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Type specimens of fossil “Architectibranchia” and Cephalaspidea (Mollusca, Heterobranchia) in the Academy of Natural Sciences of Philadelphia
expand article infoCarlo M. Cunha§|, Rodrigo B. Salvador#
‡ Drexel University, Philadelphia, United States of America
§ Ministry of Education of Brazil, Brasília, Brazil
| Universidade Metropolitana de Santos, Santos, Brazil
¶ Museum of New Zealand Te Papa Tongarewa, Wellington, New Zealand
# Staatliches Museum für Naturkunde Stuttgart, Stuttgart, Germany
Open Access

Abstract

The type specimens of fossil “Architectibranchia” and Cephalaspidea (Mollusca: Heterobranchia) deposited in the Academy of Natural Sciences of Philadelphia, USA, are listed herein. The collection includes types of circa 60 species, from the families: Acteonellidae, Acteonidae, Bullidae, Cylichnidae, Haminoeidae, Philinidae, Retusidae, Rhizoridae, Ringiculidae and Scaphandridae. The catalogue is presented in systematic order, with information on the original description, type locality, type stratum and age, catalog number in the collection, and current taxonomic status. Further taxonomic notes are offered when pertinent. Several species are illustrated here for first time. The new combinations Roxania hornii (Gabb, 1864) and Volvulella minutissima (Gabb, 1860) are proposed. Some species that have previously been assigned to Acteonidae are revised here, resulting in the following new combinations: Odostomia milium (Lea, 1846), Chrysallida sculpta (Lea, 1846) and Pyrgulina angulata (Lea, 1846). The list of taxa is also presented in other arrangements (alphabetically by specific epithets and by authorship and date) to facilitate locating information.

Key Words

Cenozoic, Cretaceous, type specimens

Introduction

The collection of fossil mollusks in the Academy of Natural Sciences of Philadelphia (ANSP; Philadelphia, PA, USA) houses about 80,000 lots, mostly of gastropods. Despite containing fossils from diverse localities worldwide, most stems from the Mesozoic and Cenozoic eras of the USA. The collection counts with original material (including type specimens) of several 19th and early 20th century paleontologists, such as Timothy A. Conrad, William M. Gabb, Henry C. Lea, Isaac Lea and Henry A. Pilsbry.

It is an international consensus that all museums should publish inventories of their type specimens. Since many invertebrate species have convoluted taxonomic histories often with inadequate descriptions and illustrations (or no illustration at all), and little modern taxonomic analysis, it is deemed that such catalogues will benefit or prompt future studies.

Some catalogues have been published regarding the Recent mollusks of the ANSP collection, focusing on specific authors or families (Borrero and Rosenberg 2015, Callomon 2015, Snyder and Callomon 2015), and the entire collection is presently searchable online. A catalogue of the ANSP invertebrate fossil types was published by Richards (1968), as well as more specific ones dealing with type specimens of a single author (e.g., Moore 1962, for Conrad’s types) or a restricted period (e.g., Johnson 1905, for the Cretaceous). These catalogues sometimes fail to cite the type specimens of some species, might present conflicting information, and, due to their broad scope, offer very little extra information on the species’ taxonomy. Therefore, we present here an annotated catalogue of the fossil “Architectibranchia” and Cephalaspidea gastropod types housed in the ANSP Invertebrate Paleontology collection. When necessary, new combinations are proposed. Moreover, some species that have previously been assigned to Acteonidae are revised, resulting in new combinations.

Material and methods

The present catalogue offers information on the original description of each species, the ANSP catalog number for the type material lots, type locality and stratum, age and current taxonomical status (following the most recent revisions, where they exist). At least one type specimen (holotype, lectotype or syntype) of each nominal species is figured here, with further type specimens figured only when they add information. Several species are figured here for the first time.

The type catalogue is arranged in three ways to facilitate locating information: (1) by current systematic position (with additional information and comments and, when necessary, proposed new combinations); (2) alphabetically by specific epithet; (3) by authorship and date of the nominal species.

Some species whose types are housed in the ANSP collection were originally classified in Architectibranchia or Cephalaspidea, but clearly do not belong to them. They are listed further below and, as some of them have never received a taxonomical reassessment, they are reclassified here.

The classification used here follows Bouchet et al. (2005, 2017), complemented (when available) by taxonomic revisions that deal specifically with the fossil taxa. The taxa represented in the ANSP collection are: Architectibranchia (Acteonellidae, Acteonidae, Ringiculidae) and Cephalaspidea (Bullidae, Cylichnidae, Haminoeidae, Philinidae, Retusidae, Rhizoridae, Scaphandridae).

Type localities and strata are provided as precisely as possible and when the original descriptions were not very precise, we added information from previous authors, specimens labels, or our own research (new information is always clearly indicated as such). Moreover, some locality and formation names were updated to conform to current conventions; the original names are also indicated. Likewise, the age of the strata are given with as much resolution as possible. However, several localities have not been studied in detail since then; in these cases, a coarser age span (e.g., period) is indicated.

Previous catalogues and species lists (e.g., Palmer 1937, Moore 1962, Richards 1968) often refer to the type specimens in different manners, without discussing the reasoning behind their choice. Herein, we indicated these previous assesments in quotation marks. One issue can be generalized here, though, regarding the use of the word “holotype” by previous catalogues when confronted by a single specimen in the collection. This practice is erroneous and all the original specimens are considered syntypes (even if there is only a single one) herein, unless explicetely indicated on the original description that a single specimen was available (in which case, it is a holotype).

Unfortunately, some of the types supposedly housed in the ANSP collection could not be found in the present study and are thus considered lost. They are: Acteon costellatus Conrad, 1833; Acteon modicellus Conrad, 1860; Bulla mortoni Forbes, 1845 and Retusa sulcata fossilis Pilsbry, 1922.

Systematic list of taxa

List of taxa by systematic arrangement

Heterobranchia

Superfamily Acteonoidea d’Orbigny, 1843

Family Acteonidae d’Orbigny, 1843
Genus Acteocina Gray, 1847

Acteocina cederstromi Richards, 1947

Figure 1A–B

Acteocina cederstromi Richards, 1947: 34, pl. 11, fig. 9.

Type locality

Bacons Castle, Virginia, depth 115 ft. (ca. 35 m); stratum: Yorktown Formation; age: Late Miocene to Middle Pliocene.

Type material

Holotype, ANSP IP16771 (as “type” in Richards 1968: 113).

Current taxonomic status

Acteocina candei (d’Orbigny, 1841) (Campbell 1993).

Figure 1. 

Types. A–B. Holotype of Acteocina cederstromi, H 4.0 mm, ANSP IP16771. C–D. Holotype of Acteocina chowanensis, H 3.8 mm, ANSP IP16754. E–F. Lectotype of Acteocina crassiplica, H 6.0 mm, ANSP IP13412. G–H. Paratype of Acteocina kirkwoodiana, H 5.0 mm, ANSP IP15935. I–J. Holotype of Acteocina puruha, H 7.6 mm, ANSP IP13684. K–L. Holotype of Acteocina subbullata, H 9.9 mm, ANSP IP3193. M–N. Syntype of Acteocina weatherlli, H 4.6 mm, ANSP IP14431. O–P. Syntype of Acteon biplicatus, H 17.2 mm, ANSP IP19466; also, syntype of A. gabbana. Q–R. Syntype of Acteon cretacea, H 19.1 mm, ANSP IP18778. S. Holotype of Acteon elegans, H 6.0 mm, ANSP IP6011. T–U. Syntype of Acteon forbesiana, H 9.2 mm, ANSP IP18777.

Acteocina chowanensis Richards, 1947

Figure 1C–D

Acteocina chowanensis Richards, 1947: 34, pl. 11. fig. 10.

Type locality

Edenton, well 11 (as “well 3” in original description), depth 46–58 ft. (ca. 14–17.5 m), U. S. Marine Base, North Carolina, USA; stratum: Chowan River Formation; age: Pliocene.

Type material

Holotype, ANSP IP16754 (as “type” in Richards 1968: 113).

Current taxonomic status

Acteocina canaliculata (Say, 1826) (Mikkelsen and Mikkelsen 1984).

Acteocina crassiplica (Conrad, 1848)

Figure 1E–F

Bulla crassiplica Conrad, 1848a: 282.

Type locality

Dr. Smith’s plantation, 6 miles northeast of Vicksburg, Warren County, Mississippi, USA; stratum: Vicksburg Group; age: Oligocene.

Type material

Lectotype, ANSP IP13412 (designation in Moore 1962: 51, Richards 1968: 119, MacNeil and Dockery 1984: 240); paralectotypes, ANSP 13413, 6 shells (as “paratype” in Richards 1968: 119, MacNeil and Dockery 1984: 240).

Current taxonomic status

Acteocina crassiplica (Conrad, 1848) (MacNeil and Dockery 1984).

Acteocina kirkwoodiana Richards & Harbison, 1944

Figure 1G–H

Acteocina kirkwoodiana Richards & Harbison, 1944: 9, figs 1, 5–6.

Type locality

Brandywine Lighthouse well, depth 385 feet [ca. 117 m], Delaware Bay, New Jersey, USA; stratum: Kirkwood Formation; age: Miocene.

Type material

Paratype, ANSP IP15935, 1 shell (Richards and Harbison 1944: 14, fig. 1, 5; as “paratypes” in Richards and Harbison 1944: figs 1, 5; as “cotype” in Richards 1968: 148).

Current taxonomic status

Acteocina kirkwoodiana Richards & Harbison, 1944.

Acteocina puruha Pilsbry & Olsson, 1941

Figure 1I–J

Acteocina puruha Pilsbry & Olsson, 1941: 13, pl. 8, fig. 1.

Type locality

Punta Blanca, Ecuador; stratum: Canoa Formation; age: Late Pliocene.

Type material

Holotype, ANSP IP13684 (as “type” in Richards 1968: 179).

Current taxonomic status

Acteocina puruha Pilsbry & Olsson, 1941.

Acteocina subbullata Pilsbry & Johnson, 1917

Figure 1K–L

Acteocina subbullata Pilsbry & Johnson, 1917: 150–151.

Type locality

Dominican Republic; stratum: “Santo Domingan Beds” (either Cercado or Gurabo Formations); age: Miocene/Pliocene.

Type material

Holotype, ANSP IP3193 (as “type” in Richards 1968: 192).

Current taxonomic status

Acteocina bullata (Kiener, 1834) (Woodring 1970).

Acteocina weatherlli (Lea, 1833)

Figure 1M–N

Acteon weatherlli Lea, 1833: 213, pl. 6, fig. 224.

Type locality

Monmouth Co., Deal, New Jersey, USA; stratum: uncertain; age: Miocene(?).

Type material

Syntype, ANSP IP14431, 1 shell (as “type” in Richards 1968: 205).

Current taxonomic status

Acteocina canaliculata (Say, 1826) (Mikkelsen and Mikkelsen 1984 as A. wetherlli [sic]).

Genus Acteon Montfort, 1810

Acteon biplicatus (Gabb, 1860)

Figure 1O–P

Acteonina biplicata Gabb, 1860a: 93, pl. 2, fig. 13 [non d’Orbigny].

Type locality

—New Jersey, USA; stratum: “Lower Green Marls” (Navesink Formation); age: Cretaceous.

Type material

Syntypes, ANSP IP19466 (as “type” in Richards 1968: 107), 1 shell, ANSP 19467, 1 shell.

Current taxonomic status

Acteon gabbana Whitfield, 1892 (Whitfield 1892, Richards and Ramsdell 1962).

Acteon costellatus Conrad, 1833

Acteon costellatus Conrad, 1833b: 45.

Type locality

Claiborne Bluff, Alabama River, Monroe County, Alabama, USA; stratum: Gosport Sand (uppermost Claiborne Group); age: Eocene.

Type material

Lost (Harris 1895a: 13, Palmer 1937: 501, Moore 1962: 50).

Current taxonomic status

Acteon costellatus Conrad, 1833 is considered a species inquirenda (Salvador and Cunha 2016).

Acteon cretacea Gabb, 1862

Figure 1Q–R

Acteon cretacea Gabb, 1862: 318.

Type locality

Crosswicks, New Jersey, USA; stratum: “Lower Green Marls” (Navesink Formation); age: Cretaceous.

Type material

Syntypes, ANSP IP18778, 2 shells (as “types” in Richards 1968: 120).

Current taxonomic status

Acteon cretacea Gabb, 1862 (Richards and Ramsdell 1962, Richard and Shapiro 1963).

Acteon elegans (Lea, 1833)

Figure 1S

Monoptygma elegans Lea, 1833: 203, pl. 6, fig. 217.

Type locality

Monroe Co., Claiborne Bluff, Alabama, USA; stratum: Gosport Sand (uppermost Claiborne Group); age: Eocene.

Type material

Holotype, ANSP IP6011 (as “lectotype” [error] in Palmer 1937: 499; as “type” in Richards 1968: 128).

Current taxonomic status

Acteon pomilius Conrad, 1833 (Palmer and Brann 1966).

Acteon forbesiana Whitfield, 1892

Figure 1T–U

Actaeon forbesiana Whitfield, 1892: 157, pl. 19, figs 17–22.

Type locality

Walnford, New Jersey, USA; stratum: “Lower Green Marls” (Navesink Formation); age: Cretaceous.

Type material

Syntypes, ANSP IP18777, 4 shells (as “types” in Richards 1968: 113).

Current taxonomic status

Acteon cretacea Gabb, 1862 (Weller 1907, Richards and Ramsdell 1962).

Acteon gabbana Whitfield, 1892

Figure 1O–P

Actaeon gabbana Whitfield, 1892: 156, pl. 19, figs 23–25.

Type locality

Tinton Falls, New Jersey, USA; stratum: “Lower Green Marls” (Navesink Formation); age: Cretaceous.

Type material

Syntypes (also of Acteonina biplicata Gabb, 1860), ANSP 19466 (as “type” in Richards 1968: 135), 1 shell, ANSP IP19467, 1 shell.

Current taxonomic status

Acteon gabbana Whitfield, 1892 (Weller 1907, Richards and Ramsdell 1962), nom. nov. pro Acteonina biplicata Gabb, 1860 [non d’Orbigny].

Acteon glans Lea, 1846

Figure 2A

Acteon glans Lea, 1846: 256, pl. 36, fig. 58.

Type locality

Petersburg, Dinwiddie County, Virginia, USA; stratum: Yorktown Formation; age: Late Miocene to Middle Pliocene.

Type material

Holotype, ANSP IP1517.

Current taxonomic status

Acteon glans Lea, 1846.

Figure 2. 

Types. A. Holotype of Acteon glans, H 1.9 mm, ANSP IP1517. B–C. Syntype of Acteon idoneus, H 11.1 mm, ANSP IP30547. D–E. Lectotype of Acteon lineatus, H 8.6 mm, ANSP IP5541. F. Holotype of Acteon nitens, H 4.0 mm, ANSP IP1516. G–H. Syntype of Acteon novellus, H 12.0 mm, ANSP IP1600. I–L. Syntype of Acteon ovoidea, H 23 mm, ANSP IP56167. 32–33. Same, H 21.0 mm, ANSP IP56166; also, holotype of A. subovoides. M–N. Lectotype of Acteon politus, H 8.2 mm, ANSP IP4266. O–P. Syntype of Acteon pomilius, H 10.1 mm, ANSP IP30546. Q–R. Syntype of Acteon punctatus, H 8.8 mm, ANSP IP5537. S–T. Holotype of Acteon subtornatilis, H 18.1 mm, ANSP IP3183.

Acteon idoneus Conrad, 1833

Figure 2B–C

Acteon idoneus Conrad, 1833b: 45.

Type locality

Claiborne Bluff, Alabama River, Alabama, USA; stratum: Gosport Sand (uppermost Claiborne Group); age: Eocene.

Type material

Syntypes, ANSP IP30547 (as “lectotype” in Palmer 1937: 500; as “lectotype?” in Moore 1962: 65), 1 shell, ANSP IP53814, 5 shells (all as “syntypes” in Richards 1968: 144). The designation as lectotype by Palmer (1937) is likely mistaken (Moore 1962; Richards 1968).

Current taxonomic status

Acteon idoneus Conrad, 1833 (Palmer 1937, Palmer and Brann 1966, Dockery 1977).

Acteon lineatus Lea, 1833

Figure 2D–E

Acteon lineatus Lea, 1833: 112, pl. 4, fig. 97.

Type locality

Claiborne, Alabama, USA; stratum: uncertain [likely Gosport Sand (uppermost Claiborne Group)]; age: Eocene.

Type material

Lectotype, ANSP IP5541, 1 shell (designation by Palmer 1937: 500; as “holotype” in Richards 1968: 152); paralectotypes, ANSP IP5442, 1 shell, ANSP IP5443, 1 shell, ANSP IP5444, 1 shell (each as “paratype” in Richards 1968: 152).

Current taxonomic status

Acteon idoneus Conrad, 1833 (Palmer 1937, Palmer and Brann 1966).

Acteon modicellus Conrad, 1860

Actaeon modicellus Conrad, 1860: 287.

Type locality

Tippah County, Mississippi, USA; stratum: “dark gray marl”; age: Late Cretaceous.

Type material

Lost (Sohl 1964).

Current taxonomic status

Acteon modicellus Conrad, 1860, nomen dubium (Sohl 1964).

Acteon nitens Lea, 1846

Figure 2F

Acteon nitens Lea, 1846: 257, pl. 36, fig. 60.

Type locality

Petersburg, Dinwiddie County, Virginia, USA; stratum: Yorktown Formation; age: Late Miocene to Middle Pliocene.

Type material

Holotype, ANSP IP1516.

Current taxonomic status

Acteon nitens Lea, 1846.

Acteon novellus Conrad, 1834

Figure 2G–H

Acteon novellus Conrad, 1834: 142.

Type locality

Suffolk, Virginia, USA; stratum: uncertain; age: Miocene/Pliocene(?).

Type material

Syntypes, ANSP IP1600, 1 shell (as “holotype” in Gardner 1948: 277, pl. 38, figs 24, 26), ANSP IP79768, 2 shells (all as “syntypes” in Moore 1962: 80; as “types” in Richards 1968: 165).

Current taxonomic status

Acteon novellus Conrad, 1834 (Moore 1962).

Acteon ovoidea Gabb, 1862

Figure 2I–L

Actaeon ovoidea Gabb, 1862: 319.

Type locality

New Jersey, USA; stratum: “Lower Green Marls” (Navesink Formation); age: Cretaceous.

Type material

Syntypes, ANSP IP56166, 1 shell, ANSP IP56167, 1 shell.

Current taxonomic status

Acteon cretacea Gabb, 1862 (Weller 1907, Richards and Ramsdell 1962).

Acteon politus (Gabb, 1869)

Figure 2M–N

Ringinella polita Gabb, 1869: 174–175, 231, pl. 28, fig. 60.

Type locality

Colusa County, California, USA; stratum: Shasta Group (Shasta Formation); age: Cretaceous.

Type material

Lectotype, ANSP IP4266 (designation in Stewart 1926: 431, fig. 18); paralectotype, ANSP IP79514, 11 shells (as “types lot” in Richards 1968: 176).

Current taxonomic status

Acteon politus (Gabb, 1869) (Anderson 1958).

Acteon pomilius Conrad, 1833

Figure 2O–P

Acteon pomilius Conrad, 1833b: 45.

Type locality

Claiborne Bluff, Alabama River, Monroe County, Alabama, USA; stratum: Gosport Sand (uppermost Claiborne Group); age: Eocene.

Type material

Syntypes, ANSP IP30546 (as “lectotype?” in Moore 1962: 88), ANSP IP53815, 5 shells (all as “syntypes” in Richards 1968: 177).

Current taxonomic status

Acteon pomilius Conrad, 1833 (Palmer and Brann 1966).

Acteon punctatus Lea, 1833

Figure 2Q–R

Acteon punctatus Lea, 1833: 111, pl. 4, fig. 96.

Type locality

Claiborne Bluff, Alabama River, Monroe County, Alabama, USA; stratum: Gosport Sand (uppermost Claiborne Group); age: Eocene.

Type material

Syntypes, ANSP IP5537, 1 shell (as “holotype” in Richards 1968: 179, and in Palmer and Brann 1966: 482), ANSP IP5538, 1 shell, ANSP IP5539, 1 shell, ANSP IP5540, 1 shell (as “paratype” in Richards 1968: 179).

Current taxonomic status

Valid as Acteon pomilius punctatus Lea, 1833 (Palmer and Brann 1966), but possible synonym of Acteon pomilius Conrad, 1833 (Harris 1895a).

Acteon subovoides Whitfield, 1892

Figure 2I–L

Actaeon subovoides Whitfield, 1892: 155, pl. 19, figs 14–16.

Type locality

New Jersey, USA; stratum: “Lower Green Marls” (Navesink Formation); age: Cretaceous.

Type material

Holotype, ANSP IP56166 [also a syntype of Actaeon ovoidea Gabb, 1862].

Current taxonomic status

Acteon cretacea Gabb, 1862 (Weller 1907, Richards and Ramsdell 1962).

Acteon subtornatilis Pilsbry & Johnson, 1917

Figure 2S–T

Acteon subtornatilis Pilsbry & Johnson, 1917: 150.

Type locality

Dominican Republic; stratum: “Santo Domingan Beds” (either Cercado or Gurabo Formations); age: Miocene/Pliocene.

Type material

Holotype, ANSP IP3183 (as “type” in Pilsbry 1922: 310, 431, pl. 23, fig. 15; Richards 1968: 193).

Current taxonomic status

Acteon subtornatilis Pilsbry & Johnson, 1917 (Pilsbry 1922).

Genus Nucleopsis Conrad, 1860

Nucleopsis subvaricatus (Conrad, 1860)

Figure 3A–B

Acteonina subvaricata Conrad, 1860: 294, pl. 47, fig. 22.

Type material

Lectotype, ANSP IP30692 (designation in Palmer 1937: 503, pl. 90, fig. 18; see also Moore 1962: 100, Salvador and Cunha 2016: figs 2A–E); paralectotypes, ANSP IP30693, 2 shells (Richards 1968: 193; Salvador and Cunha 2016: figs 2F–G).

Type locality

Claiborne, Alabama, USA; stratum: likely Gosport Sand (uppermost Claiborne Group); age: Eocene.

Current taxonomic status

Nucleopsis subvaricatus (Conrad, 1860) (Salvador and Cunha 2016).

Figure 3. 

Types. A–B. Lectotype of Nucleopsis subvaricatus, H 10.8 mm, ANSP IP30692. C–D. Holotype of Rictaxis andersoni, H 10.5 mm, ANSP IP13411. E–F. Syntype of Rictaxis oryza, H 6.8 mm, ANSP IP3181. G–H. Lectotype of Tornatellaea bella, H 12.5 mm, ANSP IP30691. I–J. Lectotype of Tornatellaea impressa, H 10.7 mm, ANSP IP4286. K–L. Holotype of Volvaria reticulata, H 7.2 mm, ANSP IP6467. M–N. Holotype of Acteonella oviformis, H 42.1 mm, ANSP IP4323. O–P. Syntype of Avellana bullata, H 25.4 mm, ANSP IP289. Q–R. Syntype of Avellana costata, H 4.5 mm, ANSP IP691. S–T. Lectotype of Biplica obliqua, H 9.9 mm, ANSP IP4263. U–V. Lectotype of Biplica mathewsonii, H 11.6 mm, ANSP IP4262.

Genus Rictaxis Dall, 1871

Rictaxis andersoni (Conrad, 1847)

Figure 3C–D

Actaeon andersoni Conrad, 1847: 287.

Type locality

Vicinity of Vicksburg, Warren County, Mississippi, USA; stratum: Vicksburg Group; age: Oligocene.

Type material

Holotype, ANSP IP13411 (Conrad 1848b: 117, pl. 11, fig. 37, Moore 1962: 38, Richards 1968: 100, MacNeil and Dockery 1984: 232, pl. 39, fig. 10).

Current taxonomic status

Rictaxis andersoni (Conrad, 1847) (MacNeil and Dockery 1984).

Rictaxis oryza (Gabb, 1872)

Figure 3E–F

Actaeonidea oryza Gabb, 1872: 245.

Type locality

Cibao region, Dominican Republic; stratum: uncertain [likely either Cercado or Gurabo Formations]; age: Miocene/Pliocene.

Type material

Syntype, ANSP IP3181 (as “type” in Pilsbry 1922: 310, pl. 23, fig. 12; Richards 1968: 167).

Remarks

Type species of genus Actaeonidea Gabb, 1872, by monotypy.

Current taxonomic status

Rictaxis oryza (Gabb, 1872) (Pilsbry 1922, Woodring 1970).

Genus Tornatellaea Conrad, 1860

Tornatellaea bella Conrad, 1860

Figure 3G–H

Tornatellaea bella Conrad, 1860: 294, pl. 47, fig. 23

Type locality

Alabama(?), USA; stratum: uncertain [likely Gosport Sand (uppermost Claiborne Group)]; age: Eocene.

Type material

Lectotype, ANSP IP30691 (designation by Palmer 1937: 502, pl. 90, fig. 21; as “holotype” in Richards 1968: 177; see also Moore 1962: 41, Salvador and Cunha 2016: fig. S1N–O).

Remarks

Type species of genus Tornatellaea, by monotypy.

Current taxonomic status

Tornatellaea bella Conrad, 1860 (Salvador and Cunha 2016).

Tornatellaea impressa (Gabb, 1864)

Figure 3I–J

Acteon impressus Gabb, 1864: 142, pl. 21, fig. 106.

Type locality

North fork of Cottonwood Creek, Shasta County, California, USA; stratum: uncertain [likely Shasta Formation]; age: Early Cretaceous.

Type material

Lectotype, ANSP IP4286 (designation by Stewart 1926: 434, pl. 24, fig. 8); paralectotypes, ANSP IP79476, 6 shells (Richards 1968: 144).

Current taxonomic status

Tornatellaea impressa (Gabb, 1864) (Anderson 1958).

Genus Volvaria Lamarck, 1801

Volvaria reticulata Johnson, 1899

Figure 3K–L

Volvaria reticulata Johnson, 1899: 71, pl. 1, fig. 1.

Type locality

Moseley’s Ferry, Brazos River, Burleson Co., Texas, USA; stratum: Stone City Beds (middle Claiborne Group); age: Middle Eocene.

Type material

Holotype, ANSP IP6467 (as “type” in Richards 1968: 182).

Current taxonomic status

Volvaria reticulata Johnson, 1899 (Palmer and Brann 1966).

Family Acteonellidae Gill, 1871 †
Genus Acteonella d’Orbigny, 1843

Acteonella oviformis Gabb, 1869

Figure 3M–N

Acteonella oviformis Gabb, 1869: 173, 232, pl. 28, fig. 58.

Type locality

Cottonwood Creek, Shasta County, California, USA; stratum: uncertain [either Shasta or Chico Formations]; age: Cretaceous.

Type material

Holotype, ANSP IP4323 (Stewart 1926: 432, pl. 21, fig. 13; Richards 1968: 168).

Current taxonomic status

Acteonella oviformis Gabb, 1869 (Anderson 1958).

Superfamily Ringiculoidea Philippi, 1853

Family Ringiculidae Philippi, 1853
Genus Avellana d’Orbigny, 1843

Avellana bullata (Morton, 1834)

Figure 3O–P

Tornitella ? bullata Morton, 1834: 48, pl. 5, fig. 3.

Type locality

Merchantville, New Jersey, USA; stratum: uncertain [likely Navesink Formation]; age: Cretaceous.

Type material

Syntypes, ANSP IP289, 1 shell, ANSP IP19702, 1 shell (as “type” in Whitfield 1892: 163, pl. 20, figs 3–4; Richards 1968: 108; Richards and Ramsdell 1962: 93).

Current taxonomic status

Avellana bullata (Morton, 1834) (Richards and Ramsdell 1962).

Avellana costata (Johnson, 1898)

Figure 3Q–R

Cinulia costata Johnson, 1897: 264 [nomen nudum].

Cinulia costata Johnson, 1898: 462, fig. 1.

Type locality

Mount Laurel well, New Jersey, USA; stratum: uncertain; age: Cretaceous.

Type material

Syntypes, ANSP IP691, 1 shell, ANSP IP79408, 2 shells (as “type” in Richards and Ramsdell 1962: 94).

Current taxonomic status

Avellana costata (Johnson, 1898) (Richards and Ramsdell 1962).

Genus Biplica Popenoe, 1957

Biplica obliqua (Gabb, 1864)

Figure 3S–T

Cinulia obliqua Gabb, 1864: 111, pl. 19, fig. 64.

Type locality

Tuscan Springs, Tehama Co., California, USA; stratum: uncertain; age: Late Cretaceous.

Type material

Lectotype, ANSP IP4263 (designation by Stewart 1926: 436, pl. 24, fig. 14; see also Richards 1968: 166; Popenoe 1957: 435).

Current taxonomic status

Biplica obliqua (Gabb, 1864) (Popenoe 1957).

Biplica mathewsonii (Gabb, 1864)

Figure 3U–V

Cinulia mathewsonii Gabb, 1864: 111, 225, pl. 19, fig. 65.

Type locality

Bull’s Head Point, Martinez, California, USA; stratum: uncertain [likely Chico Formation]; age: Cretaceous.

Type material

Lectotype, ANSP IP4262 (designation by Stewart 1926: 437, pl. 24, fig. 11).

Remarks

Popenoe (1957: 434) points out that Gabb’s material from Bull’s Head Point could represent a non-Cretaceous locality/horizon or be Paleogene material mixed with Gabb’s Cretaceous specimens.

Current taxonomic status

Biplica mathewsonii (Gabb, 1864) (Popenoe 1957).

Genus Cinulia Gray, 1840

Cinulia naticoides (Gabb, 1860)

Figure 4A–B

Actaenia naticoides Gabb, 1860c: 299, pl. 48, fig. 2.

Type locality

Mullica Hill, New Jersey, USA; stratum: uncertain [likely Navesink Formation]; age: Cretaceous.

Type material

Syntypes, ANSP IP18784, 2 shells (as “type” in Richards 1968: 164).

Current taxonomic status

Cinulia naticoides (Gabb, 1860) (Clark 1916, Richards and Ramsdell 1962).

Figure 4. 

Types. A–B. Syntype of Cinulia naticoides, H 9.0 mm, ANSP IP18784. C–D. Syntype(?) of Cinulia rectilabrum, H 9.9 mm, ANSP IP4753. E–F. Holotype of Ringicula hypograpta, H 2.0 mm, ANSP IP3841. G–H. Syntype of Ringicula lata, H 15.4 mm, ANSP IP30695. I–J. Lectotype of Ringicula mississippiensis, H 3.7 mm, ANSP IP13414. K–L. Syntype of Ringicula trapaquara, H 2.3 mm, ANSP IP6468. M–N. Lectotype of “Ringicula” varia, H 7.1 mm, ANSP IP4264. O–P. Holotype of Roxania hornii new comb., H 20.1 mm, ANSP IP4232.

Cinulia rectilabrum Gabb, 1869

Figure 4C–D

Cinulia rectilabrum Gabb, 1869: 264, pl. 35, fig. 10.

Type locality

Arivechi, Eastern Sonora, Mexico; stratum: uncertain [either Cañada de Tarachi or El Potrero Grande Units]; age: Late Cretaceous.

Type material

Syntype(?), ANSP IP4753 (as “type?” in Richards 1968: 181).

Current taxonomic status

Cinulia rectilabrum Gabb, 1869 (Stanton 1947).

Genus Ringicula Deshayes, 1838

Ringicula hypograpta Brown & Pilsbry, 1912

Figure 4E–F

Ringicula hypograpta Brown & Pilsbry, 1912: 505, text fig. 2.

Type locality

Scott’s locality 3, excavation of the lower locks at Gatun, Canal Zone, Panama; stratum: Gatun Formation; age: Late Miocene.

Type material

Holotype, ANSP IP3841 (as “type” in Richards 1968: 143).

Current taxonomic status

Ringicula (Ringiculella) semistriata d’Orbigny, 1841 (Woodring 1970).

Ringicula lata (Conrad, 1865)

Figure 4G–H

Actaeon (Nucleopsis) latus Conrad, 1865a: 34.

Type locality

Alabama(?), USA; stratum: uncertain; age: Early Eocene(?).

Type material

Syntype, ANSP IP30695, 1 shell (as “holotype” in Palmer 1937: 502, Richards 1968: 150; as “probable holotype” in Moore 1962: 69; see also Salvador and Cunha 2016: fig. S1K–M).

Current taxonomic status

Ringicula lata (Conrad, 1865) (Salvador and Cunha 2016).

Ringicula mississippiensis Conrad, 1848

Figure 4I–J

Ringicula mississippiensis Conrad, 1848a: 287.

Type locality

Dr. Smith’s plantation, 6 miles northeast of Vicksburg, Warren County, Mississippi, USA; stratum: Vicksburg Group; age: Oligocene.

Type material

Lectotype, ANSP IP13414 (designation by Moore 1962: 77; as “lectotype” in MacNeil and Dockery 1984: 235, Moore 1962: 77; as “holotype” in Richards 1968: 161); paralectotypes, ANSP IP13415, 8 shells (as “paratype” in Richards 1968: 161; MacNeil and Dockery 1984: 235; see also Conrad 1848b: 117, pl. 11, fig. 36).

Current taxonomic status

Ringicula (Ringiculella) mississippiensis Conrad, 1848 (MacNeil and Dockery 1984).

Ringicula trapaquara Harris, 1895

Figure 4K–L

Ringicula trapaquara Harris, 1895a: 76, pl. 8, fig. 7.

Type locality

Moseleys Ferry, Brazos River, Texas; stratum: lower Claiborne Formation; age: Eocene.

Type material

Syntypes, ANSP IP6468, 8 shells (as R. trapaquaria [sic], “paratype” in Richards 1968: 198).

Current taxonomic status

Ringicula trapaquara Harris, 1895 (Palmer 1937).

Ringicula varia Gabb, 1864

Figure 4M–N

Ringicula varia Gabb, 1864: 112, pl. 29, fig. 222a–b.

Type locality

Cow Creek, Shasta County, California, USA; stratum: Chico Formation; age: Cretaceous.

Type material

Lectotype, ANSP IP4264 (designation by Stewart 1926: 435, pl. 24, fig. 3; see also Richards 1968: 202).

Current taxonomic status

Uncertain, as “Ringiculavaria Gabb, 1864 (Stewart 1930).

Order Cephalaspidea P. Fischer, 1883

Superfamily Bulloidea Gray, 1827
Family Bullidae Gray, 1827
Genus Roxania Leach, 1847

Roxania hornii (Gabb, 1864), new comb.

Figure 4O–P

Bulla hornii Gabb, 1864: 143, pl. 29, fig. 235.

Type locality

Kern County, California, USA; stratum: Tejon Formation; age: Eocene.

Type material

Holotype, ANSP IP4232 (Stewart 1926: 439, pl. 29, fig. 9; Richards 1968: 142).

New taxonomic status

Roxania hornii (Gabb, 1864) new comb. This species was placed in the genus Abderospira Dall, 1898 (e.g., Stewart 1926, Keen and Bentson 1944), which is now considered a synonym of Roxania (Valdés 2008).

Genus Bulla Linnaeus, 1758

Bulla macrostoma Gabb, 1860

Figure 5A–B

Bulla macrostoma Gabb, 1860c: 301, pl. 48, fig. 15.

Type locality

Prairie Bluff, Alabama, USA; stratum: uncertain (“white limestone”); age: Cretaceous.

Type material

Holotype, ANSP IP30727 (as “type” in Richards 1968: 155).

Current taxonomic status

Bulla macrostoma Gabb, 1860 (Stephenson 1914).

Figure 5. 

Types. A–B. Holotype of Bulla macrostoma, H 26.2 mm, ANSP IP30727. C–D. Holotype of Bullopsis cretacea, H 18.0 mm, ANSP IP18924. E–F. Holotype of Retusa biforis, H 2.9 mm, ANSP IP3192. G. Paralectotype of Retusa galba, H 14.3 mm, ANSP IP53816. H. Lectotype of Retusa galba, H 15.0 mm, ANSP IP30548. I–J. Lectotype of Retusa sthillairii, H 14.0 mm, ANSP IP5486. K. Syntype of Retusa subspissa, H 4.8 mm, ANSP IP30641. L. Holotype of Volvulella cylichnoides, H 4.2 mm, ANSP IP3177. M–N. Syntype of Volvulella cylindrica, H 4.7 mm, ANSP IP3179.

Genus Bullopsis Conrad, 1858

Bullopsis cretacea Conrad, 1858

Figure 5C–D

Bullopsis cretacea Conrad, 1858: 334.

Type locality

Owl Creek, near Ripley, Tippah County, Mississippi, USA; stratum: “dark gray sandy marl” (Owl Creek Formation); age: Cretaceous.

Type material

Holotype, IPANSP 18924 (Conrad 1860: pl. 46, fig. 27; as “type” in Richards 1968: 120).

Current taxonomic status

Bullopsis cretacea Conrad, 1858 (Stephenson 1955, Sohl 1964).

Family Retusidae Thiele, 1925
Genus Retusa Brown, 1827

Retusa biforis Pilsbry & Johnson, 1917

Figure 5E–F

Retusa biforis Pilsbry & Johnson, 1917: 151

Type locality

Dominican Republic. Type stratum: “Santo Domingan Beds” (either Cercado or Gurabo Formations); age: Miocene/Pliocene.

Type material

Holotype, ANSP IP3192 (as “type” in Richards 1968: 106).

Current taxonomic status

Retusa biforis Pilsbry & Johnson, 1917 (Pilsbry 1922).

Retusa sulcata fossilis Pilsbry, 1922

Retusa sulcata var. fossilis Pilsbry 1922: 311.

Type locality

Dominican Republic; stratum: “Santo Domingan Beds” (either Cercado or Gurabo Formations); age: Miocene/Pliocene.

Type material

Holotype and paratypes, ANSP IP3186 (3 shells; lost).

Current taxonomic status

Uncertain, likely valid as Retusa sulcata fossilis Pilsbry, 1922.

Retusa galba (Conrad, 1833)

Figure 5G–H

Volvaria galba Conrad, 1833a: 34, pl. 18, fig. 2.

Type locality

Claiborne Bluff, Alabama River, Monroe County, Alabama, USA; stratum: Gosport Sand (uppermost Claiborne Group); age: Middle Eocene.

Type material

Lectotype, ANSP IP30548 (designation by Palmer and Brann 1966: 878; as “lectotype?” in Moore 1962: 62); paralectotypes, ANSP IP53816, 3 shells, ANSP IP53817, 1 shell, ANSP IP30549, 8 shells (all as “probable syntypes” in Richards 1968: 136).

Current taxonomic status

Retusa (Cylichnina) galba (Conrad, 1833) (Palmer and Brann 1966).

Retusa sthillairii (Lea, 1833)

Figure 5I–J

Bulla St. Hillairii Lea, 1833: 98, pl. 4, fig. 78.

Type locality

Monroe Co., Claiborne Bluff, Alabama, USA; stratum: Gosport Sand (uppermost Claiborne Group); age: Eocene.

Type material

Lectotype, ANSP IP5486, 1 shell (designation by Palmer 1937: 481; as “holotype” in Palmer and Brann 1966: 878); paralectotypes, ANSP IP5487, 1 shell, ANSP IP5488, 1 shell, ANSP IP5489, 1 shell, ANSP IP5490, 1 shell, ANSP IP5491, 1 shell, ANSP IP5487, 1 shell (all [including ANSP IP5486] as “types” in Richards 1968: 191).

Current taxonomic status

Retusa galba (Conrad, 1833) (Palmer and Brann 1966).

Retusa subspissa (Conrad, 1846)

Figure 5K

Bulla subspissa Conrad, 1846: 20, pl. 1, fig. 29.

Type locality

Calvert Cliffs, Maryland, USA; stratum: Calvert Formation; age: Miocene.

Type material

Syntype, ANSP IP30641, 1 shell (as “probable holotype” in Moore 1962: 100; as “type?” in Richards 1968: 193).

Current taxonomic status

Retusa subspissa (Conrad, 1846), but possible synonym of Retusa conulus (Deshayes, 1824) (Martin 1904).

Family Rhizoridae Dell, 1952
Genus Volvulella Newton, 1891

Volvulella cylichnoides (Pilsbry & Johnson, 1917)

Figure 5L

Volvula cylichnoides Pilsbry & Johnson, 1917: 151.

Type locality

Dominican Republic; stratum: “Santo Domingan Beds” (either Cercado or Gurabo Formations); age: Miocene/Pliocene.

Type material

Holotype, ANSP IP3177 (as “type” in Richards 1968: 121); paratype, ANSP IP79027, 1 shell.

Current taxonomic status

Volvulella cylichnoides (Pilsbry & Johnson, 1917), but could be a synonym of Retusa yaquensis Maury, 1917 (Pilsbry 1922).

Volvulella cylindrica (Gabb, 1872)

Figure 5M–N

Volvula cylindrica Gabb, 1872: 246 [non Carpenter, 1865; non E.A. Smith, 1871].

Type locality

Dominican Republic; type stratum: uncertain [likely either Cercado or Gurabo Formations]; age: Miocene/Pliocene.

Type material

Syntype, ANSP IP3179, 1 shell (as “type” in Richards 1968: 121).

Current taxonomic status

Volvulella persimilis (Mörch, 1875) (Dall 1889, Pilsbry 1922).

Volvulella micratracta Brown & Pilsbry, 1912

Figure 6A

Volvulella micratracta Brown & Pilsbry, 1912: 504, text fig. 1.

Type locality

Scott’s locality 3, excavation of the lower locks at Gatun, Canal Zone, Panama; stratum: Gatun Formation; age: Late Miocene.

Type material

Holotype, ANSP IP3842 (as “type” in Richards 1968: 158).

Current taxonomic status

Volvulella micratracta Brown & Pilsbry, 1912 (Woodring 1970).

Figure 6. 

Types. A. Holotype of Volvulella micratracta, H 1.5 mm, ANSP IP3842. B. Syntype of Volvulella minutissima new comb., H 2.3 mm, ANSP IP13267. C–D. Holotype of Volvulella ornata, H 3.2 mm, ANSP IP3178. E. Holotype of Volvulella parallela, H 2.1 mm, ANSP IP3188. F. Holotype of Volvulella tritica, H 4.1 mm, ANSP IP19104. G. Holotype of Atys cinctorii, H 2.6 mm, ANSP IP3185. H. Syntype of Atys sulculorum, H 2.5 mm, ANSP IP3317. I–J. Syntype of Cylichna cylindrus, H 4.7 mm, ANSP IP1554. K. Syntype of Cylichna dekayi, H 5.0 mm, ANSP IP6008. L. Syntype of Cylichna kellogii, H 5.7 mm, ANSP IP13266. M. Holotype of Cylichna recta, H 8.0 mm, ANSP IP18782. N. Syntype of Bulla occidentalis, H 10.0 mm, ANSP IP17139. O–P. Lectotype of Cylichna costata, H 16.0 mm, ANSP IP4338. Q–R. Syntype of Acteon sculptus, H 1.9 mm, ANSP IP1515. S. Syntype of Acteon milium, H 2.1 mm, ANSP IP1520. T. Holotype of Acteon angulatus, H 2.4 mm, ANSP IP1521.

Volvulella minutissima (Gabb, 1860), new comb.

Figure 6B

Volvula minutissima Gabb, 1860b: 386–387, pl. 67, fig. 52.

Type locality

Caldwell County, Texas, USA; stratum: uncertain; age: Eocene.

Type material

Syntype, ANSP IP13267, 1 shell (as “type” in Richards 1968: 159).

New taxonomic status

Volvulella minutissima (Gabb, 1860), new comb. Harris (1895b) considered the species valid. The genus Volvulella Newton, 1891 is a replacement name for Volvula A. Adams, 1850 non Gistel, 1848 (Valdés 2008).

Volvulella ornata (Pilsbry & Johnson, 1917)

Figure 6C–D

Volvula ornata Pilsbry & Johnson, 1917: 151.

Type locality

Dominican Republic; stratum: “Santo Domingan Beds” (either Cercado or Gurabo Formations); age: Miocene/Pliocene.

Type material

Holotype, ANSP IP3178 (as “type” in Richards 1968: 167); paratypes, ANSP IP81665, 6 shells.

Remarks

Pilsbry and Johnson (1917) did not clearly indicate which one of their specimens is the holotype, merely indicating the presence of a “Type”. Judging by the conventions used elsewhere in their paper, we here consider the holotype to be the specimen measured by these authors. Besides the holotype, Pilsbry and Johnson (1917) mentioned seven specimens, one of which is presently missing.

Current taxonomic status

Volvulella ornata (Pilsbry & Johnson, 1917) (Pilsbry 1922).

Volvulella parallela (Pilsbry & Johnson, 1917)

Figure 6E

Volvula parallela Pilsbry & Johnson, 1917: 151.

Type locality

Dominican Republic; stratum: “Santo Domingan Beds” (either Cercado or Gurabo Formations); age: Miocene/Pliocene.

Type material

Holotype, ANSP IP3188 (as “type” in Richards 1968: 169); paratypes, ANSP IP79026, 2 shells.

Current taxonomic status

Volvulella cylindrica parallela (Pilsbry & Johnson, 1917) (Woodring 1970).

Volvulella tritica (Olsson & Harbison, 1953)

Figure 6F

Volvula tritica Olsson & Harbison, 1953: 163, pl. 25, figs 3–3a.

Type locality

St. Petersburg, Pinellas County, Florida, USA; stratum: North St. Petersburg Beds; age: Plio-Pleistocene.

Type material

Holotype, ANSP IP19104; paratypes, ANSP IP79270, 2 shells (all as “types” in Richards 1968: 199).

Current taxonomic status

Volvulella tritica (Olsson & Harbison, 1953) (Portell and Kittle 2010).

Superfamily Haminoeoidea Pilsbry, 1895
Family Haminoeidae Pilsbry, 1895
Genus Atys Montfort, 1810

Atys cinctorii Pilsbry & Johnson, 1917

Figure 6G

Atys cinctorii Pilsbry & Johnson, 1917: 152.

Type locality

Dominican Republic; stratum: “Santo Domingan Beds” (either Cercado or Gurabo Formations); age: Miocene/Pliocene.

Type material

Holotype, ANSP IP3185 (as “type” in Richards 1968: 113).

Current taxonomic status

Atys cinctorii Pilsbry & Johnson, 1917 (Pilsbry 1922).

Atys sulculorum Pilsbry & Johnson, 1917

Figure 6H

Atys sulculorum Pilsbry & Johnson, 1917: 152.

Type locality

Dominican Republic; stratum: “Santo Domingan Beds” (either Cercado or Gurabo Formations); age: Miocene/Pliocene.

Type material

Syntypes, ANSP IP3317, 2 shells (as “type” in Richards 1968: 194).

Current taxonomic status

Atys doliolum (Maury, 1917) (Woodring 1970).

Superfamily Cylichnoidea H. Adams & A. Adams, 1854
Family Cylichnidae H. Adams & A. Adams, 1854
Genus Cylichna Lovén, 1846

Cylichna cylindrus (Lea, 1846)

Figure 6I–J

Bulla cylindrus Lea, 1846: 250–251, pl. 35, fig. 43.

Type locality

Petersburg, Dinwiddie County, Virginia, USA; stratum: Yorktown Formation; age: Late Miocene to Middle Pliocene.

Type material

Syntypes, ANSP IP1554, 1 shell (Gardner 1948: 279, pl. 38, fig. 27; as “type” in Richards 1968: 122), ANSP IP79382, 1 shell (Gardner 1948: 279, pl. 38, fig. 28).

Current taxonomic status

Cylichna cylindrus (Lea, 1846) (Gardner 1948).

Cylichna dekayi (Lea, 1833)

Figure 6K

Bulla dekayi Lea, 1833: 200, pl. 6, 215.

Type locality

Claiborne, Alabama, USA; stratum: Claiborne Group; age: Eocene.

Type material

Syntypes, ANSP IP6007, 1 shell (as “holotype” in Richards 1968: 123), ANSP IP6008, 1 shell (as “paratype” in Richards 1968: 123).

Current taxonomic status

Cylichna (Mnestia) dekayi (Lea, 1833) (Glibert 1962).

Cylichna kellogii (Gabb, 1860)

Figure 6L

Bulla kellogii Gabb, 1860b: 386, pl. 67, fig. 50.

Type locality

Texas, USA; stratum: uncertain; age: Eocene.

Type material

Syntype, ANSP IP13266, 1 shell (as “type” in Richards 1968: 148).

Current taxonomic status

Cylichna (Acrotrema) kellogii (Gabb, 1860) (Gardner 1945).

Cylichna recta (Gabb, 1860)

Figure 6M

Bulla recta Gabb, 1860c: 303, pl. 48, fig. 16.

Type locality

Burlington County, New Jersey, USA; stratum: “Lower Green Marls” (Navesink Formation); age: Cretaceous.

Type material

Holotype, ANSP IP18782 (as “type” in Richards 1968: 181).

Current taxonomic status

Cylichna recta (Gabb, 1860) (Wade 1926, Richards and Ramsdell 1962).

Superfamily Philinoidea Gray, 1850
Family Philinidae Gray, 1850
Genus Philine Ascanius, 1772

Philine gabbi Cossmann, 1895

Philine (Megistostoma) gabbi Cossmann, 1895: 127.

Type locality

Martinez, California, USA; stratum: uncertain; age: Eocene.

Type material

Syntypes ANSP IP4216 (of M. striatum Gabb, 1864), 2 shells (lost).

Remarks

This was a new name for Megistostoma striatum Gabb, 1864, when transferred to the genus Philine [non striata Deshayes, 1824]. However, the name P. gabbiana (Stoliczka, 1868) has precedence over it (see below).

Current taxonomic status

Philine (Megistostoma) gabbiana (Stoliczka, 1868) (Keen and Bentson 1944).

Philine gabbiana Stoliczka, 1868

Bullaea Gabbiana Stoliczka, 1868: 434.

Type locality

Martinez, California, USA; stratum: uncertain; age: Eocene.

Type material

Syntypes ANSP IP4216 (of M. striatum Gabb, 1864), 2 shells (lost).

Remarks

This was coined as a new name for Megistostoma striatum Gabb, 1864, when transferred to the genus Bullaea Lamarck, 1801 [non striata Deshayes]. Bullaea is presently considered a synonym of Philine.

Current taxonomic status

Philine (Megistostoma) gabbiana (Stoliczka, 1868) (Keen and Bentson 1944).

Megistostoma striatum Gabb, 1864

Megistostoma striata [sic] Gabb, 1864: 144, 229, pl. 21, fig. 108.

Type locality

Martinez, California, USA; stratum: uncertain; age: Eocene.

Type material

Syntypes, ANSP IP4216 (lost), 2 shells (Stewart 1926: 442: pl. 26, fig. 2; as “holotype” and “paratype” in Keen and Bentson 1944: 170; as “holotypes” in Richards 1968: 191).

Remarks

Type species of Megistostoma Gabb, 1864. The correct epithet is striatum (not striata), since the ending -stoma is neuter.

Current taxonomic status

Philine (Megistostoma) gabbiana (Stoliczka, 1868) (Keen and Bentson 1944), new name for M. striatum Gabb, 1864 (see above).

Family Scaphandridae G.O. Sars, 1878
Genus Ellipsoscapha Stephenson, 1941

Ellipsoscapha mortoni (Forbes, 1845)

Bulla mortoni Forbes, 1845: 63, fig. A.

Type locality

New Jersey, USA; stratum: uncertain [likely Navesink Formation]; age: Cretaceous.

Type material

Holotype not found; could be in Charles Lyell’s fossil collections (John Sime, pers. comm.), presently in the Natural History Museum (London, UK) and Oxford University Museum of Natural History (Oxford, UK).

Current taxonomic status

Ellipsoscapha mortoni (Forbes, 1845) (Richards and Ramsdell 1962, Sohl 1964).

Ellipsoscapha occidentalis (Meek & Hayden, 1856)

Figure 6N

Bulla occidentalis Meek & Hayden, 1856 (non A. Adams, 1850): 69.

Type locality

Yellowstone River (150 miles above its mouth), near Glendive, Montana, USA; stratum: Pierre Shale; age: Late Cretaceous.

Type material

Syntypes, ANSP IP17139, 2 shells.

Current taxonomic status

Ellipsoscapha occidentalis (Meek & Hayden, 1856) (Sohl 1967). Substitution of the junior primary homonym is not mandatory if the conditions of Article 23.9.5 ICZN (1999) are met, but a request for a ruling of the Commission under its plenary powers to validate the junior homonymous name would be necessary.

Genus Scaphander Montfort, 1810

Scaphander costatus (Gabb, 1864)

Figure 6O–P

Cylichna costata Gabb, 1864: 143, pl. 2, fig. 107.

Type locality

Martinez, Contra Costa County, California, USA; stratum: Tejon Formation s. l.; age: Eocene.

Type material

Lectotype, ANSP IP4338 (designation by Stewart 1926: 437, pl. 27, fig. 3; see also Richards 1968: 118); paralectotypes, ANSP IP79477, 8 shells.

Remarks

Type species of subgenus Mirascapha Stewart, 1927.

Current taxonomic status

Scaphander (Mirascapha) costatus (Gabb, 1864) (Squires 1984).

Species transferred to other gastropod groups

Here are listed the species (with type material in the ANSP collection) that were originally classified in “Architectibranchia” and Cephalaspidea, but that actually do not belong to them. Some of them have already undergone taxonomical revision and are listed concisely in Table 1, while others are reclassified below.

Species (with type material in the ANSP collection) that were originally classified in Architectibranchia and Cephalaspidea, but that after revision were reclassified in other groups. The species are arranged in alphabetical order of the specific epithets, with information on their original description, type specimens in the ANSP collection, current taxonomic status, and references of such status. Abbreviations: Hol = holotype; Lec = lectotype; Pal = Paralectotype(s); Par = paratype(s); Syn = syntype(s).

Original description Type material (ANSP) Current taxonomic status Family Reference(s)
calafia, “Acteonina” Stewart, 1926: 432, pl. 21, fig. 12 4287 (Hol) Paosia calafia (Stewart, 1926) Pseudomelaniidae Squires and Saul (2004)
californica, Acteonina Gabb, 1864: 128, pl. 29, fig. 230a–b 4259 (Pal) Paosia californica (Gabb, 1864) Pseudomelaniidae Squires and Saul (2004)
curta, Globiconcha Gabb, 1862: 319 31393 (Hol) Tylostoma elevatum (Shumard, 1853) Tylostomatidae Stanton (1947)
elevatus, Acteon Lea, 1833: 113, pl. 4, fig. 98 5545 (Lec); 5546 to 5549 (Pal) Pyramidella larvata Conrad, 1833 Pyramidellidae Palmer (1937), Palmer and Brann (1966)
globosus, Acteon Lea, 1846: 255, pl. 36, fig. 55 1518 (Syn) Iselica globosa (Lea, 1846) Amathinidae Lee (2015)
granulatus, Acteon Lea, 1846: 255, pl. 36, fig. 54 1533 (Syn) Odostomia granulatus (Lea, 1846) Pyramidellidae Holmes (1860)
laevis, Actaeon Lea, 1841: 94, pl. 1, fig. 4 lost (Hol) Odostomia laevis (Lea, 1847) Pyramidellidae Palmer (1937)
linteus, Solidulus Conrad, 1858: 334, pl. 35, fig. 10 lost (Hol) Eoacteon linteus (Conrad, 1858) Acteoninidae Stephenson (1955), Sohl (1964)
magnoplicatus, Acteon Lea, 1841: 94, pl. 1, fig. 5 13158 (Hol) Odostomia (Evalea) melanella (Lea, 1833) Pyramidellidae Palmer (1937)
melanellus, Acteon Lea, 1833: 113, pl. 4, fig. 99 5550 (Lec); 5551 to 5557 (Pal) Odostomia (Evalea) melanella (Lea, 1833) Pyramidellidae Palmer (1937)
pygmaeus, ?Acteon Lea, 1833: 114, pl. 4, fig. 101 5559 (Hol) Pyramidella larvata Conrad, 1833 Pyramidellidae Palmer (1937), Palmer and Brann (1966)
simplex, Acteon Lea, 1843: 8 1519 (Syn) Odostomia simplex (Lea, 1843) Pyramidellidae Ward and Blackwelder (1987)
striatus, Acteon Lea, 1833: 114, pl. 4, fig. 100 5558 (Hol) Odostomia (Evalea) melanella alveata (Lea, 1833) Pyramidellidae Palmer (1937), Palmer and Brann (1966)
turbinatus, Acteon Lea, 1843: 256, pl. 36, fig. 56 1522 (Syn) Odostomia turbinatus (Lea, 1846) Pyramidellidae Ward and Blackwelder (1987)

Panpulmonata

Superfamily Pyramidelloidea Gray, 1840
Family Pyramidellidae Gray, 1840
Genus Chrysallida Carpenter, 1856

Chrysallida sculpta (Lea, 1846), new comb.

Figure 6Q–R

Acteon sculptus Lea, 1846: 257, pl. 36, fig. 59.

Type locality

Petersburg, Dinwiddie County, Virginia, USA; stratum: Yorktown Formation; age: Late Miocene to Middle Pliocene.

Type material

Syntypes, ANSP IP1515, 2 shells (as “types” in Richards 1968: 186).

Taxonomical reassessment

This species is better classified in the genus Chrysallida due to the following conchological features (Robba 2013): a weak or absent columellar fold, teleoconch sculpture consisting of collabral ribs and spiral cords of similar strength. Most species of Chrysallida are strongly sculptured, as the present specimens, with the ribs forming nodes where crossing the spiral cords and tending to fade away near the base.

Genus Odostomia Fleming, 1813

Odostomia milium (Lea, 1846), new comb.

Figure 6S

Acteon milium Lea, 1846: 257, pl. 36, fig. 61.

Type locality

Petersburg, Dinwiddie County, Virginia, USA. Type stratum: Yorktown Formation. Age: Late Miocene to Middle Pliocene.

Type material

Syntypes, ANSP IP1520, 1 shell, ANSP IP79385, 1 shell (lost, not found in the vial).

Taxonomical reassessment

This species is better classified in the genus Odostomia due to the following conchological features (Robba 2013): verall ovate-conical shell with moderately convex whorls, suture with a blunt subsutural margin, slit-like umbilicus, and prominent columellar fold.

Genus Pyrgulina A. Adams, 1863

Pyrgulina angulata (Lea, 1846), new comb.

Figure 6T

Acteon angulatus Lea, 1846: 256, pl. 36, fig. 57.

Type locality

Petersburg, Dinwiddie County, Virginia, USA. Type stratum: Yorktown Formation. Age: Late Miocene to Middle Pliocene.

Type material

Holotype, ANSP IP1521.

Taxonomical reassessment

This species is better classified in the genus Pyrgulina due to the following conchological features (Robba 2013): elongated (somewhat turreted) conical shell, convex whorls, distinct subsutural shelf, deep suture, preence of a columellar fold present, teleoconch sculpture consisting of axial ribs (including on the base) overridden by spiral threads.

Campbell (1993: 68) considers Acteon angulatus Lea 1846 a synonym of Melanella angulata (Lea, 1846) (Eulimidae), which is clearly not the case. The present species lacks the diagnostic conchological features of the latter genus (and of eulimids in general), such as: a high spire, a smooth teleoconch with fine axial marks, and the absence of a columellar fold (Warén 1984, Geiger 2016).

List of taxa by species-group names

Here is presented a list of the species whose types can be found in the ANSP collection, arranged alphabetically by specific epithet. Species appear first in their original generic allocation and then in their current systematic position. An “*” after the specific epithet indicates that the type material is lost.

andersoni , Actaeon Conrad, 1848a: 287. Rictaxis andersoni (Conrad, 1848). Acteonidae.

bella , Tornatellaea Conrad, 1860: 294. Tornatellaea bella Conrad, 1860. Acteonidae.

biforis , Retusa Pilsbry & Johnson, 1917: 151. Acteon gabbana Whitfield, 1892. Retusidae.

biplicata , Acteonina Gabb, 1860a: 93. Acteocina canaliculata (Say, 1826). Acteonidae.

bullata , Tornitella? Morton, 1834: 48. Avellana bullata (Morton, 1834). Ringiculidae.

calafia , “Acteonina” Stewart, 1926: 432. Paosia calafia (Stewart, 1926). Pseudomelaniidae.

cederstromi , Acteocina Richards, 1947. Acteocina candei (d’Orbigny, 1841). Acteocinidae.

chowanensis , Acteocina Richards, 1947: 34. Acteocina canaliculata (Say, 1826). Acteocinidae.

cinctorii , Atys Pilsbry & Johnson, 1917: 152. Atys cinctorii Pilsbry & Johnson, 1917. Haminoeidae.

costata , Cinulia Johnson, 1898: 462. Avellana costata (Johnson, 1898). Ringiculidae.

costata , Cylichna Gabb, 1864: 143. Scaphander (Mirascapha) costatus (Gabb, 1864). Scaphandridae.

costellatus *, Acteon Conrad, 1833b: 45. Status uncertain. Acteonidae.

crassiplica , Bulla Conrad, 1848a: 282. Acteocina crassiplica (Conrad, 1848). Bullidae.

cretacea , Acteon Gabb, 1862: 318. Acteon cretacea Gabb, 1862. Acteonidae.

cretacea , Bullopsis Conrad, 1858: 334. Bullopsis cretacea Conrad, 1858. Bullidae.

cylichnoides , Volvula Pilsbry & Johnson, 1917: 151. Volvulella cylichnoides (Pilsbry & Johnson, 1917). Rhizoridae.

cylindrica , Volvula Gabb, 1872: 246 [non Carpenter, 1865]. Volvulella persimilis (Mörch, 1875). Rhizoridae.

cylindrus , Bulla Lea, 1846: 250. Cylichna cylindrus (Lea, 1846). Cylichnidae.

dekayi , Bulla Lea, 1833: 200. Cylichna (Mnestia) dekayi (Lea, 1833). Cylichnidae.

elegans , Monoptygma Lea, 1833: 203. Acteon pomilius Conrad, 1833. Acteonidae.

forbesiana , Actaeon Whitfield, 1892: 157. Acteon cretacea Gabb, 1862. Acteonidae

fossilis *, Retusa sulcata Pilsbry, 1922: 311. Retusa sulcata fossilis Pilsbry, 1922. Retusidae.

gabbana , Actaeon Whitfield, 1892: 156. Acteon gabbana Whitfield, 1892. Acteonidae.

gabbi , Philine (Megistostoma) Cossmann, 1895: 127. Philine (Megistostoma) gabbiana (Stoliczka, 1868). Philinidae.

gabbiana , Bullaea Stoliczka, 1868: 434. Philine (Megistostoma) gabbiana (Stoliczka, 1868). Philinidae.

galba , Volvaria Conrad, 1833a: 34. Retusa galba (Conrad, 1833). Retusidae.

glans , Acteon Lea, 1846: 256. Acteon glans Lea, 1846. Acteonidae.

hornii , Bulla Gabb, 1864: 143. Roxania hornii (Gabb, 1864) new comb. Scaphandridae.

hypograpta , Ringicula Brown & Pilsbry, 1912: 505. Ringicula (Ringiculella) semistriata d’Orbigny, 1841. Ringiculidae.

idoneus , Acteon Conrad, 1833b: 45. Acteon idoneus Conrad, 1833. Acteonidae

impressus , Acteon Gabb, 1864: 142. Tornatellaea impressa (Gabb, 1864). Acteonidae.

kirkwoodiana , Acteocina Richards & Harbison, 1944: 9. Acteocina kirkwoodiana Richards & Harbison, 1944. Acteocinidae.

kellogii , Bulla Gabb, 1860b: 386. Cylichna (Acrotrema) kellogii (Gabb, 1860). Cylichnidae.

latus , Actaeon (Nucleopsis) Conrad, 1865a: 34. Ringicula lata (Conrad, 1865). Ringiculidae.

lineatus , Acteon Lea, 1833: 112. Acteon idoneus Conrad, 1833. Acteonidae.

macrostoma , Bulla Gabb, 1860c: 301. Bulla macrostoma Gabb, 1860. Bullidae.

mathewsonii , Cinulia Gabb, 1864: 111. Biplica mathewsonii (Gabb, 1864). Ringiculidae.

micratracta , Volvulella Brown & Pilsbry, 1912. Volvulella micratracta Brown & Pilsbry, 1912. Rhizoridae.

minutissima , Volvula Gabb, 1860b: 386. Volvulella minutissima (Gabb, 1860) new comb. Rhizoridae.

mississippiensis , Ringicula Conrad, 1848a: 287. Ringicula (Ringiculella) mississippiensis Conrad, 1848. Ringiculidae.

modicellus *, Actaeon Conrad, 1860: 287. Acteon modicellus Conrad, 1860. Acteonidae.

mortoni *, Bulla Forbes, 1845: 63. Haminoea mortoni Forbes, 1845. Haminoeidae.

naticoides , Actaenia Gabb, 1860c: 299. Cinulia naticoides (Gabb, 1860). Ringiculidae.

nitens , Acteon Lea, 1846. Acteon nitens Lea, 1846. Acteonidae.

novellus , Acteon Conrad, 1834: 142. Acteon novellus Conrad, 1834. Acteonidae.

obliqua , Cinulia Gabb, 1864: 111. Biplica obliqua (Gabb, 1864). Ringiculidae

occidentalis , Bulla Meek & Hayden, 1856: 69. Ellipsoscapha occidentalis (Meek & Hayden, 1856). Scaphandridae.

ornata , Volvula Pilsbry & Johnson, 1917: 151. Volvulella ornata (Pilsbry & Johnson, 1917). Rhizoridae.

oryza , Acteonidea Gabb, 1872: 245. Rictaxis oryza (Gabb, 1872). Acteonidae.

oviformis , Acteonella Gabb, 1869: 173. Acteonella oviformis Gabb, 1869. Acteonellidae.

ovoidea , Actaeon Gabb, 1862: 319. Acteon cretacea Gabb, 1862. Acteonidae.

parallela , Volvula Pilsbry & Johnson, 1917: 151. Volvulella cylindrica parallela (Pilsbry & Johnson, 1917). Rhizoridae.

polita , Ringinella Gabb, 1869: 174. Acteon politus (Gabb, 1869). Acteonidae.

pomilius , Acteon Conrad, 1833b: 45. Acteon pomilius Conrad, 1833. Acteonidae.

punctatus , Acteon Lea, 1833: 111. Acteon pomilius punctatus Lea, 1833. Acteonidae.

puruha , Acteocina Pilsbry & Olsson, 1941: 13. Acteocina puruha Pilsbry & Olsson, 1941. Acteocinidae.

recta , Bulla Gabb, 1860c: 303. Cylichna recta (Gabb, 1860). Cylichnidae.

rectilabrum , Cinulia Gabb, 1869: 264. Cinulia rectilabrum (Gabb, 1869). Ringiculidae.

reticulata , Volvaria Johnson, 1899: 71. Volvaria reticulata Johnson, 1899. Acteonidae.

sthillairii , Bulla Lea, 1833: 98. Retusa (Cylichnina) galba (Conrad, 1833). Retusidae.

striata, Megistostoma Gabb, 1864: 144. Philine (Megistostoma) gabbiana (Stoliczka, 1868). Philinidae.

subbullata , Acteocina Pilsbry & Johnson, 1917: 150. Acteocina bullata (Kiener, 1834). Acteocinidae.

subovoides , Actaeon Whitfield, 1892: 155. Acteon cretacea Gabb, 1862. Acteonidae.

subspissa , Bulla Conrad, 1846: 20. Retusa subspissa (Conrad, 1846). Retusidae.

subtornatilis , Acteon Pilsbry & Johnson, 1917: 150. Acteon subtornatilis Pilsbry & Johnson, 1917. Acteonidae.

subvaricata , Acteonina Conrad, 1860: 294. Nucleopsis subvaricatus (Conrad, 1860). Acteonidae.

sulculorum , Atys Pilsbry & Johnson, 1917: 152. Atys doliolum (Maury, 1917). Haminoeidae.

trapaquara , Ringicula Harris, 1895a: 76. Ringicula trapaquara (Harris, 1895a). Ringiculidae.

tritica , Volvula Olsson & Harbison, 1953: 163. Volvulella tritica (Olsson & Harbison, 1953). Rhizoridae.

varia , Ringicula Gabb, 1864: 112. Ringicula varia Gabb, 1864. Ringiculidae.

weatherlli , Acteon Lea, 1833: 213. Acteocina canaliculata (Say, 1826). Acteocinidae.

List of taxa by authorship

Here the list of types is arranged by author (alphabetically) and date; see “References” section for full citation. Species appear in their original generic allocation; see text for current systematic status and placement. An “*” indicates that the type material is lost.

Brown, A. P. & Pilsbry, H. A.

1912 hypograpta, Ringicula

1912 micratracta, Volvulella

Conrad, T. A.

1833a galba, Volvaria

1833b costellatus, Acteon*

1833b idoneus, Acteon

1833b pomilius, Acteon

1834 novellus, Acteon

1846 subspissa, Bulla

1848a andersoni, Actaeon

1848a crassiplica, Bulla

1848a mississippiensis, Ringicula

1858 cretacea, Bullopsis

1860 bella, Tornatellaea

1860 modicellus, Actaeon*

1860 subvaricata, Acteonina

1865a latus, Actaeon (Nucleopsis)

Cossmann, M.

1895 gabbi, Philine (Megistostoma)

Forbes, E.

1845 mortoni, Bulla*

Gabb, W. M.

1860a biplicata, Acteonina

1860b kellogii, Bulla

1860b minutissima, Volvula

1860c macrostoma, Bulla

1860c naticoides, Actaenia

1860c recta, Bulla

1862 cretacea, Acteon

1862 curta, Globiconcha

1862 ovoidea, Actaeon

1864 costata, Cylichna

1864 hornii, Bulla

1864 impressus, Acteon

1864 mathewsonii, Cinulia

1864 obliqua, Cinulia

1864 striata, Megistostoma

1864 varia, Ringicula

1869 oviformis, Acteonella

1869 polita, Ringinella

1869 rectilabrum, Cinulia

1873 cylindrica, Volvula

1873 oryza, Acteonidea

Harris, G. D.

1895a trapaquara, Ringicula

Johnson, C. W.

1898 costata, Cinulia

1898 reticulata, Volvaria

Lea, H. C.

1846 cylindrus, Bulla

1846 glans, Acteon

1846 nitens, Acteon

Lea, I.

1833 dekayi, Bulla

1833 elegans, Monoptygma

1833 lineatus, Acteon

1833 punctatus, Acteon

1833 sthillairii, Bulla

1833 weatherlli, Acteon

Meek, F. B. & Hayden, F. V.

1856 occidentalis, Bulla

Morton, S. G.

1834 bullata, Tornitella?

Olsson, A. A. & Harbison, A.

1953 tritica, Volvula

Pilsbry, H. A.

1922 fossilis, Retusa sulcata*

Pilsbry, H. A. & Johnson, C. W.

1917 biforis, Retusa

1917 cinctorii, Atys

1917 cylichnoides, Volvula

1917 ornata, Volvula

1917 parallela, Volvula

1917 subbullata, Acteocina

1917 subtornatilis, Acteon

1917 sulculorum, Atys

Pilsbry, H. A. & Olsson, A. A.

1941 puruha, Acteocina

Richards, H. G.

1947 cederstromi, Acteocina

1947 chowanensis, Acteocina

Richards, H. G. & Harbison, A.

1944 kirkwoodiana, Acteocina

Stoliczka, F.

1868 gabbiana, Bullaea

Whitfield, R. P.

1892 forbesiana, Actaeon

1982 gabbana, Actaeon

1982 subovoides, Actaeon

Acknowledgements

We are deeply grateful to Francisco Borrero and Paul Callomon (ANSP) for helping with images; to Gary Rosenberg (ANSP) for helping with nomenclature; to John Sime (ANSP) for helping with curatorial matters; to Cathy Buckwalter and Alexandria Capone (ANSP) for helping in tracking down some of the old literature; and to Paul Callomon (ANSP), Alexander Nützel (Bayerische Staatssammlung für Paläontologie und Geologie, Germany) and Luiz R. L. Simone (Museu de Zoologia da Universidade de São Paulo, Brazil) for the helpful comments that greatly improved this article. The comprehensive coverage of crucial literature associated with this study was only possible due to initiatives facilitating online open-access to older scientific works, most notably the Biodiversity Heritage Library and the Internet Archive projects. This work was partly supported by a doctoral grant from CNPq (Conselho Nacional de Desenvolvimento Científico e Tecnológico, Brazil) to R.B.S. (proc. #245575/2012-0) and a post-doc grant from CAPES (Coordenação de Aperfeiçoamento de Pessoal de Nível Superior, Brazil) to C.M.C. (proc. #8739/13-7).

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