Parvanachis Radwin 1968

Parvanachis Radwin, 1968 Anachis (Parvanachis) Radwin, 1968: 145 –147 (type by original designation Buccinum obesum C.B. Adams, 1845). Diagnosis. Shell small (3–8 mm long), globose fusiform, with dominant axial ridges and underlying spiral grooves. Spire generally flat-sided with incised sutures. Aperture edge thickened and denticulate, with a shallow posterior notch anterior to the suture. The radula is typical for columbellids, with sigmoid lateral teeth, having a triangular basal secondary lateral cusp with a strong hook toward the radular membrane. The female gonoduct has a short thin-walled vestibule, lacks a bursa copulatrix, has a single gland mass, and has a coiled gonopericardial duct and gonopericardial-pallial duct, both encased in muscle. The male gonoduct is typical for columbellids, but lacks the secondary seminal vesicle found in many species. The penis is simple in form with a long or short filament-like tip. Remarks. Parvanachis has been usually considered either as a genus or as a subgenus of Anachis since its description. Keen (1971) still gave it subgeneric status. Radwin (1977 a, 1977 b) raised Parvanachis to a full genus in his papers on Western Atlantic columbellids, though was apparently thinking about this earlier, as he lists Parvanachis as a genus in a Festivus paper in 1974 but does not discuss it. His justification (1977 b) was based on Parvanachis’ distinct shell and radular morphology. Lopes et al. (1971) referred Anachis obesa to Zafra A. Adams, 1860, an Indo-Pacific taxon, and Faber (2004) similarly regarded Parvanachis as a subgenus of Zafra.Published as part of Maintenon, Marta J., 2014, Taxonomic revision of the species of Parvanachis Radwin, 1968 (Gastropoda: Columbellidae) from the Gulf of Panama, pp. 201-225 in Zootaxa 3753 (3) on page 209, DOI: 10.11646/zootaxa.3753.3.1, http://zenodo.org/record/25331

Parvanachis diminuta C.B Adams 1852

Parvanachis diminuta (C.B Adams, 1852) Figures 1 B, 5 A, 5 D Columbella diminuta C.B. Adams, 1852: p. 85, not figured. Reeve, 1858: sp. 115, pl. 20 fig. 115. Turner, 1956: p. 45, pl. 7 fig. 2. Mitrella (Astyris) diminuta Adams. Mörch, 1860: p. 94. Anachis rufotincta Carpenter, 1857: pp. 511–512, not figured. Anachis diminuta. Carpenter (1863): p. 344, not figured. Strong & Hertlein, 1939: p. 184, not figured. Keen, 1958: p. 381, sp. 429 (illustration from Turner, 1956). Columbella (Seminella) diminuta (C.B. Adams). Tryon (1883): p. 177, pl. 58 fig. 44. Columbella (Anachis) diminuta C.B. Adams. Kobelt, 1897: p. 163, taf. 22, figs. 13, 14. Anachis (Parvanachis) diminuta (C.B. Adams, 1852). Keen, 1971: p. 584, sp. 1204 (illustration from Turner, 1956). Abbott, 1974: p. 197, sp. 2064 (not figured). Parvanachis diminuta (C.B. Adams, 1852). Skoglund, 1992: p. 89. Types. Lectotype, MCZ 186401, from Panama. A photo of the type was available from W. Thorsson to confirm ID. Taxonomic history. Charles Baker Adams did not illustrate this species when he described it; Reeve illustrated it in 1858. In both cases its shell was described as pale, with a dark base, which is characteristic of this species, and its identity has not been in question. Carpenter (1863) considered it a species of Anachis (which he allied with Pisania based on the operculum) while Tryon (1883) placed it in Columbella (Seminella) and based his illustration on that of Reeve. Anachis rufotincta (with a more reddish anterior end) was named by Carpenter (1857) from Mazatlan, and as reported by Tryon (1883) and later authors, is a synonym. Diagnosis. Small species with a biconic axially ribbed shell, mottled pale tan with a narrow white spiral below the suture and a dark reddish or purple-brown anterior tip. Material. Over 100 specimens of this species were collected on the bottoms of rocks in muddy sand. Two were sectioned, two dissected. USNM has dry specimens from Baja California Sur to Panama. Shell (Fig. 1 B): Shell small, biconic, 3.5 to 4.3 mm long (avg. 3.84 mm) and 1.55 to 1.95 mm wide (avg. 1.70 mm) in 12 specimens measured. Adult shells with 4 to 4.25 teleoconch whorls (avg. 4.00). Protoconch smooth, off white, 3.25 to 3.5 whorls (avg. 3.14). Dominant sculpture of axial ridges, except on first two teleoconch whorls where strong spiral ridges make the sculpture cancellate. One weak subsutural groove cuts across tops of axial ridges. Shell off-white with vague chestnut blotches that give an overall mottled tan affect, a dark purplish brown base and usually a narrow white band slightly below the suture. Aperture edge thickened, with a few denticles internally. Shallow posterior sinus present in aperture edge. Parietal wall denticulate, anterior edge of callus detached. Aperture white, with purple flush on anterior columellar wall. Body coloration. Body cream colored with white specks overall. Operculum (Fig. 5 A, tip broken): Operculum ovoid with a terminal nucleus, darker centrally, muscle scar bilobed, no keel. Radula (Fig. 5 D): One radula obtained whole had 123 tooth rows. Lateral teeth in adult specimens dissected 18 to 20 µm long, 7 µm wide. Lateral teeth with three pointed secondary cusps, the basal cusp pointed and curved toward the membrane, and separated from the distal two by a wide gap. Reproductive anatomy: Male system typical for Parvanachis. Ciliated duct to mantle cavity present anterior to coiled seminal vesicle. Anterior spermiduct ciliated, moderately muscular, passes forward embedded in body wall from mantle cavity duct to penis base. Long spermiduct loop in hemocoel leaves body wall near penis base and runs back alongside proboscis, then doubles back and enters penis. Spermiduct of internal loop wide, with clustery mucoidal epithelium that continues along most of the length of the penis except for a non-secretory region at the base. Penis tip with subepithelial mucus glands in addition to secretory epithelium. Penis long, relatively thick, simple with a short filament tip. Penis curled back in the posterior mantle cavity at rest. Female system typical for Parvanachis, including a single gland mass, with muscular gonopericardial duct and no bursa copulatrix. Gland mass stains red (eosinophilic) in the middle, and pale purple (with haematoxylin) on both ends. The female sectioned had an egg capsule in the vestibule, which was otherwise short and thin-walled. Gonopericardial duct wide and coiled, epithelium cuboidal with large basal nuclei; duct has a wide, very thickwalled medial region (Figs. 3 F, G) with a large mass of red-stained material on one side (Fig. 3 G), which might be partially dissolved sperm. The epithelium in that region is different than the rest, and the epithelial characteristics are not readily discernible, but the tissue appears to be secretory. Gonopericardial-pallial duct (gpp) splits off in this same region (Fig. 3 F). Pericardium in the specimen sectioned with a large amount of unoriented sperm.Published as part of Maintenon, Marta J., 2014, Taxonomic revision of the species of Parvanachis Radwin, 1968 (Gastropoda: Columbellidae) from the Gulf of Panama, pp. 201-225 in Zootaxa 3753 (3) on pages 209-211, DOI: 10.11646/zootaxa.3753.3.1, http://zenodo.org/record/25331

Parvanachis pygmaea Sowerby I 1832

Parvanachis pygmaea (Sowerby I, 1832) Figure 6 A–D, 7 A, 7 E Columbella pygmaea Sowerby, 1832: p. 119, not figured. 1844 a: p. 141, pl. 40 fig. 163. Duclos in Chenu, 1848: pl. 25 figs. 1– 2. Adams, C.B., 1852: p. 97. Reeve, 1858: sp. 128, pl. 22, figs. 128–129. Anachis pygmaea (Sowerby, 1832). Carpenter, 1857: p. 510; 1863: p. 344. Baker, Hanna & Strong, 1938: p. 250, not figured. Strong & Hertlein, 1939: p. 184, not figured. Keen, 1958, p. 384, sp. 448 (illustration from Sowerby, Thes. Conch.). Columbella (Seminella) pygmaea (Sowerby). Tryon, 1883: p. 166, pl. 56 figs. 91, 92. Columbella (Anachis) pygmaea Sowerby. Kobelt, 1897: p. 158, taf. 22, figs. 1, 2. Anachis dalli Bartsch, 1931: pp. 2–3, pl. 1 fig. 2. Anachis (Parvanachis) pygmaea (Sowerby, 1832). Keen, 1971: p. 585, sp. 1209. Abbott, 1974: p. 197, sp. 2068 (not figured). Parvanachis pygmaea (Sowerby, 1832). Skoglund, 1992: p. 89. Types. Four probable syntypes marked by Keen in 1964, NHMUK 1966389, collected by Cuming at Monte Cristo (not St. Elena), Ecuador (in Guayaquil province, according to C.B. Adams, 1852) on dead shells. Taxonomic history. Sowerby (1832) reports Cuming collecting this species on dead shells in sandy mud at Santa Elena, at 10 fms. Adams (1852) reports finding them under stones at the low water mark. Keen (1971) figures one of Cuming’s specimens from Monte Cristo, as well as Sowerby’s illustration; no material from St. Elena was found at NHMUK. Sowerby’s (1844) illustration is not specific enough to match a particular specimen, but is consistent with the Monte Cristo specimens. Carpenter (1857) refers to Anachis pygmaea as being variable in color, from ‘light horn with a few purple-brown patches to nearly uniform black brown’; he probably has this species confused with P. adamsi (described below), which is similar in size and whose markings sometimes resemble those of P. pygmaea. In 1864 he refers to his and Adams’ material as Anachis pygmaea var. auriflua, but no formal description or illustration has been found to determine an identity for that name, so it must remain a nomen nudum. One lot of Carpenter’s specimens (USNM 716231, Carpenter Tablet 2429), is cited in Carpenter (1857) as showing P. pygmaea in different stages of growth, but is not this species; however it has fewer specimens than reported (6 juveniles rather than 7 juveniles and 3 adults) so perhaps the lot was split. Bartsch (1931) named Anachis dalli (holotype USNM 368144) from Taboga Island, Panama. Photos of the likely syntypes of C. pygmaea and the type specimen of A. dalli were both examined, and though Bartsch briefly notes that his species was previously misidentified as A. pygmaea, he does not specify how they differ, and no differences between the types are apparent. Diagnosis. Small (4 to 7 mm) species with a biconic, axially ribbed shell, straw colored with a white band immediately below the suture and a spiral row of dark markings overlying or slightly below the white band. A second band of dark markings is present below the periphery. Material. Parvanachis pygmaea was common at most localities (6 of 11, Table 1) though not in high numbers. 45 specimens were collected on and under rocks in muddy sand or sand; most were large juveniles or adults. USNM has these from Baja California to Ecuador. Three were sectioned, three dissected. Shell (Figs. 6 A–D): Shell biconic, 4.2 to 6.8 mm long (avg. 5.41 mm) and 2.05 to 2.85 mm wide (avg. 2.40 mm) in 26 specimens measured. Adult shells with 4.5 to 5.5 teleoconch whorls (avg. 4.8). Protoconch smooth, offwhite, with 3 to 3.25 (avg. 3.17 in eight specimens) whorls. Shell sculptured primarily with axial ridges (15 to 18 on the penultimate whorl), weak, evenly spaced spiral grooves between axial ridges on the spire whorls. No welldeveloped subsutural cord except on the first two whorls, unlike P. adamsi, which is the same size and very similar otherwise. Apex usually eroded, unlike similar species. Shell easily recognized by its striking and relatively invariant color pattern. Shell glossy white or yellowish, with a row of purplish brown blotches on the anterior half of each whorl, usually strongest on every other axial ridge, and a white band between the dark spots and the suture. Body whorl with a second row of axially elongated dark blotches below the periphery. Aperture edge thickened, with a few denticles internally. Aperture edge with a shallow posterior sinus. Parietal callus with a weakly denticulate ridge. Aperture color reflects that of the shell exterior, anterior end of shell paler. Body coloration: Body cream colored, with dense black mottling and white speckles. Front corners of the foot without markings. Siphon mottled, with a black band; tentacles cream colored, with black bands on the base and middle. Operculum (Fig. 7 A): Operculum oblong, with a keel and bilobed muscle scar, and usually a terminal nucleus (one photographed appeared to be damaged, nucleus was sub-terminal). Operculum center is darker tan. Radula (Fig. 7 E): Radula similar to those of P. adamsi (Fig. 7 G) and P. nigricans (Fig. 7 H). Lateral teeth about 40 µm long in adults, with three pointed secondary cusps; the basal cusp is pointed down and embedded in radular membranes. Center plates are rectangular, about 30 to 40 µm wide by 10 to 25 µm deep. The three specimens (two adult males, one adult female) dissected had radulae with 125 to 132 tooth rows. Reproductive anatomy: Spermiduct typical for Parvanachis, with the addition of a flap-like projection at the penis near the base and a penial pouch. Coiled seminal vesicle with a long posterior portion having normal darkstaining sperm, anterior portion with mixed dark-staining and pink-staining sperm cells. Epithelia in both portions flat. Short mantle cavity duct present adjacent to kidney. Spermiduct from mantle Cavity duct to penis base encased in body wall muscle. At penis base, spermiduct leaves body wall and forms a long loop adjacent to the proboscis. Spermiduct loop epithelium (Fig. 4 C) tall and mucoidal, becomes non-secretory at penis base. Penis long and narrow, with a short filament tip and a flap of tissue protruding to one side near the base (Fig. 4 F). A semilongitudinal section of the penis where it narrows is shown in Fig. 4 F; at this point the duct also narrows and becomes non-secretory. Penial spermiduct straight. Tissue flap at base does not involve duct or any modified epithelia, so ostensibly has a more mechanical function. Distal portion of penis at rest stored in a pouch in the mantle cavity roof. Female reproductive system typical for Parvanachis; bursa copulatrix absent, coiled gonopericardial duct present. Glands comprise a single mass, which stains red anteriorly and darker purple posteriorly, with a pale purple band across the middle closer to the posterior end. Gonopericardial duct splits from oviduct just inside posterior end of gland mass. Gonopericardial duct muscular and coiled on the oviduct side, with a tall secretory epithelium; then widens to become almost as wide as the glandular mass itself, with a pouched, mucoid wall (Figs. 3 C–E). Duct narrows before entering the pericardium, which in the specimen sectioned was full of sperm and cellular material. Gonopericardial-pallial duct splits from pericardium adjacent to where gonopericardial duct enters. Vestibule short, non-muscular. The specimen sectioned had an egg capsule in the vestibule, so details of the vestibule epithelium and form could not be ascertained with certainty. Remarks. The 12 year gap between description of this species and its illustration, combined with lack of specimens from the reported type locality and the obvious similarity between several of these species, creates some uncertainty in this species’ original identity. It is possible, given the similarity between P. pygmaea, P. mullineri and P. adamsi (discussed below), to suggest that the unillustrated Columbella pygmaea Sowerby, 1832, is actually what I identify here as P. mullineri or P. adamsi. Reeve’s (1858) figure, interestingly enough, shows the dark band adjacent to the suture, consistent with P. mullineri. However, Sowerby’s illustration from 1844, consistent with the existing syntypes, sets the identity of the species.Published as part of Maintenon, Marta J., 2014, Taxonomic revision of the species of Parvanachis Radwin, 1968 (Gastropoda: Columbellidae) from the Gulf of Panama, pp. 201-225 in Zootaxa 3753 (3) on pages 213-217, DOI: 10.11646/zootaxa.3753.3.1, http://zenodo.org/record/25331

Parvanachis dichroma Maintenon, 2014, n. sp.

Parvanachis dichroma n. sp. (Figures 1 C, D, 5 B, 5 E) Types. Holotype, Fig. 1 C, (USNM 1231302), Chumical, Panama, 29 Jan. 2006, 4.43 mm long. 59 paratypes, USNM 1231303–1231306, see Table 1. Etymology. named to note the strong contrast between the white shell and its brown markings. Diagnosis. Small species with a biconic axially ribbed shell, bright white with two spiral rows of chestnut markings and a reddish-chestnut anterior tip. Material. This species was found intertidally on and under rocks in a habitat with fine, pale muddy sand. About 60 were collected; they occurred more densely than similar species where they were found and tended not to be mixed with other small columbellid species. Three adults were sectioned (one male, two females with one female cross sectioned, one longitudinally sectioned), three males dissected. Shell (Figs. 1 C, D): Shell small, biconic, 4.1 to 5.1 mm long (avg. 4.50 mm) and 2.00 to 2.20 mm wide (avg. 2.05 mm) in 15 specimens measured. Adult shells with 4 to 5 teleoconch whorls (avg. 4.3). Protoconch smooth, white, with 3 to 3.25 whorls and no real sinus. Predominant sculpture consists of rounded, slightly sinuous axial ridges (16 to 19 on the body whorl), with spiral grooves between ridges on the spire whorls and often on the body whorl. A weak subsutural groove cuts across the tops of the axial ridges. Shell translucent, white. Spire whorls have a single band of wavy axial chestnut makings in the middle of the whorl, with an additional row present on the basal half of the body whorl. These form two large blobs on the outer lip. Tip of siphonal canal in adults dark purplish brown. Aperture white. Aperture edge thickened, denticulate, with a shallow posterior notch. Parietal wall with a ridge of denticles. Body coloration: Anterior part of body cream colored except for white specks overall. Behind the mantle edge, preserved specimens are black, which can easily be seen through the shell; this may be an artifact of preservation. Operculum (Fig. 5 B): Operculum ovoid, with bilobed scar, no keel. Nucleus terminal, with a darker spot in the middle. Radula (Fig. 5 E): In four adult male specimens dissected, lateral teeth 28 to 34 µm long by 12 to 15 µm wide, with rectangular center plates. Two radulae that were complete had 137 and 155 tooth rows. Lateral teeth have three pointed secondary cusps, the bottom one pointed and curved down, and separated from the distal two by a wide gap. Reproductive anatomy: Male reproductive anatomy typical for the genus. Primary seminal vesicle coiled, contains both pink staining and typical dark staining sperm, with a low epithelium overall. Spermiduct gives off short mantle cavity duct adjacent to the kidney. Anterior spermiduct embedded in the body wall has a thin coat of circular muscle. Anterior spermiduct enters hemocoel near penis base and forms a long loop adjacent to the proboscis. The loop has a complex pale purple-staining mucoid epithelium that continues into the penis base, after which the duct becomes non-secretory. Penis tip densely supplied with subepithelial mucus cells. Penis long and very narrow, with a short filament tip (as in Fig. 2 B). It rests in the mantle cavity. Female reproductive anatomy typical for the genus, with a short glandular complex having an eosinophilic anterior portion and a pale purple-staining posterior portion (Fig. 3 A). Gland mass similar in dimensions from anterior to posterior. To the posterior right of the gland mass is a thick walled, coiled gonopericardial duct, part of which may function in sperm storage or absorption. Reproductive system has no bursa; vestibule short, with a simple, non-muscular wall. Remarks. This species closely resembles Parvanachis oxillia (Duclos in Chenu, 1846) in terms of the size and color pattern. The identity of the latter has been difficult to determine, because the Chenu publication contains only figures, and localities are not given for most species. The syntype lot of Columbella oxillia, MNHN 6414, is labeled with a locality of Panama, but it does not specify whether the material came from the Pacific or the Caribbean. A photograph of one of the syntypes (http://coldb.mnhn.fr/CatalogNumber/ MNHN /IM/ 6414) suggests that P. oxillia has a narrower form and somewhat more acute spire, with a more varicose lip and a relatively large posterior labial denticle. In this it more closely resembles P. obesa (see Fig. 1 A) than it does the current species, and Rosenberg (2009) lists it as a synonym of the Caribbean P. obesa.Published as part of Maintenon, Marta J., 2014, Taxonomic revision of the species of Parvanachis Radwin, 1968 (Gastropoda: Columbellidae) from the Gulf of Panama, pp. 201-225 in Zootaxa 3753 (3) on pages 211-212, DOI: 10.11646/zootaxa.3753.3.1, http://zenodo.org/record/25331

Parvanachis nigricans

Parvanachis nigricans (Sowerby, 1844 a) new comb. Figures 7 D, 7 H, 8 E–H Columbella nigricans Sowerby I, 1844 a: p. 143, pl. 40 fig. 172. Duclos in Chenu, 1848: pl. 21 figs. 7-8. Adams, C.B., 1852: pp. 95-96. Reeve, 1858: sp. 137, pl. 23, fig. 137. Mitrella (Astyris) nigricans Sowerby, 1844. Mörch, 1860: p. 94 (not figured). Anachis nigricans (Sowerby, 1844). Carpenter, 1863: p. 345. Strong & Hertlein, 1939: p. 184, not figured. Keen, 1958: p. 383, sp. 444 (illustration from Reeve, 1858). Abbott, 1974: p. 197, sp. 2067 (not figured). Columbella (Seminella) nigricans (Sowerby). Tryon, 1883: p. 168, pl. 57, fig. 2 Columbella (Seminella) nigricans Sowerby. Kobelt, 1897: p. 137, taf. 20, figs. 5, 6. Anachis (Costoanachis) nigricans (Sowerby, 1844). Keen, 1971: p. 581, sp. 1186. Costoanachis nigricans (Sowerby, 1844). Skoglund, 1992: p. 88. Types. Five syntypes, NHMUK 1966396, collected by Cuming in the Galapagos. Sowerby’s figure is not good enough to match to a specimen. Keen (1971) figures the syntypes along with Reeve’s illustration, however the photo quality is not good enough for specimen identification. Photos of the syntypes from K. Monsecour and the British Museum were used to help identify the species. Taxonomic history. Sowerby described this species from the Galapagos, as blackish with a whitish suture and pale aperture edge; Reeve (1858) echoes this. Adams (1852) recognized the extensive variation in color and pattern found in Panama shells, but insomuch as he reported having 120 mostly dark colored large specimens and 500 small ones with more variable coloration, he undoubtedly had both this and P. adamsi described above. Tryon (1883) described it as dark, but allowed for a lighter band on the body whorl. Keen (1971) also follows Sowerby’s lead in describing the species as dark brown to blackish. Diagnosis. Medium sized biconic axially ridged species 7 to 9 mm long, with variable coloration consisting of a tan to dark purple-brown base color, with oblique lighter markings and occasionally a white subsutural band. Sculpture denser and more strongly developed than other species. Material. This species was found intertidally on the sides of rocks, more commonly on offshore islands. USNM has material from Baja California to Panama. About 70 specimens were collected from five localities. Two adults were sectioned and three were dissected. Shell (Figs. 8 E–H): Shell medium-sized, biconic, 6.80 to 8.80 mm long (avg. 7.63 mm) and 2.65 to 3.80 mm wide (avg. 3.09 mm) in 19 specimens measured. Some adults from Isla San Pedro Gonzalez were smaller. Adults have 5.5 to 6.5 (avg. 5.8) teleoconch whorls. Protoconch similar in color to teleoconch but generally paler, with 3 to 3.5 (avg. 3.2 in 16 specimens) whorls. Primary sculpture of dense axial ridges (16 to 19 on the penultimate whorl) with spiral grooves between axial ridges and a well-developed subsutural groove. Shell coloration variable, tan or grayish to dark purple-brown, with oblique paler markings. Some with a spiral subsutural white band and/ or a spiral band of whitish blotches below periphery. Aperture reflects outer color, with labial denticles anterior to the posterior notch. Parietal ridge with denticles located over the spirals underneath. Body coloration: Body and foot cream colored, with black mottling and white dots overall. Head dark, cephalic tentacles pale, with black bands at middle. Siphon with a black ring near tip and black specks closer to body. Body under shell mostly pale, with a dark mantle edge. Operculum (Fig. 7 D): Operculum oblong oval, with terminal nucleus, keel and bilobed scar. Pigmentation similar overall. Radula: Radula (Fig. 7 H) narrow, with lateral teeth about 50 µm long. Center plates only slightly wider than deep. Each lateral tooth with three evenly spaced secondary cusps. Basal cusp pointed but embedded in membranes. Radulae in two male specimens had 148 and 153 tooth rows. Reproductive anatomy: Male anatomy similar to that of P. adamsi. Seminal vesicle with a low epithelium, contains both pink and purple staining sperm (Fig. 4 B). Wide, densely ciliated mantle cavity duct present at back of body cavity. Spermiduct narrows on entering the body wall; body wall spermiduct with a thick muscle coat and low, non-secretory epithelium. Anterior spermiduct with a single long loop in the body cavity adjacent to the proboscis. Anterior spermiduct loop with secretory reddish and pale purple-staining complex columnar epithelium (Fig. 4 D). Penis long and narrow (Fig. 2 D), with a tissue flap at the base (Fig. 4 G) consisting of connective tissue and mucus cells, similar to the rest of the penis wall. Spermiduct in penis base sinuous, with a low, nonsecretory epithelium (Fig. 4 F). Spermiduct in middle region of penis wide and thickly muscular, with mixed complex columnar and mucoidal secretory cells (Fig. 4 F). Distal third of penis narrows (Fig. 4 F) to very long filament tip, duct with non-secretory epithelium. Penis at rest curled into back of mantle cavity, lacking a penial pouch. Females with a gonopericardial duct but no bursa copulatrix. Gland mass stains red overall, with a purple staining band through middle toward the posterior end. Anterior end around lumen stains more purplish-red. Gland lumen pouched. Vestibule short, not muscular, with a smooth wall. Gonopericardial duct long and coiled, with a low epithelium close to the gland mass. Pericardial end of duct increases in diameter and becomes thicker walled and slightly folded; this portion may act in sperm digestion, however no sperm was present in the system of the female sectioned. Remarks. This species is similar to both P. adamsi and P. pygmaea ¸ but is larger and more strongly sculptured than both. It has traditionally been counted as a species of Costoanachis perhaps because it is slightly larger than other species of Parvanachis included here, but its shell characteristics, anatomy and radular morphology are more similar to the Parvanachis species herein than to any Anachis or Costoanachis species investigated. The following species has traditionally been considered a Parvanachis, but its anatomy is quite different, and more consistent with species of Anachis, as described above.Published as part of Maintenon, Marta J., 2014, Taxonomic revision of the species of Parvanachis Radwin, 1968 (Gastropoda: Columbellidae) from the Gulf of Panama, pp. 201-225 in Zootaxa 3753 (3) on pages 220-221, DOI: 10.11646/zootaxa.3753.3.1, http://zenodo.org/record/25331

Parvanachis mullineri Poorman 1983

Parvanachis mullineri (Poorman, 1983) Figures 6 E–H, 7 B, 7 F Anachis (Parvanachis) mullineri Poorman, 1983: pp. 8–9, figs. 4, 7. Types. Holotype, SDMNH 81612, from Bahia de Santiago, Colima, Mexico. Taxonomic history. This species has been mostly unrecognized since its relatively recent description. The specimens from Panama do not match it perfectly, but given the phenotypic variation in columbellids and the different localities, it seems likely that they are the same thing. The illustrated holotype appears very similar to the Panama specimen in Fig. 6 E. Poorman emphasizes the distinction of this species based on the varicose outer lip, pronounced anal sulcus and lamellose edge of the columellar callus. The latter two features are not particularly unusual in columbellids, but the varicose outer aperture edge is unusual, though it appears not to be present in all specimens. Diagnosis. Small (4 to 6 mm long) species with a biconic, axially ribbed shell, often with an unusually thick outer lip. Shell straw colored with a band of alternating white and dark blotches immediately below the suture and a wide unmarked band at the periphery. A second band of dark axial markings is present below the periphery. Material. A total of nine specimens were collected at Isla Venado and Punta Calafate, including five adults. In the Smithsonian’s dry collection, a few lots of these were found from locations ranging from southern Baja California to Ecuador. Because there were relatively few specimens and variation in the shell is important, these were not sectioned and the two specimens dissected were broken rather than dissolving the shell. Shell (Figs. 6 E–H): Shell biconic, 4.1 to 5.3 mm long (avg. 4.90 mm) and 1.8 to 2.5 mm wide (avg. 2.27 mm) in five adult specimens measured. Adult shells have 3.75 to 4.25 teleoconch whorls (avg. 4.15). Protoconch smooth, off white, with 3.5 to 3.75 (avg. 3.55 in six specimens) whorls and a narrow sinus. Shell sculptured primarily with axial ridges (15 to 19 on the body whorl), and spiral grooves between axial ridges on the spire whorls. Shells similar to P. pygmaea but differing slightly in pattern. Shell base color straw, with a wide band below the suture consisting of occasional white spots overlain on a dark band or blotches. The body whorl has a band of axially elongated dark blotches below the periphery and usually a separate row of spots on the base. The anterior tip of the shell is white, often with a spiral band of small dark markings. Aperture edge thickened, varicose in some specimens, with a few denticles internally and a couple large dark blobs on the dorsal aperture edge. Aperture edge with a shallow posterior sinus. Parietal callus with a weakly denticulate ridge. Aperture color reflects that of the shell exterior. Body coloration: Same as P. pygmaea. Operculum (Fig. 7 B): Operculum oblong, with a keel and a bilobed muscle scar and a terminal nucleus. Radula (Fig. 7 F): Radula similar to that of P. pygmaea. Lateral teeth about 38 µm long in two subadults dissected, with three pointed secondary cusps; the basal cusp is pointed down and embedded in radular membranes. Center plates are rectangular, 30 –35 µm wide by 7.5 µm deep. The subadults dissected had 131 and 139 tooth rows. Remarks. This species is very similar to Parvanachis pygmaea and was found in the same locations, though it was much less common. The shell is slightly smaller and stouter, with about one fewer teleoconch whorl and onehalf more protoconch whorls. Though there is overlap, these differences are both statistically significant; T = 3.70, P = 0.001, DF = 23 for the mean number of teleoconch whorls and T = 4.00, P = 0.002, DF = 12 for the mean number of protoconch whorls. P. mullineri also has less erosion at the apices, whereas the apex of P. pygmaea from the same locality typically is eroded; this may indicate a slight difference in habitat or shell chemistry. The color pattern in the Panama specimens of P. mullineri has the subsutural bands of white and dark markings mixed, with a wide unmarked spiral band below that, and typically two rows of markings below the periphery. P. pygmaea tends to have the two bands of markings mostly if not completely separated (dark anterior and white posterior), with a narrower unmarked band below. The labial edge in this species also tends to be thicker than in P. pygmaea, to the point of being varicose in some specimens, with large dark blotches on the dorsal aperture edge.Published as part of Maintenon, Marta J., 2014, Taxonomic revision of the species of Parvanachis Radwin, 1968 (Gastropoda: Columbellidae) from the Gulf of Panama, pp. 201-225 in Zootaxa 3753 (3) on page 217, DOI: 10.11646/zootaxa.3753.3.1, http://zenodo.org/record/25331

Parvanachis adamsi Maintenon, 2014, n. sp.

Parvanachis adamsi n. sp. Figures 7 C, 7 G, 8 A–D Types. Holotype, Fig. 8 A, (USNM 1231317), Chumical, Panama, 29 Jan. 2006, 5.93 mm long. 227 paratypes, USNM 1231318–1231323; see Table 1. Etymology. This species is named for Charles Baker Adams, who did extensive work on the Panamic fauna. Taxonomic history. C.B. Adams (1852) discusses under Columbella nigricans large typical specimens, and a number of smaller atypical specimens with a wide variation in color, that are almost certainly this species. Similarly, as mentioned above, Carpenter (1857) describes P. pygmaea as being quite variable in shell coloration with many essentially dark overall, which likely indicates that he had this new species mixed in with it. In 1864, he reported his previous findings of Anachis pygmaea in the Mazatlan collection, and Adams’ specimens of Columbella pygmaea, as being Anachis pygmaea var. auriflua, which he apparently intended to describe in the Annals and Magazine of Natural History; but this never occurred and no types were designated, thus that name is a nomen nudum and there is no way to be certain what he intended to use the name for. P. adamsi and P. nigricans especially are difficult to initially sort out without access to a large number of specimens. Diagnosis. Biconic axially ridged species 4 to 7 mm long, with variable coloration consisting of a straw to dark brown base color, usually with oblique dark markings and a white subsutural band. Axial ridges large, less dense than other species, with 14 to 16 on the penultimate whorl. Material. About 200 specimens were collected, in the middle to high intertidal zone, on the sides of rocks. The large numbers were collected due mostly to the species' high degree of variability. Two adult specimens were sectioned, and three dissected. Shell (Figs. 8 A–D): Shell biconic, 4.15 to 6.85 mm long (avg. 5.78 mm) and 1.93 to 2.95 mm wide (avg. 2.59 mm) in 24 adult specimens measured. Adults have 4 to 6 teleoconch whorls (avg. 4.9). Primary sculpture of heavy, well separated axial ridges with underlying spiral grooves best developed on the first spire whorl, mostly lacking on the anterior whorls. Protoconch off white, with 3 to 3.5 (avg. 3.25 in 11 specimens) whorls. Shell color and pattern varies; pale beige or yellowish to tan, usually with oblique brown or black lines, spots or blotches, and often a white spiral band below the suture (but note the specimen in Fig. 8 D is almost entirely without markings). Anterior tip of siphonal canal pale. Aperture similar in color to exterior, with labial denticles anterior to the posterior canal, weakly denticulate parietal ridge. Body coloration: Body cream colored with diffuse black mottling and white dots, and black spots on sole of foot. Siphon cream colored with black bands at the middle and tip, cephalic tentacles have medial and basal black bands Operculum (Fig. 7 C): Operculum ovoid, nucleus terminal. Keeled, bilobed muscle scar. Pigmentation slightly darker near keel. Radula (Fig. 7 G): Radula narrow, with 124 to 130 tooth rows in three adults dissected. Lateral teeth 38 µm long in both by 18 µm across the basal cusp on all three. Center plates rectangular, 33 µm wide and 13 µm deep in one specimen. Lateral teeth with three secondary cusps, gap between basal cusps slightly wider than that between distal cusps. Basal cusp broadly triangular, comes to a strong hooked point. Reproductive anatomy: Male reproductive system similar to that of P. nigricans. The coiled seminal vesicle (Fig. 4 A) contains pink-staining atypical sperm cells that are mixed with dark purple staining sperm cells, and the epithelium is flat overall. A wide, densely ciliated duct to the mantle cavity is present at the back of the anterior body cavity. The spermiduct anterior to the pallial duct enters the body wall and narrows, and the portion that runs along the body wall has a thick muscle coat and a low, nonsecretory epithelium. The anterior spermiduct makes a long loop in the body cavity adjacent to the proboscis. The spermiduct in the loop has a tall, pale purple-staining mucoidal epithelium with terminal nuclei. The spermiduct at the penis base is non-secretory, but toward the tip the duct becomes wide with subepithelial mucoid cells. The penis is wide at the base, the duct sinuous (as in Fig. 2 D). A flap of tissue projects from the side of the penis near the middle; internally this has no obvious differentiation from the rest of the penis wall, but the duct straightens after that point. The penis tip is filament-like. The distal portion of the penis at rest is stored in a penial pouch. Female system typical for Parvanachis, with a gonopericardial duct, and no bursa copulatrix. Gland mass stains red overall, with a purple staining band through the middle closer to the posterior end. The gland lumen is pouched. Anterior end of gland mass with a short (almost nonexistent) flat-walled vestibule. Gonopericardial duct coiled, with ciliated cuboidal epithelium near gland mass. Closer to the pericardium duct becomes thicker walled and slightly folded. Reproductive system in female sectioned contained no sperm, however pericardium was full of cellular material and secretions. Remarks. This species is very similar to both P. nigricans and to P. pygmaea and P. mullineri. Color and pattern can be similar in all four, but this species has the oblique shell color elements seen in P. nigricans. P. nigricans is larger and has coarser sculpture; it has denser axial ridges (16 to 19 on the penultimate whorl), better developed spiral grooves over most of the shell, and a more strongly developed subsutural groove, but weaker parietal denticles. Parvanachis pygmaea is also similar and roughly the same size, but with a more stereotypic color pattern lacking oblique elements, and having more axial ridges on the penultimate whorl (15 to 18 rather than the 14 to 16 in this species), and a better developed subsutural spiral groove. P. pygmaea, P mullineri and P. adamsi tend to have lighter shells with dark markings, while P. nigricans tends to have a colored shell with pale markings. This species also has the long filament tip on the penis tip seen in P. nigricans, but lacking in P. pygmaea.Published as part of Maintenon, Marta J., 2014, Taxonomic revision of the species of Parvanachis Radwin, 1968 (Gastropoda: Columbellidae) from the Gulf of Panama, pp. 201-225 in Zootaxa 3753 (3) on pages 218-220, DOI: 10.11646/zootaxa.3753.3.1, http://zenodo.org/record/25331

Parvanachis minibrunnea Maintenon, 2014, n. sp.

Parvanachis minibrunnea n. sp. (Figures 1 F, 1 G, 5 C, 5 F) Types. Holotype, Fig. 1 F, (USNM 1231307), Isla Venado, Panama, 30 Jan. 2006, 3.23 mm long. 32 paratypes, USNM 1231308–1231313, see Table 1. Etymology. Named for being very small and brown. Taxonomic history. This species has been identified in collections as Parvanachis albonodosa (Carpenter, 1857). Carpenter originally described Anachis albonodosa as greenish white, marked with spots and zigzag lines of brown. The species is based on two specimens (syntypes) in the Reigen collection from Mazatlan, where they were found on sessile bivalves: an adult or subadult in the British Museum, and a broken juvenile at the US National Museum (USNM 716233, Fig. 1 E). Both are greenish white with brown lines and lack spiral grooves. Tryon (1883) discusses these briefly. Brann (1966) has a drawing of the British Museum syntype showing the shell pattern (pl. 56, fig. 654). Keen (1958, 1968) noted the shell characteristics, and stated that A. albonodosa has never been figured or recognized since its original description. She figures the British Museum syntype (1968, 1971; Keen’s two photos (1971) are of the same specimen). In 1971 however, (perhaps after discussions with Radwin, who she credits for help with the columbellid section of the book), Keen amends her original description of A. albonodosa to describe the shell as light brown with a greenish tint, with white tips to the ribs, which brings the description more in line with P. minibrunnea. She kept her original illustration of the British Museum syntype, which is indeed greenish white with dark markings rather than brown with white markings; but detail in the small greyscale photo is difficult to see. Closer examination of the types of Anachis albonodosa Carpenter indicates that Keen was initially correct. Anachis albonodosa is a very different species, and no other name has been found for the small brown shells with a band of white spots. The two syntypes of A. albonodosa bear no resemblance to any species I have seen from the eastern Pacific. They are however very similar to species of Zafra from the Indo-west Pacific, none of which have otherwise been reported to occur in the Eastern Pacific. A. albonodosa appears to be an adventitious species of Zafra. In light of this, I herein provide a new name for the previously misidentified small brown Panamic species. Diagnosis. Smallest of the Panamic Parvanachis. Shell narrow, red-brown with a narrow, spiral white band slightly below the suture. Animal cream colored with white specks. Material. Over 40 specimens were collected under near shore rocks in muddy sand at Isla Venado, Bique and Chumical. Four were sectioned and four dissected. Shell (Figs. 1 F, G): Smallest species of Parvanachis; shells of 12 individuals measured 3.05 to 4.05 mm long (avg. 3.30 mm), 1.35 to 1.55 mm wide (avg. 1.43 mm). Adults with 3.25 to 4 teleoconch whorls (avg. 3.58). Protoconch smooth, cream colored at the apex but darker brown toward the teleoconch, 3.25 to 4 whorls (avg. 3.6, n = 10). Axial sculpture dominant overall, with spiral grooves visible between rounded axial ridges. First teleoconch whorl with similarly strong axial and spiral sculpture so is effectively cancellate. Weak subsutural groove cuts across the axial ridges. Shell red-brown, with a narrow, spiral white band just below the subsutural groove. First teleoconch whorl with three darker spiral bands. Aperture edge thickened, with three vague pale bands on the outer edge, and a few denticles internally. Shallow posterior sinus present in the aperture edge. Parietal callus with axial ridge and sometimes weak denticles, anterior edge of the callus detached. Aperture shows color and markings seen on outside of shell. Body coloration: Body cream colored, with white speckles. Operculum (Fig. 5 C): Operculum oval with a terminal nucleus, pale yellow overall. Muscle scar bilobed, keel weak. Radula (Fig. 5 F): Lateral teeth of four adult (two male, two female) specimens dissected 18–20 µm long by 7.5 µm wide, with rectangular center plates. Adult female dissected had a radular ribbon with at least 125 tooth rows. Lateral teeth delicate, with three pointed secondary cusps, the bottom one pointed and curved down, and separated from the distal two by a wide gap. Reproductive anatomy: Female reproductive anatomy similar to that of other Parvanachis species, with a gonopericardial duct but no bursa copulatrix. Gland mass comprises a single unit that stains red anteriorly, pale purple posteriorly. Capsule gland lumen flat but thick, ciliated. Gonopericardial duct coiled, relatively short and of small diameter close to gland mass, widens closer to pericardium, no obvious glandular portion to the duct. Pericardium in the specimens examined full of sperm, reddish droplets and other pale staining material, wall thick. Anterior end of the gland mass and vestibule with an egg capsule in the vestibule in both females sectioned so details of the form and epithelium could not be ascertained; vestibule short and not muscular. Male reproductive system similar to that of P. dichroma, having a long spermiduct loop adjacent to the proboscis and a simple penis with a short filament tip (as in Fig. 2 B). Coiled seminal vesicle with primarily dark purple staining sperm, no secondary vesicle. Part of the body wall spermiduct adjacent to the pallial duct very thick, coiled, muscular. Spermiduct loop in body cavity with a tall, pale purple staining mucoidal epithelium and lots of mucoid secretions in the duct. Penial spermiduct straight, wide, nonsecretory, no obvious subepithelial secretory tissues. Penis in sectioned specimens curled back in the mantle cavity, without a pouch. The following four species (P. pygmaea, P. mullineri, P. adamsi n. sp. and P. nigricans n. comb.) are discussed together because they are difficult to tell apart, and all have a unique penis morphology not present in the previous species. They are also generally larger. These four species may comprise a monophyletic group, but not all regional species of Parvanachis have been investigated, so I am reluctant to suggest a new genus level group until a more global and more inclusive analysis can be undertaken.Published as part of Maintenon, Marta J., 2014, Taxonomic revision of the species of Parvanachis Radwin, 1968 (Gastropoda: Columbellidae) from the Gulf of Panama, pp. 201-225 in Zootaxa 3753 (3) on pages 212-213, DOI: 10.11646/zootaxa.3753.3.1, http://zenodo.org/record/25331

Lettres et entretiens : extraits

Università degli Studi di Triest

Taxonomic revision of the species of Parvanachis Radwin, 1968 (Gastropoda: Columbellidae) from the Gulf of Panama

Maintenon, Marta J. (2014): Taxonomic revision of the species of Parvanachis Radwin, 1968 (Gastropoda: Columbellidae) from the Gulf of Panama. Zootaxa 3753 (3): 201-225, DOI: 10.11646/zootaxa.3753.3.