South side of Castle Harbor, opposite Tucker’s Town, Bermuda.

Holotype (monotypy), ANSP A7030, 1 specimen (dehydrated but subsequently put back on alcohol, 33.2 mm long preserved); ANSP 66519, 1 shell (35 mm) (A. Heilprin coll. 1888), removed from body of holotype by Heilprin.

Heilprin (1888) stated that he examined a single specimen. We do not illustrate the rehydrated alcohol specimen as it is a blackened mass without visible distinguishing features.

Aplysia dactylomela Rang, 1828 (fide Valdés et al. 2013).

Figure 1. 

Aplysiida types. A. Holotype of Aplysia aequorea (=A. dactylomela), shell, L 35 mm, ANSP 66519 in dorsal view. B. Same, in ventral view. C–G. Types of A. badistes, C. Shell, L 18.7 mm, ANSP 187712 in dorsal view. D. Same, in ventral view. E. Same species, whole specimen, L 45.8mm, ANSP A7028 in dorsal view. F. Same, in ventral view. G. Same, in right lateral view. H–J. Syntypes of A. benedicti (=Aplysia argus), whole specimens ANSP A7027. H. Specimen L 79 mm in dorsal view. I. Same, in right lateral view. J. Specimen L 76 mm in right lateral view.

Venetian Causeway, Biscayne Bay, Florida, USA.

Holotype and paratype: ANSP A7028, 2 specimens (one whole except for removal of buccal mass, measuring 45.8 mm long preserved and the other dissected, measuring 46.7 mm long preserved); ANSP 187712, 1 shell (18.7 mm) and gizzard plates of dissected specimen of A7028 (H. A. Pilsbry coll., iv/1951), 1 slide with radulas of dissected specimens.

Aplysia badistes Pilsbry, 1951 is the type species of subgenus Metaplysia Pilsbry, 1951. Pilsbry stated “Type and another specimen are 187712 ANSP”. Pilsbry described the internal anatomy from one specimen but extracted the radula from both. Someone separated the shell and gizzard plates as ANSP 188279, and labeled them “Paratype”, a term not used in the publication. We have restored them to the original catalogue number (187712) as it is not possible to tell which specimen Pilsbry intended to be the holotype.

A. juliana Quoy & Gaimard, 1832 (fide Eales 1960).

Apia Harbor, Upolu, Samoa.

Syntypes: ANSP A7027, 2 specimens + 1 slide with radulae and jaw mounted (leg., C. N. E. Eliot, 19–21/vii/1899) [1 specimen (Eliot 1900: fig 2a), preserved 76 mm long, whole, not dissected; another specimen, preserved 79 mm long, with radula removed].

Aplysia argus Rüppell & Leuckart, 1830 (fide Alexander & Valdes, 2013).

Sandwich Islands [Hawaii, USA].

Potential syntypes: ANSP 20686, 1 shell + 2 fragments (11.3 mm long, “ex auct”), ANSP 20687, 3 shells (7.1, 7.2 and 7.5 mm long, “ex auct”), MCZ 31442 and 298486 (Johnson 1994).

It is not known whether Pease collected the specimens deposited at ANSP and MCZ before or after the species description was published. Therefore, we consider them potential syntypes.

Aplysia parvula Mörch, 1863 (fide Eales 1960).

Figure 2. 

Aplysiida types. A–B. Syntype of Syphonota elongata (=Aplysia parvula), A. Shell, L 11.3 mm, ANSP 20686 in dorsal view. B. Same, in ventral view. C–E. Syntypes of Tethys floridensis (=A. fasciata), C. Shell, L ca 45 mm, ANSP 63529 in dorsal view. D. Same species, whole specimen, L ca 90 mm, ANSP A7034 in right lateral view. E. Same, in dorsal view. F. Syntype of Dolabrifera fusca (= D. dolabrifera), shell, L 7.7, ANSP 20710 in dorsal view. G. Same, in ventral view. H–K. Syntypes of D. jacksoniensis (= D. brazieri), H. Shell, L 6.7 mm, ANSP 64931 in dorsal view. I. Same, in ventral view. J. Same species, whole specimen, L ca 16.7 mm, ANSP A7040 in dorsal view. K. Same in ventral view.

Key West, Florida, USA.

Syntypes: ANSP A7034, 2 specimens (one 90 mm long preserved, with shell, not dissected; other 76 mm long preserved, shell removed, penis everted) (H. C. Machette, leg. 1893) [1 specimen (Fig. 2C, D), penis not everted; another specimen, penis everted (Fig. 2E)]; ANSP 63529, 1 shell (fragmented, ca 45 mm long).

Pilsbry (1895) stated that “the types are two well preserved alcohol specimens”. We judge the original drawing of a whole animal (Pilsbry 1896a: pl. 37, fig. 19) to be a composite of the two specimens.

Aplysia fasciata Poiret, 1789 (fide Medina et al. 2005).

Polynesia.

Syntype: ANSP 20710 (Pease 1868b: pl. 12, fig. 27; Valdés et al 2017: fig. 5I, J), 1 shell (7.7 mm long) “ex auct”, no locality data. Syntypes, MCZ 297870, 3 shells, from Tahiti (Johnson 1994).

Pease (1868a) neglected to state a locality within Polynesia, although he described the habitat as “under stones, in the upper region of the laminarian zone”.

Dolabrifera dolabrifera Rang, 1828 (fide Valdés et al. 2017).

Port Jackson, New South Wales, Australia.

Syntypes: ANSP A7040, 2 specimens (one 16.7 and another 11.3 mm long preserved); ANSP 64931 (Pilsbry 1896b: pl. 44, figs 38–41; Valdés et al. 2017: fig. 8E, F), 1 shell (6.7 mm long; ex J. C. Cox 1893).

Both alcohol specimens have had the shell removed, but only one shell is present in the corresponding dry lot.

Dolabrifera brazieri G. B. Sowerby II, 1870 (fide Valdés et al. 2017).

San Juan del Sur, Nicaragua.

Syntypes (all leg. J. E. Bransford): ANSP A7048, 2 specimens (both dry, one ca 25.5 mm long preserved with shell 7.8 mm long and another ca 24.6 mm long preserved, shell removed); ANSP 67517 (Pilsbry, 1896b: figs 12–14; Valdés et al 2017: fig. 15A, B), 1 shell (9.2 mm long), taken from body in A7048; ANSP 67518 (Pilsbry 1896b: fig. 15), 1 shell (8.2 mm long), corresponding alcohol specimen missing.

Pilsbry (1896a) gave measurements for two specimens preserved in ethanol, but three shells and two bodies are present in the ANSP collection. The ANSP ledger lists 63517 from “San Juan Del Sur Nicaragua” (the type locality) as containing two alcohol specimens and one dry and 63518 from “Nicaragua” as having one alcohol and one dry. These were catalogued on 21 January 1896 and so are part of the material that Pilsbry examined as the description was published on 13 March 1896. The ledger lists 63517 as “Types” but does not list 63518 as a “type”. The original ANSP label for 63517 says “Shell of the TYPE”; whereas that for 63518 does not mention type status. Although Pilsbry intended 63517 to be the holotype, he did not designate a holotype in the original description, so we consider all of the material to be syntypes.

ANSP A7048 contains a note saying, “August 1969 badly dried” and the back of the most recent Academy label says “Treated with Tri-sodium phosphate and stepped up to 70% ETOH. Unfortunately, the sample subsequently dried out again, and so has been combined with the dry lot.

Dolabrifera nicaraguana Pilsbry, 1896 (fide Valdés et al. 2017).

Figure 3. 

Aplysiida types. A–E. Syntypes of Dolabrifera nicaraguana, A. Shell, L 8.2 mm, ANSP 67518 in dorsal view. B. Same, in ventral view. C. Same species, shell, L 9.2 mm, ANSP 67517 in dorsal view. D. Same in lateral view. E. Same in ventral view. F–I. Syntypes of Tethys panamensis (= A. dactylomela), F. whole specimen, L ca 52 mm, ANSP A7042 in right lateral view. G. Same in dorsal view. H. Shell, L 16 mm, ANSP 20693 in ventral view. I. Same in dorsal view.

Panama.

Syntype: ANSP A7042 (Pilsbry 1896b: fig. 45), 1 specimen (52 mm long preserved), ANSP 20693 (Pilsbry, 1896b: figs 46–48), 1 shell (16 mm long; J. A. McNeil coll.).

Recently, the Caribbean species Aplysia dactylomela was genetically differentiated from Indo-Pacific ones (Alexander and Valdés 2013), while the East Pacific remains unsampled (Uribe et al. 2013). Aplysia panamenis (Pilsbry, 1895) is the representative synonym of this area.

Aplysia dactylomela Rang, 1828 (fide Eales 1960).

Cape May, New Jersey [USA].

Holotype (monotypy): ANSP A2250 (ca 105 mm long preserved, shell 60 mm long; H. Lemon coll., 25/x/1894). Pilsbry stated that he examined a single specimen.

Aplysiidans possess a large gland near the mantle floor, called the opaline gland (or gland of Bohadsch), that produces a milky liquid when the animal is disturbed (Gosliner 1994). The gland can be multiporous (e.g. A. depilans, A. juliana and A. parvula) or uniporous (e.g. A. dactylomela and A. fasciata) (Beeman 1968; Cunha and Simone 2019; pers obs.). The opaline gland in Aplysia perviridis is unique in having both conditions, uniporous and multiparous (figs 32, 33). Pilsbry (1895) described the opaline gland as “projecting externally as a pedunculated oval body”, but that appears to be the result of contraction upon fixation, as suggested by Pilsbry: “but perhaps evaginated, in which case it would have one large orifice”.

The opaline gland of a single young specimen collected in 1950 from Sea Isle City, New Jersey, which is about 35 km north of the type locality was dissected to confirm this condition (ANSP A2235, ~60 mm long preserved). Despite the lack of a “pedunculated oval body” the condition of gland is the same (Fig. 4B, C). The multiporous part is more developed in the holotype than in the young specimen.

Aplysia perviridis (Pilsbry, 1895) was accepted as a synonym of A. willcoxi Heilprin, 1886 by Eales (1960). However, with the anatomical differences of the opaline gland, we treat it as a valid species herein.

Aplysia perviridis (Pilsbry, 1895) comb. nov.

Figure 4. 

Aplysiida types. A–E. Holotype of Tethys willcoxi var. perviridis (=A. perviridis), A. Whole specimen, L. ca 105 mm, ANSP A2250 in right lateral view. B. Same, detail of opaline gland in dorsal view showing the uniporous and multiparous apertures. C. Same, opaline deflected showing the opaline gland spread on the roof of haemocoel. D. Shell of the same, L 60 mm, in ventral view. E. Same in dorsal view. F–H. Syntype of Tethys pilsbryi (= A. pilsbryi). F. Whole specimen, L 97 mm, ANSP A7036 in right lateral view. G. Same in dorsal view. H. Same, detail of pallial cavity showing the upturned edge to the rim of the mantle border (ed); legend: pa - parapodium, sr - seminal groove, ha - hermaphrodite aperture, ed - upturned edge, ma - mantle; gi - gill, as - anal siphon. Scale bar: 5 mm.

Silam, north coast of Yucatan, Mexico.

Syntypes, ANSP A7036, 2 specimens (one 97 mm long preserved with shell and other 94 mm long preserved with shell removed, both dissected and radula removed) + 1 slide with radulae and jaw mounted (Heilprin Expedition, 1890) [one specimen (Letson 1898: figs 2–3), whole, lacking the left rhinophore (added in the figure), dissected ventrally, shell and radula not removed; other specimen, shell and radula removed].

Eales (1960) synonymized this species with A. cervina Dall & Simpson, 1901 because of general morphological similarity, but it differs in having an upturned edge to the rim of the mantle where the purple gland lies (Fig. 4H: ed). This remarkable characteristic is unique in the genus Aplysia. Letson (1898) also described the presence of “a few inconspicuous scattered warts” on the integument, not reported by Eales (1960). However, both features were confirmed in juvenile specimens from Yucatan (ANSP A1993, 10 specimens measuring 25–50 mm). Additionally, A. pilsbryi has a shorter shell (ca 1.2 times long as wide), while the shell of A. cervina is more elongated (ca 1.6 times long as wide) (MacFarland 1909).

Aplysia pilsbryi (Letson, 1898). Kobelt (1897: 240) introduced the combination to the genus Aplysia.

West coast of Mexico.

Holotype (original designation, “the type”): ANSP A7029, 1 specimen (ca 92 mm long preserved; W.H. Jones, leg.) [shell removed and figured by Pilsbry (1896b: pl. 55, fig. 5) but not found, should be ANSP 66307].

Pilsbry (1895) cited plate 55, figures 4 and 5, but when the plate was published the next year, the captions showed figures 5 and 6, the latter being correct. The specimen has the sole covered by small black dots (Fig. 5D), which is not mentioned in the original description nor by Eales (1960).

Aplysia robertsi (Pilsbry, 1895) (fide Eales 1960).

Figure 5. 

Aplysiida types. A–D. Holotype of Tethys robertsi (= A. robertsi), A. Whole specimen, L ca 92 mm, ANSP A7029 in dorsal view. B. Same in right lateral view. C. Same in ventral view. D. Same, detail showing the sole covered by small black dots. E–G. Syntype of Dolabrifera swiftii (=D. dolabrifera), D. Shell, L 10.5 mm, ANSP 67519 in dorsal view. F. Same in ventral view. G. Same in lateral view. H–L. Syntype of Tethys pulmonica var. tryoniana (=A. pulmonica), H. Whole specimen, L ca 122 mm, ANSP A7037 in right lateral view. I. Same in dorsal view. J. Same in ventral view. K. Shell, L ca 58 mm, ANSP 66306, in dorsal view. L. Same in ventral view. Scale bar: 5 mm

West Indies.

Syntype: ANSP 67519 (Valdés et al 2017: fig. 12K, L), 1 shell (10.5 mm long, donated by Robert Swift).

In the plate caption, which was published later than the description (September 23 vs March 13), Pilsbry suggested the name may be a synonym of D. ascifera Rang.

Dolabrifera ascifera (Rang, 1828) (fide Valdés et al. 2017).

Upolu, Samoa Island.

Syntype, ANSP A7037 (Pilsbry, 1896b: pl. 57, fig. 27), 1 specimen (ca122 mm long preserved), ANSP 66306 (Pilsbry, 1896b: pl. 57, figs 24–26), 1 shell (fragmented, ca 58 mm long; ex G. W. Tryon from Dr Gräffe, Museum Godeffroy).

In 1895 Pilsbry cited plate 57, figures 54–57, but when the plate was published the next year, the captions showed figures 24–27, the latter being correct. The specimen was purchased by G. W. Tryon from the Godeffroy Museum Catalogue. Pilsbry cited “Aplysia sp.? Museum Godeffroy, Catalog IV, p. 105, No. 1107a”, but apparently examined only one specimen. Pilsbry did not dissect the specimen but removed the shell.

According to Pilsbry (1895), A. pulmonica and Tethys pulmonica var. tryoniana (both from Samoa) are similar, differing just in that A. pulmonica lacks black maculations and has a more extensive posterior junction of the parapodia. We follow Eales (1960), keeping Tethys pulmonica var. tryoniana as a synonym of A. pulmonica until more material is available.

According to a recent study by Alexander and Valdes (2013), suggesting that “the ring doesn’t mean a thing”, A. pulmonica Pease, 1860 is a synonym of A. argus Rüppell & Leuckart, 1830. The photographs of Hawaiian specimens identified as A. dactylomela and A. pulmonica in their paper (Alexander and Valdes 2013: fig. 2) suggest that the material of A. pulmonica has rings in the color pattern, although it is supposed to lack them. Photographs of live specimens of the main material used in that publication (A. dactylomela, CPIC 00297 and A. pulmonica, CPIC 00313) confirm that both possess black rings in the body color. The black rings are more evident in the material they identified as A. dactylomela (paler) than in the ones they identified as A. pulmonica (darker).

Their Hawaiian material does belong to a single species, as they concluded based on their molecular data. The dark morph is consistent with the description of A. grandis Pease, 1860 (type locality Hawaii), which was not cited by the authors, in having a purplish-brown color, pale along the flanks, everywhere above densely crowded with minute white dots, and foot projecting posteriorly, where it is rounded. The dark morph is not consistent with the original description and illustration of A. pulmonica, which differs in being coarsely reticulated with black veins, lacking white dots, having oral tentacles short, and the posterior junction of the parapodia producing a sac-like structure. Bebbington (1977) included A. grandis in the synonymy of A. dactylomela, and not of A. pulmonica. In synonymizing A. pulmonica with A. argus, Alexander and Valdes (2013) ignored that most previous authors regarded A. dactylomela and A. pulmonica in the Indo-Pacific as distinct species. We regard their use of A. pulmonica for their Hawaiian material as a misidentification, and resume use of A. pulmonica as a valid species.

Little Gasparilla Bay, west coast of Florida, USA.

Syntypes (ex Heilprin 1887): A2262, 5 specimens with radula associated and 3 loose shells; ANSP 64260, SEM stub with fragment of radula.

The ledger entry for ANSP 64260 says six specimens in alcohol. Heilprin (1887b: 20) noted that a half dozen specimens were collected with dip-nets by Joseph Willcox and the expedition cook (Moses Natteal, p. iii), so one syntype is currently missing. The alcohol lot was subsequently renumbered as A2262, but we retained the dry number for an SEM stub with a radular fragment. All specimens were previously dissected, with the radula having been removed and kept with the specimens.

ANSP 63628 contains shells (and fragments) of about seven individuals collected by Willcox, from Marco, Florida donated by Joseph Leidy. We do not consider these parts of the type series as the locality was not mentioned in the original description.

Aplysia fasciata Poiret, 1789 (fide Medina et al. 2005; Rosenberg et al. 2009).

Figure 6. 

Aplysiida types. A–D. Syntype of Aplysia willcoxi (=A. fasciata). A. Whole specimen, L = ca 114 mm, ANSP A2262, in dorsal view. B. Same in right lateral view. C. Shell in ventral view, L = 50 mm. D. Same, in dorsal view.