Bancroft, 230 pp. 431, doi.10.1007/s10661-017-6127-8. Latreille, P.A. 1825. Familles naturelles du Hickman, C.S., and J.H. Lipps. 1983. règne animal, exposée succinctement et dans un Foraminiferivory: selective ingestion of ordre analytique, avec l’indication de leurs foraminifera and test alterations produced by genres. Paris. J.B. Bailliére. 570 pp. the neogastropod Olivella. Journal of Link, D.H.F. 1807-1808. Beschreibung der Foraminiferal Research 13(2), p. 108-114. Naturalien-Sammlung der Universität zu Howe, H.V. 1922. Faunal and stratigraphic Rostock. Rostock, Adlers Erben. 1 Abt. [Part 1], relationships of the Empire Formation, Coos p. 1-50; 2 Abt. [Part 2], p. 51-100; 3 Abt. [Part Bay, Oregon. University of California 3], p. 101-165; Abt. 4 [Part 4], 30 pp; Abt. 5 [Part 5], 38 pp [1808]; Abt. 6 [Part 6], 38 pp. ISSN 0738-9388

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Linnaeus, C. 1758. Systema Naturae per regna Oldroyd, T.S. 1925. The fossils of the lower tria naturae, secundum classes, ordines, genera, San Pedro fauna at Nob Hill cut, San Pedro, species, cum characteribus, differentiis, California. Proceedings of the U.S. National synonymous, locis. Editio decima, reformata. Museum 65(2535), 36 pp. Laurentius Salvius. Holmiae. ii, 824 pp., Onuf, C.P. 1972. Aspects of the population http://www.biodiversitylibrary.org/ biology of the intertidal snail Olivella biplicata; item/10277#page/3/mode/1up. distribution, nutrition and effects of natural Loel, W. and W.H. Corey. 1932. The enemies. Santa Barbara, Calif. University of Vaqueros Formation, lower Miocene of California at Santa Santa Barbara, unpublished California. I Paleontology. University of Ph.D. dissertation, 223 pp. California Publications, Department of Osmont, V.C. 1905. Geological section of the Geological Sciences Bulletin 22(3), p. 31-410. Coast Ranges north of the Bay of San Francisco. Marrat, F.P. 1868. On some new species of University of California Publications, Bulletin Oliva. Annals and Magazine of Natural History, of the Department of Geology 4(3), p. 39-87. 4th series, 2, p. 212-214. Packard, E.L. 1918. Molluscan fauna from San Marrat, F.P. 1870-1871. Monograph of the Francisco Bay. University of California genus Oliva. In G.B. Sowerby II (ed.), Publication in Zoology 14(2), p. 199-452. Thesaurus Conchyliorum, vol. 4 (29-30), p. 1- Palmer, K.V.W. 1958. Type specimens of 46 [1871], pl. 328–341 [1870], p. 342-351 marine Mollusca described by P.P. Carpenter [1871]. London, privately published. from the West Coast (San Diego to British Martin, B. 1916. The Pliocene of middle and Columbia). Geological Society of America northern California. University of California Memoir 76, p. 376 pp. Publications, Department of Geology Bulletin Palmer, K.V., and D.C. Brann. 1965. 9(15), p. 215-259. Catalogue of the Paleocene and Eocene McLean, J.H. 2007. Shelled Gastropoda. In Mollusca of the southern and eastern United Carlton, J.T., ed., The Light and Smith Manual. States. Part 1. Pelecypoda, Amphineura, Intertidal Invertebrates from Central California Peteropoda, Scaphopoda and Cephalopoda. to Oregon (4th ed.), Berkeley, Calif., University Bulletins of American Paleontology 48, 471 pp. of California Press, p. 713–753. Perry, F.A. 1993. Fossil invertebrates and Nomland, J.O. 1917. The Etchegoin Pliocene geology of the marine cliffs at Capitola, of middle California. University of California California. Santa Cruz, CA. Santa Cruz City Publications, Department of Geology Bulletin Museum, 30 pp. 10(14), p. 191-254. Powell, C.L., II. 1998. The Purisima Formation Oldroyd, I.S. 1924-1927. The marine shells of and related rocks (upper Miocene - Pliocene), the west coast of North America. Stanford, CA. greater San Francisco Bay area, central Stanford University Press. Pelecypoda and California. Review of literature and USGS Brachiopoda 1, 247 pp., 57 pls. (1924); collections (now housed at the Museum of Scaphopoda and gastropods II(1), 297 pp., 29 Paleontology, University of California, pls. (1927); II(2), p. 304 pp., pls. 30-72 (1927); Berkeley). U. S. Geological Survey Open-File II(3), p. 339 pp., pls. 73-108 (1927). Report 98-594, 102 pp., http://wrgis.wr. Oldroyd, T.S. 1918. Olivella biplicata usgs.gov/open-file/of98-594/. angelena, var. nov. The Nautilus 32(1), p. 34- Powell, C.L., II. 2008. Outcrops and mollusks 35. of the “Margaritan” California provincial Oldroyd, T.S. 1921. Some varieties of western molluscan stage in the northern Salinas Valley, Olivellas. The Nautilus 34(4), p. 117-119. ISSN 0738-9388

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Monterey and San Benito counties, central Sowerby, G.B. 1847-1887 (1880). Thesaurus California. PaleoBios 27(3), p. 86-125. conchyliorum, or, Monographs of genera of Powell, C.L., II, L.B. Grant, and S.W. shells, edited by G.B. Sowerby, Jr., completed Conkling. 2005. Paleoecologic analysis and by G.B. Sowerby, III. London, 5 volumes, age of a Late Pleistocene fossil assemblage issued in 44 parts. from Upper Newport Bay, Newport Beach, Squires, R.L. 2012. Late Pliocene megafossils Orange County, California. The Veliger 47(3), of the Pico Formation, Newhall area, Los p. 183-192. Angeles County, southern California. Natural Powell, C.L., II, and D. Stevens. 2000. Age History Museum of Los Angeles County, and paleoenvironmental significance of mega- Contributions in Science 520, p. 73-93. invertebrates from the "San Pedro" Formation Stevens, F. 1929. Notes on the marine in the Coyote Hills, Fullerton and Buena Park, Pleistocene deposits of San Diego County, Orange County, Southern California. U.S. California. Transactions of the San Diego Geological Survey Open-File Report 319, 83 pp. Society of Natural History 5(16), p. 247-255. Puri, H.S., and R.O. Vernon. 1964. Summary Swainson, W. 1829-1833. Zoological of the geology of Florida and a guidebook to Illustrations, or original figures and the classic exposures. Florida Geological descriptions of new, rare, or interesting animals, Survey Special Publication 5, 312 pp. selected chiefly from the classes of ornithology, Reeve, L.A, 1850. Monograph of the genus entomology, and conchology, and arranged Oliva. Conchologia Iconica, 6, pls. 1-30. according to their apparent affinities. Second Richards, H.G. and K.V.W. Palmer. 1953. series. Baldwin & Cradock, London. v. 1-3, pl. Eocene Mollusks from Citrus and Levy 1-30 (1829), pl. 31-45 (1830), pl. 46-85 (1831), Counties, Florida. Florida Geological Survey pl. 86-96 (1832), pl. 97-136 (1833), Geological Bulletin 35, 95 pp. http://www.biodiversitylibrary.org/bibliography Rosenberg, G., and R.E. Petit. 2003. /42278. Kaicher’s Card Catalogue of World-Wide Touring, R.M. 1959. Structure and stratigraphy Shells. A collation with discussion of species of the La Honda and San Gregorio Quadrangles, named therein. The Nautilus 117(4), p. 99-120. San Mateo County, California. Stanford Roth, B. 1979. Late Cenozoic marine University, California. Ph.D. dissertation, 228 invertebrates from northwest California and pp. southwest Oregon. Berkeley, CA. University of Troschel, F.H. 1866-1893. Das Gebiss der California, Ph.D. dissertation, 792 pp. Schnecken zur Begründung Einer Natürlichen Russell, M.P. 1991. Modern death assemblages Classification. Nicolaische Verlags- and Pleistocene fossil assemblages in open Buchhandlung, Berlin. 2(1), p. 1-64, pls. 1-4 coast high energy environments, San Nicolas (1866); 2, p. 65-96, pls. 5-8 (1868); 3, p. 97- Island, California. Palaios 6(2), p. 179-191. 132, pls. 9-12 (1869); 4, p. 133-180, pl. 13-16 Smith, A.G., and Gordon, M., Jr. 1948. The (1875); 5, p. 181-216, pls. 17-20 (1876); 6, p. marine mollusks and brachiopods of Monterey 217-240, pls. 21-24 (1879); 7, p. 251-334, pls. Bay, California, and vicinity. Proceedings of 25-28 (1891); 8, p. 337-409, pls. 29-32 (1893). the California Academy of Sciences, 4th series, Valentine, J.W. 1956. Upper Pleistocene 26(8), p. 147-245. Mollusca from Potrero Canyon, Pacific Sowerby, G.B., I. 1825. A catalogue of the Palisades, California. Transactions of the San shells contained in the collections of the late Diego Society of Natural History 12(10), p. Earl of Tankerville. E.J. Stirling for G.B. 181-205. Sowerby, London. 92 pp. ISSN 0738-9388

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Valentine, J.W. 1957. Late Pleistocene faunas Wenz, W. 1938-1944. Handbuch der from the northwestern coast of Baja California, Paläozoologie. Berlin, Gebrüder Borntraeger, Mexico. Transactions of the San Diego Society 8vo., ed. 6. Gastropoda. Teil 1, I-viii240 pp., of Natural History 12(16), p. 298-308. Allgemeiner Teil und Prosobranchia (March Valentine, J.W. 1961. Paleoecologic molluscan 1938); teil 2, p. 241-480, Prosobranchia und geography of the California Pleistocene. Mesogastropoda (1938); teil 3, p. 481-720, University of California Publications in Prosobranchia (July 1939); teil 4, p. 720-960, Geological Sciences 34(7), p. 309-442. Prosobranchia (August 1940); teil 5, p. 961- Valentine, J.W. 1966. Numerical analysis of 1200, Prosobranchia (October 1941); teil 6, p. marine molluscan ranges on the extratropical 1201-1506, Prosobranchia (October 1943); teil northeastern Pacific shelf. Limnology and 7, p. 1507-1639, Inhalt, literature, index Oceanography 11(2), p. 198-211. (November 1944). Vanatta, E.G. 1915. Notes on Oliva. The Williamson, M.B. 1892. An annotated list of Nautilus 29(6), p. 67-75. the shells of San Pedro Bay and vicinity. U.S. Vedder, J.G., and R.M. Norris. 1963. Geology National Museum Proceedings 15(898), p. 179- of San Nicolas Island, California. U.S. 220. Geological Survey Professional Paper 369, 65 Wood, W. 1828. Supplement to the Index pp. Testaceologicus; or a catalogue of shells, Vervaet, F.L.J. 2018.The living Olividae British and Foreign. Illustrated with 480 figures. species described by Pierre-Louis Duclos. Vita Richard Taylor for W. Wood, London. 59 pp. Malacologica 17, 111 pp. Woodring, W.P., and M.N. Bramlette. 1950 Walker, S.E. 1992. Criteria for recognizing [1951]. Geology and paleontology of the Santa marine hermit crabs in the fossil record using Maria District, California. U.S. Geological gastropod shells. Journal of Paleontology 66(4), Survey Professional Paper 222, 185 pp. p. 535-558. Woodring, W.P., M.N. Bramlette, and S.W. Weaver, C.E. 1912. A preliminary report on Kew. 1946. Geology and paleontology of Palos the Tertiary paleontology of western Verdes Hills, California. U.S. Geological Washington. Washington Geological Survey Survey Professional Paper 207, 145 pp. Bulletin 15, 80 pp. Yates, L.G. 1886. Catalogue of marine shells in the Lorenzo G. Yates’ collection, Santa Barbara, Cal. Privately printed, Santa Barbara, 81 pp. ISSN 0738-9388

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Plate 1. A-F. Callianax strigata. A-C. LACM 1973-45.6 Esperanza Inlet, Vancouver Island, British Columbia, Canada: A = 15.1 mm, B = 15.0 mm, C = 15.4 mm. D-F. LACM 1949-20.11 outer Bodega Bay, Sonoma County, California. D = 15.5 mm, E = 14.0 mm, F = 11.3 mm. G-K. Callianax biplicata. G = Santa Monica Bay, Los Angeles County, California, 27.5 mm (Berschauer Collection). H-K = Laguna Beach, Orange County, California (Berschauer Collection); H = 22,2 mm, I = 25.1 mm, J = 28.2 mm, K = 21.6 mm. ISSN 0738-9388

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Plate 2. A-F. Callianax alectona. A = LACM 1968-174.8, Vancouver Island, British Columbia, Canada, 26.4 mm. B-C = LACM 1939-129.25, Pyramid Cove, San Clemente Island, Los Angeles County, California; B = 19.7 mm, and C = 18.3 mm. D-F = LACM 1968-173.11, Vancouver Island, British Columbia, Canada; D = 21.5 mm, E = 15.6 mm, F = 20.0 mm. G-L. Callianax diegensis. G-I = LACM 158258, Alamitos Bay, Los Angeles County, California; G = 15.0 mm, H = 13.6 mm, I = 13.8 mm. J-L = San Pedro, Los Angeles County, California (Berschauer Collection); J = 14.8 mm, K = 16.6 mm, L = 14.6 mm. Callianax strigata (Reeve, 1850) photographed by Bob Abela. This 11.0 mm specimen was collected while diving of Bodega Head, Sonoma County, California in about 30 ft on clean, ﬁne sand along the exposed side of the breakwater. 01 September 1991.

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