Search

Yamazaki, Midori, Yamazaki, Kazunori, Rundell, J., Ueshima, Rei (2015): Systematic review of diplommatinid land snails (Caenogastropoda, Diplommatinidae) endemic to the Palau Islands. (3) Description of eight new species and two new subspecies of Hungerfordia. Zootaxa 4057 (4): 511-538, DOI: http://dx.doi.org/10.11646/zootaxa.4057.4.3

Hungerfordia nodulosa M. Yamazaki and Ueshima sp. nov. (Figs. 8, 12K 1���2) Materials examined. Holotype. Southern coast of western part of Bkulschuul island (Loc. Uo 8 - 2), Ulong islands, Palau. Collected by R. U. and M. Y. on Feb. 25, 2011. UMUTZ-MG-B 1149 T. Paratypes. Ulong: Loc. Uo 7 -1, 6 specimens, 27 /I/ 2010, UMUTZ-MG-B0934; Loc. Uo 8 -2, 11 specimens, 25 /II/ 2011, UMUTZ-MG-B 1149; Loc. Uo 8 -4, 22 specimens, 27 /II/ 2011, UMUTZ-MG-B 1226. Diagnosis. Shell sinistral, large for Hungerfordia species, conical. Shell color usually yellow on upper whorls, whitish on the last whorl, with two brown bands behind the aperture: upper band extended below the suture; lower band winding around the columella. Axial ribs developed on teleoconch; ribs on upper whorls (higher than the last whorls) strong, very widely spaced, weakly folded and shortly protruded at the peripheries of the whorls, elevated by axial protrusions of shell wall (along the ribs); ribs above the aperture closely spaced, low; ribs on the last 3 / 4 whorls strong, widely spaced, folded, partially and strongly elevated by axial protrusions of shell wall only at the peripheries of the whorls, with nodular swellings. Interspace between the widely spaced ribs with some closely spaced growth wrinkles below the suture. Aperture rather large; columellar axis within the aperture slightly oblique. Columellar tooth present, moderately developed. Peristome double. Outer peristome strongly developed, widely expanded beyond the inner peristome. Inner peristome squarish circular. Thickened lip developed behind the aperture, moderately wide; dorsal margin distinct, abruptly expanded from the preceding whorl as the outer peristome. FIGURE 8. Hungerfordia nodulosa sp. nov. (A 1���10) A topotypic paratype, Loc. Uo 8 - 2, B 1149 a: (A 10) sculpture on the penultimate whorl; (B 1���6) holotype, B 1149 T; (C) brownish color variant, a paratype from Loc. Uo 8 - 4, B 1226; (D 1���2) outer surface, and (D 3���4) inner surface of operculum, a topotypic paratype, B 1149. Scale bars, 1mm. All specimens, UMUTZ-MG. Description. Shell (Figs. 8 A���C, 12 K 1���2). Shell sinistral, thick, large for Hungerfordia species, conical, last whorl widest or last 2 whorls almost equally widest when excluding the thickened lip and peristomes; sides almost straight. Shell color usually yellow (occasionally reddish brown) on most whorls (higher than the last whorl), whitish on the last whorls, with two vague brown bands behind the aperture: upper band developed below the suture behind the outer peristome, becoming broader toward the aperture; lower band winding around the columella on the last 5 / 8 whorls, longer than the upper band. Whorls 5���6, slightly convex, but seemingly angulate at the peripheries of widely spaced ribs due to axial protrusions of the shell wall along the obliqued ribs, broadest (in longitudinal width) on the penultimate or the last whorl in dorsal view; last 3 / 4 whorls with characteristic nodular swellings along the ribs; last 1 / 8 whorls (including the thickened lip) expanded upward, ascending onto the penultimate whorl in left lateral view; last 1 / 4 whorls behind the outer peristome somewhat dorso-ventrally depressed at the base. Suture impressed; suture of a part of the last whorl occasionaly running slightly inside the angulate base (=basal keel) of the penultimate whorl. Constriction located above the parietal/collumellar junction of the aperture, internally without palatal plica except a low and axial crest caused by the constriction, associated with abrupt changes of the ribbing pattern. Tuba 7 / 8 whorls. Protoconch 3 / 4 ��� 1 + 1 / 4 whorls, almost smooth, very weakly and very finely punctated on the lower whorl. Apical septum absent. Axial ribs strongly developed on teleoconch, distinct throughout whorls, oblique to the coiling axis: ribs on first 1 / 3 ��� 1 / 2 whorls of teleoconch weak, closely spaced, very low, almost straight; ribs on the following 1 / 4 ��� 1 / 3 whorls of initial teleoconch gradually becoming stronger and more widely spaced; ribs on the subsequent whorls except the last whorl (ribs on the penultimate and the higher 2.5���3 whorls) strongly developed, very widely spaced (7���11 ribs in a whorl), elevated by axial protrusions of shell wall along the ribs, weakly folded and slightly protruded at the peripheries of the whorls, sharply edged, but never pointed at the shoulder, gradually becoming more closely spaced and less folded on the last 1 + 1 / 4 to last 1 whorls; ribs above the aperture (ribs on the last 1 + 1 / 8 to last 3 / 4 whorls around the constriction) closely spaced, low (but distinct), almost straight (not folded at the peripheries of the whorls); ribs on the last 3 / 4 whorls strongly developed, abruptly becoming very widely spaced again, folded and slightly protruded at the peripheries of the whorls, partially and characteristically elevated by axial protrusions of shell wall along the ribs, strongly elevated only along the central and folded part of the ribs (elevated only at the peripheries of the whorls), associated with nodular swellings, never elevated at the upper margin (between the suture and the periphery) and the lower (basal) margin (between the periphery and the columella). Interspace between the widely spaced ribs with interstitial growth wrinkles below and above the suture: upper growth wrinkles always developed below the suture on most whorls, closely spaced, moderately long, disappearing near the peripheries of the whorls; lower growth wrinkles occasionally present on the last 1 + 1 / 8 to last 1 + 1 / 4 ��� 2 whorls, weakly developed just above the suture of the next whorl, very short, much shorter than the upper wrinkles, closely spaced, occasionally and partially connected with the upper wrinkles by weak growth lines. Spiral sculpture present throughout teleoconch whorls, fine, prominent on the upper whorls, somewhat weakened on the last 3 / 4 whorls, sinuous in high magnification. Basal keel absent on the last whorl, but present on the penultimate and the higher few whorls, a trace of angulate base in younger stages, hardly visible in adult shell, but occasionally visible just above the suture of the last whorl in oblique umbilical view, narrow, associated with a very weak spiral depression. Axial protrusions of the shell wall (along the ribs) on the last 3 / 4 whorls characteristically developed, strongly developed only at the periphery of the whorls (developed only along the central and the folded parts of the ribs), never developed at the base and below the suture, with nodular swellings; abapertural side almost flat or faintly concave in oblique umbilical view, but seemingly convex in mid-umbilical view due to the obliqued axial ribs; adapertural side prominently convex, almost equally wide as the abapertural side in umbilical view. Base smooth except around the columella, convex on the last 1 to last 2 / 3 whorls, somewhat dorso-ventrally depressed on the last 1 / 4 whorls behind the outer peristome, with a weak circum-columellar fold; circum-columellar fold very weakly develped on the last 3 / 4 whorls around the columella, with fine and weak growth lines on its surface, expanded beyond the inner peristome at the columellar side in front view. Umbilicus closed in adult. Aperture rather large, not protruded, slightly tilted downward against the coiling axis in left lateral view; columellar axis within aperture rather oblique (tilted outward) against the coiling axis in front view. Columellar tooth present, moderately developed, emerged at the aperture, but never extended onto the inner peristome, deeply extended inward about 3 / 4 whorls. Peristome double. Outer pesitome strongly developed at the dorsal margin of the thickened lip, expanded beyond the inner peristome in front view; inner surface with fine and concentric growth lines; outer surface with fine spiral sculptures. Inner peristome squarish circular, moderately expanded, but less expanded at the upper palatal side, slightly expanded beyond the thickened lip at the basal side (in left lateral view), strongly protruded from the outer peristome; parietal margin located below the middle level of the upper body whorl. Thickened lip developed behind the aperture, moderately wide, with very fine growth lines on its outer surface, white, lacking of periostracum, almost cylidrically extended in umbilical view; dorsal margin distinct, abruptly expanded from the preceding whorl, with a developed outer peristome; upper most margin (= suture) ascending onto the penultimate whorl toward the aperture in left lateral view; base extended almost horizontally toward the aperture in left lateral view; palatal margin protruded beyond the penultimate whorl in apical view. Dimensions. Shell height 5.3���6.2 mm, shell diameter 3.2���3.7mm, suture width 2.5���2.7 mm, peristome height 1.9���2.4 mm. Operculum (Figs. 8 D 1���4). Operculum corneous, multispiral, circular, semi-transparent, amber-colored, thin, flat; outer surface smooth; inner surface with a very low, arcuate ridge near the columellar margin. Penis. Penis absent. Radula. Not examined. Distribution and ecology. Endemic to Palau: Ulong islands. The species inhabits limestone rocks. Remarks. H. nodulosa is similar to H. ringens (Crosse, 1866), but differs in the following ways: the axial ribs on the last 3 / 4 whorls are widely spaced, prominently folded and protruded at the peripheries of the whorls, and with characteristic nodular swellings; lateral fold, dorsal fold, and dorsal groove never developed on last 3 / 4 whorls; the axial ribs are also widely spaced at the dorsal side of the penultimate whorl (Figs. 12 I, K). H. nodulosa is also similar to H. spinoscapula Yamazaki & Ueshima, 2015, which coexists with the species in sympatry. H. nodulosa, however, differs from H. spinoscapula by the axial ribs being never sharply pointed at the shoulder, the strongly developed outer peristome, the absence of lateral fold and dorsal fold on the last 3 / 4 whorls, and the larger and broader shell in the sympatric populations (Figs. 12 J, K). Although most specimens of H. nodulosa are readily distinguished from H. robiginosa Yamazaki & Ueshima, 2015 by the yellow shell color, the brownish color variant of the species is very similar to H. robiginosa (Figs. 12 K 2, L). Brownish variant of H. nodulosa, however, differs from H. robiginosa by the following features: the axial ribs on the last 3 / 4 whorls are distinctly elevated at the peripheries of the whorls, associated with characteristic nodular swellings; the axial ribs on the penultimate and the higher 2 whorls are more widely spaced; the outer peristome is widely expanded beyond the inner peristome. Etymology. The specific name, derived from Latin nodulosus, refers to the characteristic nodular swellings on the last 3 / 4 whorls., Published as part of Yamazaki, Midori, Yamazaki, Kazunori, Rundell, J. & Ueshima, Rei, 2015, Systematic review of diplommatinid land snails (Caenogastropoda, Diplommatinidae) endemic to the Palau Islands. (3) Description of eight new species and two new subspecies of Hungerfordia, pp. 511-538 in Zootaxa 4057 (4) on pages 527-530, DOI: 10.11646/zootaxa.4057.4.3, http://zenodo.org/record/241686, {"references":["Crosse, H. (1866) Note sur les Mollusques opercules terrestres des iles Pelew ou Palaos. Journal de Conchyliologie, 14, 346 - 350.","Yamazaki, M., Yamazaki, K. & Ueshima, R. (2015) Systematic review of diplommatinid land snails (Caenogastropoda, Diplommatinidae) endemic to the Palau Islands. (2) Taxonomic revision of Hungerfordia species with low axial ribs. Zootaxa, 3976 (1), 1 - 89."]}

Hungerfordia fragilipennis M. Yamazaki and Ueshima sp. nov. (Figs. 5, 12R 1���2) Materials examined. Holotype. Eastern slope in front of Ngkur (Loc. Gk 6 - 2), northern Ngeruktabel island, Palau. Collected by R. U. on Oct. 22, 2007. UMUTZ-MG-B0737T. Paratypes. Ngeruktabel: Loc. Gk 6 -2, 56 specimens, 22 /X/ 2007, UMUTZ-MG-B0737; Loc. Gk 7 -6, 11 specimens, 30 /I/ 2010, UMUTZ-MG-B1004, 1012; Loc. Gk 7 -7, 2 specimens, 30 /I/ 2010, UMUTZ-MG-B 1024; Loc. Gk 7 -8, 8 specimens, 30 /I/ 2010, UMUTZ-MG-B 1034. Diagnosis. Shell sinistral, very large for Hungerfordia species, subconical with almost cylindrical last 2 whorls. Shell color transparent, upper whorls occasionally yellow-colored. Constriction located above the aperture, associated with disappearance of the wing-like ribs. Apical septum present. Apical whorls frequently decollated in adult. Axial ribs on most whorls strongly protruded as wings, very widely spaced; wing-like projections of the ribs high, very wide, arising at the suture, extended below to the suture of the next whorl, thin, very fragile, frequently broken or lost in adult; ribs above the aperture abruptly becoming closely spaced and very low, without any winglike projections. Interspace between the wing-like ribs almost smooth, polished, with fine and short growth wrinkles below the suture on the last 1.5-3 / 4 whorls. Spiral sculptures absent on most whorls. Umbilicus closed or very narrowly open. Aperture hardly or slightly tilted upward against the coiling axis. Columellar tooth very weak. Peristome double. Outer peristome strongly developed, widely expanded beyond the inner peristome in front view. Inner peristome squarish circular, widely expanded. Interspace between the inner and outer peristomes wide. Description. Shell (Figs. 5 A���E, 12 R 1���2). Shell sinistral, very large for Hungerfordia species, rather thin for winged Hungerfordia species, subconical with almost cylindrical last 2 whorls, penultimate whorl widest (last 2 whorls almost equally widest, but penultimate whorl slightly wider than the last whorl) when removing the protruded ribs and peristomes. Apical whorls (above the apical septum) frequently decollated in adult. Shell color transparent, upper whorls occasionally yellow-colored with the color becoming deeper towards the apical septum, apical whorls above the septum always uncolored. Whorls 6���7, convex on apical whorls, less convex on the last 2��� 3 whorls; last 1 / 16 ��� 1 / 8 whorls (including the interspace between the outer and inner peristomes) expanded upward and ascending onto the penultimate whorl in left lateral view. Suture impressed. Constriction located above the parietal/columellar junction of the aperture, associated with abrupt changes of the ribbing pattern, internally without palatal plica except a low axial-crest caused by the constriction. Tuba 7 / 8 whorls. Protoconch 1 + 1 / 4 whorls, almost smooth, very weakly and very finely punctated on the lower whorl. Apical septum present, constructed inside the apical whorls, constructed between a half whorl before and a half whorl after the protoconch/ teleoconch boundary. Axial ribs developed on the teleoconch, highly developed as wings on most whorls: ribs undeveloped on the first 1 / 16 whorls of the teleoconch; ribs on the following 1 / 4 whorls of initial teleoconch weak, rather moderately spaced, very low, without wing-like projections; ribs on the subsequent whorls except above the aperture (ribs on most whorls except above the aperture) strongly developed, widely and highly protruded as ���wings���, very widely spaced (9���12 ribs in a whorl), not synchronized with those on the previous whorls, with simply curved or very weakly angulate base-lines, weakly elevated by axial protrusions of shell wall along the ribs; wing-like projections of the ribs lamellar, high, thin, very fragile, frequently broken or lost in adult, very wide, arising at the suture, extended below to the suture of the next whorl, weakly curved or very faintly folded at the peripheries of the whorls, very weakly pointed with a rounded edge, protruded slightly upward on most whorls, becoming protruded rather downward on the last 3 / 4 whorls; ribs above the aperture (ribs on the last 1 + 1 / 8 ��� 1 whorls to the last 3 / 4 whorls around the constriction) abruptly becoming very closely spaced and very low, without any wing-like projection, weak, fine. Interspace between the wing-like ribs almost smooth, glossy, without any axial sculpture except some faint growth lines on most whorls, with fine and short growth wrinkles below the suture on the last 1.5���3 / 4 whorls (except above the aperture). Spiral sculptures almost undeveloped on the teleoconch, very faintly and very finely developed only on the initial 1 / 2 ��� 1 whorls of teleoconch, absent on the subsequent whorl (most teleoconch whorls). Base convex, without basal keel. Axial protrusions of shell wall (along the ribs) hardly developed on the last 3 / 4 whorls, very weak in umbilical view (visible after removal of the winged ribs). Umbilicus very narrowly opened or closed in adult, though hidden by the winged ribs in fresh specimens. Aperture hardly or slightly tilted upward against the coiling axis (in left lateral view), weakly protruded from the previous whorl; columellar axis within aperture almost vertical. Columellar tooth very weakly developed, very low, penetrated inward about 3 / 4 whorls. Peristome double, expanded. Outer peristome strongly developed, widely expanded beyond the inner peristome in front view, occasionally with a rounded baso-columellar edge, reflexed backward, abruptly expanded from the preceding whorl; inner surface smooth, polished, with fine concentric growth lines. Inner peristome squarish circular, widely expanded in all around (particularly at the parietal side), widely separated and protruded from the outer peristome, somewhat thickened and weakly reflexed; parietal margin located below or near the middle level of the upper body whorl. Interspace between the inner and outer peristomes rather wide, with very fine growth wrinkles or riblets on the outer surface; uppermost margin (= suture) ascending onto the penultimate whorl toward the aperture in left lateral view; base also ascending toward the aperture in left lateral view. Dimensions. Shell height 6.0 to> 6.6 mm (the largest specimen has a partially decollated apex), diameter 3.7��� 4.3 mm, suture width 2.5���2.7 mm, peristome height 2.0��� 2.5 mm. Operculum (Figs. 5 F 1���4). Operculum corneous, multispiral, circular, transparent, amber colored, thin, flat or slightly concave; outer surface smooth; inner surface with a low, weak, arcuate ridge near the columellar margin. Penis. Penis absent. Radula. Not examined. Distribution and ecology. Endemic to Palau: Eastern coast of northern part of Ngeruktabel island. The species frequently inhabits vertical walls of large limestone rocks, occasionally on limestone rubble. Remarks. H. fragilipennis is most similar to H. papilio stenoptera Yamazaki & Ueshima, 2013 (Figs. 12 R, S), but differs in the following points: the wing-like ribs are shorter, more fragile (frequently lost), and never synchronized with those on the previous whorls even on the apical whorls; the apical septum is present; the apical whorls above the septum is frequently decollated; the shell is larger, broader, and thinner; the umbilicus is distinctly narrower. H. fragilipennis is unique among winged Hungerfordia species in having rather thin shell, which is responsible for the very fragile wing-like ribs and the frequently decollated apical whorls. Etymology. The specific name, derived from Latin fragilis and pennis, refers to the very fragile wing-like ribs. FIGURE 5. Hungerfordia fragilipennis sp. nov. (A 1���6) A topotypic paratype, Loc. Gk 6 - 2, B0737a; (B) apical whorls of a decollated specimen, showing the apical septum, a topotypic paratype, B0737b; (C 1���6) holotype, note that the wing-like projections of the ribs are completely lost and that the apical whorls are decollated, B0737T; (D) yellowish variant, a topotypic paratype, B0737c; (E 1���4) a paratype from Loc. Gk 7 - 6, a specimen with well preserved wing-like ribs and a complete apex, B 1012; (F 1���2) outer surface, and (F 3���4) inner surface of operculum, a topotypic paratype, B0737. Scale bars, 1mm (A, C���E), 0.5 mm (B, F). All specimens, UMUTZ-MG., Published as part of Yamazaki, Midori, Yamazaki, Kazunori, Rundell, J. & Ueshima, Rei, 2015, Systematic review of diplommatinid land snails (Caenogastropoda, Diplommatinidae) endemic to the Palau Islands. (3) Description of eight new species and two new subspecies of Hungerfordia, pp. 511-538 in Zootaxa 4057 (4) on pages 520-522, DOI: 10.11646/zootaxa.4057.4.3, http://zenodo.org/record/241686, {"references":["Yamazaki, K., Yamazaki, M. & Ueshima, R. (2013) Systematic review of diplommatinid land snails (Caenogastropoda, Diplommatinidae) endemic to the Palau Islands. (1) Generic classification and revision of Hungerfordia species with highly developed axial ribs. Zootaxa, 3743 (1), 1 - 71. http: // dx. doi. org / 10.11646 / zootaxa. 3743.1.1"]}

Hungerfordia brachyptera M. Yamazaki and Ueshima sp. nov. (Figs. 6, 12Q) Materials examined. Holotype. 600m North of Metkerel a Chudel (Loc. Mc 6 - 3), northeastern part of Mecherchar island, Palau. Collected by R. U. on Oct. 25, 2007. UMUTZ-MG-B0842T. Paratypes. Mecherchar: same data as holotype, 2 specimens, UMUTZ-MG-B0842. Diagnosis. Shell sinistral, slender conical, large to very large for Hungerfordia species. Shell color white or transparent, upper whorls yellow colored. Apical septum present. Axial ribs on most whorls, strongly developed, highly protruded as wings, very widely spaced, elevated by axial protrusions of shell wall; wing-like projections of the ribs wide, rather short for winged Hungerfordia species, hardly or very weakly folded at the peripheries of whorls; ribs above the aperture closely spaced, very low, without wing-like projections; ribs on the last 3 / 4 whorls very widely spaced (with the interspace much wider than those on the upper whorls), with strong wing-like projections, very strongly elevated by axial protrusions of shell wall along the ribs. Interspace between the winged ribs with fine growth wrinkles below the suture. Spiral sculpture present throughout teleoconch whorls. Basal keel developed on the penultimate and the higher 1���2 whorls, a trace of angulate base in younger stage, obviously visible just above the suture of the last whorl, but disappearing on the last whorl itself. Aperture slightly protruded, weakly tilted downward in left lateral view. Columellar tooth very weak. Peristome double. Outer peristome strongly developed, widely expanded beyond the inner peristome, with a rounded baso-columellar edge. Inner peristome moderately expanded; parietal margin located below the middle level of the upper body whorl. Interspace between the outer and inner peristomes wide, rather smooth on the outer surface. Description. Shell (Figs. 6 A���D, 12 Q). Shell sinistral, large to very large for Hungerfordia species, slender conical, last whorl widest when removing the protruded ribs and peristomes. Shell color white or transparent, upper whorls occasionally yellow with the color becoming deeper towards the apical septum, apical whorls above the apical septum always uncolored. Whorls 7���8, convex; last 1 / 8 whorls (including interspace between the outer and inner peristomes) expanded upward, ascending onto the penultimate whorl in left lateral view. Suture impressed; suture on the last 1���3 / 4 to last 1 / 3 whorls deeply impressed, running slightly inside the basal keel of the penultimate whorl. Constriction located above the parietal/columellar junction of the aperture, associated with abrupt changes of the ribbing pattern, internally without palatal plica except a low axial-crest caused by the constriction. Tuba 7 / 8 whorls. Protoconch 1 + 1 / 3 whorls, almost smooth, lower whorl faintly and very finely punctated. Apical septum present, constructed inside the apical whorls, constructed at 0.5���1 + 1 / 3 whorls after the protoconch/teleoconch boundary. Axial ribs developed on the teleoconch, oblique to the coiling axis, strongly protruded as wings on most whorls; ribs on the first 1 / 4 ��� 1 / 3 whorls of the initial teleoconch closely spaced, low, almost straight (not folded at the peripheries of the whorls); ribs on the subsequent whorls higher than the last whorl (ribs on most whorls) strongly protruded as ���wings���, very widely spaced (8���10 ribs in a whorl), not synchronized with those on the previous whorls, hardly or weakly folded at the peripheries of the whorls, strongly elevated by axial protrusions of shell wall along the ribs; wing-like projections of the ribs strongly developed, but rather short for winged Hungerfordia species, rather wide, obtusely pointed at the shoulder, weakly curved upward, arising at the suture on upper whorls, becoming to be arising at some distance below the suture on the lower whorls, protruded rather upward, lower margin extended below to the suture of the next whorl, with fine spiral sculptures on the outer surface; ribs above the aperture (ribs on the last 1 to last 3 / 4 whorls around the constriction) abruptly changed, closely spaced, very low, without any wing-like projections, not folded; ribs on the last 3 / 4 whorls strongly developed as ���wings��� again, very widely spaced, with the interval much wider than those on the upper whorls (2 ribs in a quarter whorl), very highly elevated by axial protrusions of shell wall along the ribs (particularly at the first winged rib which reappears after the constriction, and the second rib located at right side of the aperture); wing-like projection of the ribs strongly developed and shaped as those on the upper whorls, protruded laterally or rather downward, arising at or near the suture. Interspace between the wing-like ribs with fine growth wrinkles below the suture, which are occasionally extended below to the suture of the next whorl. Spiral sculptures present throughout teleoconch whorls, fine, rather prominent, extended onto outer surface of the winglike ribs and outer peristome, sinuous in high magnification. Basal keel developed on the penultimate and the higher 1���2 whorls, a trace of angulate base in younger stages, recognizable just above the suture of the last 2���3 whorls in adult shell, most obvious just above the suture of the last whorl in oblique umbilical view, but disappearing on the last whorl itself. Base not convex, rather rugged by the strong axial protrusions of shell wall FIGURE 6. Hungerfordia brachyptera sp. nov. (A 1���9, D) A topotypic paratype, Loc. Mc 6 - 3, B0842a: (A 8) sculptures on the penultimate whorl, left lateral view, showing that the basal keel of the penultimate whorl (indicated by arrowheads) is developed just above the suture of the last whorl: (A 9) oblique umbilical view, note that the suture of the last whorl is running slightly inside the basal keel of the penultimate whorl (indicated by arrowheads), note also that the axial protrusions of shell wall are strongly developed along the ribs on the last 3 / 4 whorls; (B 1���5) holotype, B0842T; (C 1���2) a topotypic paratype, B0842b: (C 2) umbilical view, only the last whorl shown, note the strongly developed axial protrusions of shell wall along the ribs; (E 1���2) outer surface, and (E 3���4) inner surface of operculum, topotypic paratype, B0842a. Arrowheads indicate the basal keel of the penultimate whorl. Scale bars, 1mm. All specimens, UMUTZ-MG. along the ribs. Umbilicus closed in adult. Axial protrusions of shell wall (along the wing-like ribs) on the last 3 / 4 whorls very strongly developed, in particular, highly developed at a strong rib just after the constriction and the next rib protruded toward the right lateral side of the aperture; abapertural side concave in umbilical view; adapertural side concave or only slightly convex in umbilical view, almost similarly wide as the abapertural side. Aperture weakly protruded (at the parietal side), tilted downward against the coiling axis in left lateral view; columellar axis within aperture slightly oblique to the coiling axis. Columellar tooth very weak, very low, recessed deeply inside the aperture. Peristome double. Outer peristome strongly developed, very widely expanded beyond the inner peristome in front view, abruptly reduced at the lower columellar side to form a rounded baso-columellar edge, with fine concentric growth lines on the inner surface, with fine spiral sculptures on the outer surface. Inner peristome squarish circular, rather widely expanded in all around, weakly protruded from the previous whorl at the parietal side, strongly protruded from the outer peristome, parietal margin located below the middle level of the upper body whorl. Interspace between the inner and outer peristomes wide, widest at the upper palatal side and narrowed toward the base in left lateral view, almost smooth and with fine growth lines on the outer surface; uppermost margin (= suture) ascending onto the penultimate whorl toward the aperture in left lateral view. Dimensions. Shell height 5.9���6.3 mm, diameter 3.3���3.6 mm, suture width 2.2���2.3 mm, peristome height 1.8 ��� 2.0 mm. Operculum (Figs. 6 E 1���4). Operculum corneous, multispiral, circular, thin, transparent, amber colored, slightly concave; outer surface smooth; inner surface with a low, thin, weak, arcuate ridge near the columellar margin. Penis. Penis absent. Radula. Not examined. Distribution and ecology. Endemic to Palau: Mecherchar island. The species inhabits vertical walls of large limestone rocks, and limestone rubble. Remarks. H. brachyptera is distinguished from other winged Hungerfordia species in the following points: the characteristic basal keel is developed on the penultimate whorls and the higher 1���2 whorls (most prominent just below the suture of the last whorl); the wing-like ribs are shorter, less folded; the ribs on the last 3 / 4 whorls are very widely spaced, very strongly elevated by axial protrusions of shell wall along the ribs; the ribs above the aperture are closely spaced, very low, and without any wing-like projections; the inner peristome is not so widely expanded; apical septum is present. Etymology. The specific name, derived from Latinized Greek brachys and pterus, refers to the wing-like ribs that are rather short for winged Hungerfordia species., Published as part of Yamazaki, Midori, Yamazaki, Kazunori, Rundell, J. & Ueshima, Rei, 2015, Systematic review of diplommatinid land snails (Caenogastropoda, Diplommatinidae) endemic to the Palau Islands. (3) Description of eight new species and two new subspecies of Hungerfordia, pp. 511-538 in Zootaxa 4057 (4) on pages 523-525, DOI: 10.11646/zootaxa.4057.4.3, http://zenodo.org/record/241686

Hungerfordia spiroperculata M. Yamazaki and Ueshima sp. nov. (Figs. 3, 4, 12 M) Materials examined. Holotype. West of Bkulsechuul island, Ulong islands (Loc. Uo 7 - 2), Palau. Collected by R. U. and M. Y. on Jan. 27, 2010. UMUTZ-MG-B0942T. Paratypes. Ulong: Loc. Uo 7 -2, 4 specimens, 27 /I/ 2010, UMUTZ-MG-B0942; ditto, 42 specimens, 25 /II/ 2011, UMUTZ-MG-B 1146; Loc. Uo 8 -3, 1 specimen, 25 /II/ 2011, UMUTZ-MG-B 1168; ditto, 16 specimens, 27 /II/ 2011, UMUTZ-MG-B 1223. Diagnosis. Shell sinistral, medium-sized for Hungerfordia species, slender fusiform, with spiny projections. Shell color yellowish brown. Constriction located above the aperture, followed by an abrupt whorl swelling (= ventral protrusion), which is sharply edged along a rib in umbilical view. Tuba 3 / 4 whorls, with coarse granular projections on the inner surface. Axial ribs very strongly developed, strongly folded and highly protruded as spines at the periphery of whorls, rather widely spaced, absent or greatly reduced above the aperture, but developed again on the last 3 / 4 whorls; spiny projection of the ribs very long, slender, almost tubular, with an opened and blunt terminus, white. Interspace between the spiny ribs smooth. Apical septum present. Spiral sculpture almost absent on teleoconch whorls, but faintly developed only on outer surface of the spiny projections. Aperture squarish circular, with a canal-like depression at the baso-columellar edge. Columellar tooth weakly developed, hidden by the columellar lip in front view. Peristome double. Outer peristome expanded, obtusely angulate at the palatal side. Inner peristome narrower than the outer peristome, with a prominent baso-palatal lip, with a short columellar lip just after the columellar tooth. Interspace between the inner and outer peristomes very narrow. Operculum distinctive, unusually shaped for Hungerfordia species; outer side elevated, with a strong spiral fringe; inner side with a strong arcuate ridge. Description. Shell (Figs. 3 A���E, 12 M). Shell sinistral, medium-sized for Hungerfordia species, slender fusiform, penultimate whorl widest when removing the protruded ribs and peristomes. Shell color yellowish brown, apical whorls above the apical septum always uncolored. Whorls 6���7.5, somewhat convex, obtusely angulate at the periphery of the penultimate whorl; last 1 / 8 whorls (including the outer and inner peristomes) weakly expanded both upward and downward, weakly ascending onto the penultimate whorl in left lateral view. Suture impressed. Constriction located above the parietal/columellar junction of the aperture, followed by an abrupt whorl swelling (= ventral protrusion), associated with abrupt disappearance or reduction of the spiny ribs, internally without palatal plica except for a low axial-crest caused by the constriction. Tuba 7 / 8 whorls, with coarse granular projections on the inner surface. Protoconch 1 ��� 1 + 1 / 4 whorls, almost smooth, very weakly and very finely punctated on lower whorl. Apical septum present, constructed inside at 1 + 1 / 8 ��� 1.5 whorls after the protoconch/ teleoconch boundary. Axial ribs developed on teleoconch, strongly protruded as tubular spines on most whorls; ribs on the first 1 / 8 whorls of teleoconch hardly developed; ribs on the following 0���1 / 4 whorls of the initial teleoconch low, moderately spaced, only slightly protruded, almost straight (not folded); ribs on the subsequent whorls except above the aperture (ribs on most whorls) strong, highly protruded as tubular spines, rather widely spaced (14���19 ribs in a whorl), not synchronized with those on previous whorls, strongly curled and folded at the peripheries of the whorls; upper margin of the ribs above the tubular spines (between the suture and spines) very low, never elevated nor protruded (attached to the basal wall), indistinct, hardly visible; lower margin below the spines (between the spines and the suture of the next whorl) also very low, very weakly elevated or faintly flared, but never protruded; spiny projections on the ribs almost tubular, with a blunt and opened terminus, with a narrow longitudinal slit at the adapertural side, very high (long), slender, almost straight, narrow, arising near the peripheries of the whorls (at some distance below the suture), disappearing just below the peripheries of the whorls, with the lower margin close to or slightly overlapped with the upper margin at the base, protruded rather upward on most whorls, becoming protruded downward on last 3 / 4 whorls, very weakly and very finely striated on the outer surface; ribs on last 1 to last 3 / 4 whorls above the aperture (around the constriction) abruptly weakened, completely absent or greatly reduced, variable in the degrees of the reduction, rarely with vestigial spiny projections; ribs on the last 3 / 4 whorls developed once more, strongly protruded as tubular spines as those on the upper whorls. Interspace between the spiny ribs almost smooth on upper whorls, with or without very weak and very fine growth lines below and above the suture in high magnification, occasionally with prominent and fine growth wrinkles below the suture on the last 3 / 4 whorls. Spiral sculpture hardly developed on the teleoconch, hardly visible on the whorls in low magnification, but occasionally present as very faint and very fine vestiges in high magnification, weakly but more prominently developed on the outer surface of the tubular spines, sinuous in FIGURE 3. Hungerfordia spiroperculata sp. nov. (A 1���6) A topotypic paratype, Loc. Uo 8 - 1, B 1146 a; (B 1���5) a topotypic paratype after removal of the spiny ribs, B 1146 b: (B 2) aperture, front view, showing the baso-palatal lip (blue arrowhead) and the columellar lip (yellow arrowhead), note that the columellar tooth (red arrowhead) is hidden by the columellar lip; (B 3) basal part of the columella after removal of the palatal wall of the aperture, left lateral view, note that the columellar lip (yellow arrowhead) occurs after the end of the columellar tooth (red arrowhead); (B 4) umbilical view, showing the strong ventral protrusion after the constriction, arrowheads are same as in Fig. B 2; (B 5) inner surface of the ventral protrusion on the last 3 / 4 whorls, showing the presence of granular projections inside the shell; (C) partially opened apical whorls, showing the presence of the apical septum, a topotypic paratype, B 1146 c; (D 1���5) holotype, B 1146 T; (E 1���2) a yellowish paratype with vestigial spiny ribs above the aperture, Loc. Uo 7 - 2, B0942; (F 1���2) outer surface, and (F 3���4) inner surface of operculum, a topotypic paratype, B 1146 d; (G 1���2) operculum, a topotypic paratype, B 1146 e. Scale bars, 1mm (A, B 1���4, C, D, E), 0.5 mm (B 5, F, G). All specimens, UMUTZ-MG. FIGURE 4. Hungerfordia spiroperculata sp. nov. (A 1) Radulae; (A 2) radula with a central tooth coming off from the basal membrane; (B) buccal mass, note the very long radular ribbon which forms a large coil within the radular sac; paratype, UMUTZ-MG-B0942. Scale bars, 50 ��m (A 1), 20 ��m (A 2), 500 ��m (B). high magnification. Base rather convex, with a prominent circum-columellar fold; circum-columellar fold developed around the columella on the last 5 / 8 whorls, becoming stronger and more prominently ridged toward the aperture in umbilical view. Ventral protrusion strongly developed, occurring at a short distance after the constriction, caused by an axial protrusion of shell wall along a spiny rib, which firstly reappears after the constriction, hidden by spiny projection of the rib in fresh specimens, sharply edged along the rib in umbilical view (visible after removal of the spiny projection of the rib), highly projected toward the ventral side. Axial protrusion of shell wall along the ribs on last 3 / 4 whorls very weakly developed after the ventral protrusion, hidden by the tubular spines (only visible after removal of the tubular spines); abapertural side very narrow; adapertural side convex, much wider than the abapertural one in umbilical view. Umbilicus closed in adult. Aperture hardly tilted against the coiling axis in left lateral view, not protruded, located at base of the previous whorl (hardly expanded onto the upper body whorl in front view), characteristically shaped with a canal-like depression at the basocolumellar edge, which is caused by development of a baso-palatal lip and a columellar lip; columellar axis within aperture almost vertical. Columellar tooth weakly developed, recessed inside the aperture, rather sharply edged just before the aperture, hidden by a columellar lip in front view, never extended onto the inner peristome, penetrated inward about 3 / 4 whorls. Peristome double. Outer peristome developed, shortly expanded beyond the inner peristome at the palatal side in front view, obtusely angulate at the palatal side, weakly protruded at the upper palatal side in left lateral view, rather gradually expanded from the preceding whorl. Inner peristome weakly expanded, squarish circular with rounded palatal and baso-columellar edges, characteristically shaped by a basopalatal lip and a columellar lip, leaving a canal-like depression at the baso-columellar edge between these lips, somewhat narrower than the outer peristome, hardly or slightly protruded from the outer peristome; parietal margin located at the base of the upper body whorl (located far below the middle level of the upper body whorl); basopalatal lip strongly developed inside the inner peristome, a narrow ridge extended along and inside the palatal and basal margin of the inner peristome, weakly but prominently protruded forward in umbilical view, becoming stronger toward the base, abruptly truncated at the base at a short distance before the columella; columellar lip weakly developed at the columellar side of the inner peristome (near the base), a weak and axial ridge developed near the base of the columella, short, low, seemingly a columellar tooth in front view, but a distinct structure developed after the end of the columellar tooth, shortly protruded forward in umbilical view. Interspace between the inner and outer peristome very narrow or almost without space at the palatal and basal sides; columellar side shortly erected, weakly grooved on its outer side. Dimensions. Shell height 4.5���5.1 mm, diameter 3.2���4.1 mm, suture width 1.8���2.1 mm, peristome height 1.3��� 1.5 mm. Operculum (Figs. 3 F 1���4, G 1���2). Operculum corneous, multispiral, circular, not flat, semi-transparent, amber colored, unusually shaped for Hungerfordia species; outer side smooth and flat only at the central part, elevated at the marginal part, with a wide and thin spiral fringe; inner side with a strong arcuate ridge neat the columellar margin; arcuate ridge highly protruded, thick, C-letter shaped. Penis. Penis absent. Radula (Figs. 4 A 1���2, B). Radula of specialized type of taenioglossate; ribbon very long, forming a large coil within the radular sac. Central tooth very large, with a very wide, large, and blunt central cusp, with 1���2 pair(s) of vestigial lateral cups, basal plate longitudinally elongated and strongly constricted at the base. Lateral teeth also large, with a large, very wide and blunt major cusp, with a vestigial inner cusp, without outer cusp, basal plate obliquely elongated. Inner marginal teeth shorter than the lateral teeth, elongated, with a large major cusp, with two small and slender inner cusps, with a small outer cup. Outer marginal teeth shorter than the inner marginal, with a large, wide and blunt major cusp, with two small and slender inner cusps, without outer cusp. Distribution and ecology. Endemic to Palau: Ulong islands. The species inhabits limestone rocks. Remarks. H. spiroperculata is readily distinguished from other spiny Hungerfordia species by the following features: the development of a strong baso-palatal lip inside the inner peristome; the presence of a short columellar lip just after the columellar tooth; the development of a strong ventral protrusion after the constriction; the presence of granular projections inside the tuba; the spiny projections of the ribs being tubular and slender; the slender shell with rather closely spaced spiny ribs. H. echinata tubulispina Yamazaki & Ueshima, 2013 is most similar to the species in having tubular projections of the ribs (Figs. 12 M, N), but is distinguishable by the features mentioned above. H. spiroperculata is also unique among the genus in having a peculiar operculum (Figs. 3 F 1���4, G 1���2): the operculum is not flat, with a strongly protruded spiral fringe on the outer surface, with a strongly developed arcuate ridge on the inner side. These unusual opercular features are rather similar to those of the other Palauan diplommatinid genus, Palaina (Eupalaina), but differs from the latter in that the operculum is not deeply concave nor thickened (Yamazaki et al., 2013). Despite such superficial similarities to Palaina in the operculum, the characteristic features of the shell, the radulae, and the absence of penis in the species consistently support the taxonomic placement in Hungerfordia. Occurrence of such an exceptional type of operculum within Hungerfordia shows that operculum morphology is not necessarily conserved within the genus. Etymology. The specific name, derived from the Latin spiro + operculatus, refers to the characteristic operculum with a strong spiral fringe., Published as part of Yamazaki, Midori, Yamazaki, Kazunori, Rundell, J. & Ueshima, Rei, 2015, Systematic review of diplommatinid land snails (Caenogastropoda, Diplommatinidae) endemic to the Palau Islands. (3) Description of eight new species and two new subspecies of Hungerfordia, pp. 511-538 in Zootaxa 4057 (4) on pages 517-520, DOI: 10.11646/zootaxa.4057.4.3, http://zenodo.org/record/241686, {"references":["Yamazaki, K., Yamazaki, M. & Ueshima, R. (2013) Systematic review of diplommatinid land snails (Caenogastropoda, Diplommatinidae) endemic to the Palau Islands. (1) Generic classification and revision of Hungerfordia species with highly developed axial ribs. Zootaxa, 3743 (1), 1 - 71. http: // dx. doi. org / 10.11646 / zootaxa. 3743.1.1"]}

Hungerfordia chilorhytis M. Yamazaki and Ueshima sp. nov. (Figs. 10, 12E) Material examined. Holotype . Belual a kelat, a northern islet of Ngemelis islands (Loc. Gm 7 - 1), Palau. Collected by R. U. and M. Y. on Jan. 28, 2010. UMUTZ-MG-B0944T. Paratypes. Ngemelis : same data as holotype, 28 specimens, UMUTZ-MG-B0944. Diagnosis. Shell sinistral, small for Hungerfordia species, slender conical or fusiform. Last 3 / 4 whorls characteristically sculptured by strong dorsal grooves. Shell color yellow. Axial ribs low, closely spaced on most whorls (higher than the last 3 / 4 whorls), irregularly spaced and partially indistinct on the last 3 / 4 whorls. Spiral sculpture present throughout teleoconch whorls. Dorsal grooves strongly developed on the last 3 / 4 whorls, axial (parallel to the axial ribs), 5���6, deeply impressed, widely spaced. Interspace between the axial grooves convex, somewhat protruded like ���axial folds���, with 2���5 of ribs on its surface. Base convex, with a prominent circumcolumellar fold; circum-columellar fold strongly developed, almost radially straight in umbilical view. Outer peristome absent. Inner peristome somewhat distorted, obtusely angulate at the periphery, squarish circular, with a characteristic baso-palatal lip. Baso-palatal lip developed along the palatal and basal margin of the inner peristome, a ridge-like thickening, abtuptly truncated at a short distance before the columella (at the base), leaving a canal-like depression at the baso-columellar edge of the inner peristome. Columellar tooth moderately developed. Thickened lip developed behind the aperture, wide; dorsal margin distinct; palatal side abruptly expanded from the preceding whorl, then gradually narrowed toward the aperture in umbilical view. Endemic to an islet of Ngemelis islands. Description. Shell (Figs. 10 A���C, 12 E). Shell sinistral, thick, small for Hungerfordia species, slender conical or fusiform, penultimate whorl widest when excluding the thickened lip; sides almost straight or slightly concave on the upper whorls, convex on the last 2 whorls. Shell color yellow. Whorls 6���6.5, convex, broadest (in longitudinal width) on the penultimate or the last whorl in dorsal view: last 3 / 4 whorls characteristically sculptured, with strong axial dorsal grooves; last 1 / 8 whorls (including the thickened lip) expanded upward, ascending onto the penultimate whorl in left lateral view. Suture impressed. Constriction located above the parietal/columellar junction of the aperture, internally without palatal plica except a low, axial crest caused by the constriction, without FIGURE 10. Hungerfordia chilorhytis sp. nov. (A 1���8) A topotypic paratype, Loc. Gm 7 - 1, B0944a: (A 7) aperture, showing the developed baso-palatal lip of the inner peristome; (B 1���6) holotype, B0944T; (C) a topotypic paratype, B0944b; (D 1���2) outer surface, and (D 3���4) inner surface of operculum, a topotypic paratype, B0944c. Scale bars, 1mm (A 1���6, A 8, B, C), 0.5 mm (A 7, D). All specimens, UMUTZ-MG. parietal tooth. Tuba 7 / 8 whorls. Protoconch 1 ��� 1 + 1 / 8 whorls, almost smooth, very weakly and finely punctated on the lower part of whorl. Apical septum absent. Axial ribs present on teleoconch, almost straight (never folded at the peripheries of the whorls), oblique to the coiling axis: ribs on most whorls (higher than the last 3 / 4 whorls) distinct, closely spaced (13���22 ribs in a quarter whorl), low; ribs on the last 3 / 4 whorl irregularly spaced, partially indistinct. Dorsal grooves strongly developed on the last 3 / 4 whorls, axial (parallel to the axial ribs), deeply impressed, 5���6, widely spaced, arising at short distance below the suture, becoming weaker toward the columella, internally protruded as ���axial folds��� within the aperture. Interspace between the dorsal grooves convex, somewhat protruded like ���axial dorsal folds���, wide, frequently edged along a rib in umbilical view, with 2���5 ribs on its surface, also with short and closely spaced growth lines or reduced ribs just below the suture. Spiral sculpture present on teleoconch, continued throughout whorls, but somewhat weakened on the last 3 / 4 whorls, fine, rather prominent, sinuous in high magnification. Base rather convex, with a prominent circum-columellar fold; circumcolumellar fold rather strongly developed on the last 3 / 4 whorls, almost radially straight in umbilical view, extended along the columellar side of the thickened lip, strongest and broadest just after the constriction, then gradually becoming weaker toward the columella. Umbilicus closed in adult. Aperture not protruded, slightly tilted downward against the coiling axis in left lateral view; columellar axis within aperture almost vertical. Columellar tooth moderately developed, emerged at the aperture, but never extended onto the inner peristome, deeply extended inward about 3 / 4 whorls. Peristome single. Outer peristome never developed at the dorsal margin of the thickened lip. Inner peristome characteristically shaped and sculptured, squarish circular, obtusely angulate at the periphery, moderately expanded at the parietal, columellar, and basal sides, depressed and less expanded at the upper palatal side (in front view), with a characteristic baso-palatal lip; parietal margin located below the middle level of the upper body whorl. Baso-palatal lip developed along the palatal and basal margin of the inner peristome, a ridgelike thickening which is continued from the angular edge to the base of the inner peristome, strongly developed at the base, but abtuptly truncated at a short distance before the columella (at the base), leaving a canal-like depression at the baso-columellar edge of the inner peristome (in umbilical view), with fine glanular sculptures on its surface. Thickened lip developed behind the aperture, rather wide in left lateral view, expanded beyond the inner peristome in front view, white, lacking of periostracum, with fine growth lines on its outer surface; palatal side slightly or hardly narrowed toward the aperture in umbilical view; dorsal margin distinct, abruptly expanded from the preceding whorl at the palatal side; uppermost margin (= suture) ascending onto the penultimate whorl toward the aperture in left lateral view; base extended almost horizontally or slightly ascending toward the aperture in left lateral view; palatal margin somewhat protruded beyond the penultimate whorl in apical view. Dimensions. Shell height 2.8���3.2 mm, shell diameter 1.7 ���2.0 mm, suture width 1.4���1.6 mm, peristome height 0.9���1.1 mm. Operculum (Figs. 10 D 1���4). Operculum corneous, multispiral, circular, semi-transparent, amber-colored, thin, flat; outer surface smooth; inner surface with a low, C-letter shaped ridge at the columellar side. Penis. Penis absent. Distribution and ecology. Endemic to Palau: Belual a kelat, a northern islet of Ngemelis islands. The species inhabits limestone rocks. Remarks. H. chilorhytis is similar to H. crenata Yamazaki & Ueshima, 2015 in the sculpture and the ribbing pattern, but differs in the distinctly larger shell size and the presence of a baso-palatal lip inside the inner peristome (Figs. 12 D, E). Eymology. The species name derived from Latinized Greek, cheilos + rhytis, refers to the inner peristome with a characteristic baso-palatal lip., Published as part of Yamazaki, Midori, Yamazaki, Kazunori, Rundell, J. & Ueshima, Rei, 2015, Systematic review of diplommatinid land snails (Caenogastropoda, Diplommatinidae) endemic to the Palau Islands. (3) Description of eight new species and two new subspecies of Hungerfordia, pp. 511-538 in Zootaxa 4057 (4) on pages 532-534, DOI: 10.11646/zootaxa.4057.4.3, http://zenodo.org/record/241686, {"references":["Yamazaki, M., Yamazaki, K. & Ueshima, R. (2015) Systematic review of diplommatinid land snails (Caenogastropoda, Diplommatinidae) endemic to the Palau Islands. (2) Taxonomic revision of Hungerfordia species with low axial ribs. Zootaxa, 3976 (1), 1 - 89."]}

Hungerfordia goniobasis exserta M. Yamazaki and Ueshima subsp. nov. (Figs. 1, 12G) Materials examined. Holotype. 0.1 km north of Ngeruleomel beach (Uo 8 - 5), Ulong island, Palau. Collected by R. U. and M. Y. on Feb. 27, 2011. UMUTZ-MG-B 1242 T. Paratypes. Ulong: same data as holotype, 15 specimens, UMUTZ-MG-B 1242. Diagnosis. Shell similar to other subspecies of H. goniobasis, but differs in the following features. Inner peristome strongly protruded from the outer peristome. Interspace between the inner and outer peristomes wide. Outer peristome widely expanded, distinctly expanded from the basal keel at the baso-columellar edge. Basal keel short, weaker, located close to the columella. Distribution endemic to Ulong islands. Description. Shell (Figs. 1 A���C, 12 G). Shell sinistral, small for Hungerfordia species, slender conical, penultimate whorl widest (slightly wider than the last whorl) when removing the protruded ribs. Shell color usually white, rarely reddish brown. Whorls 7���7.5, almost regularly coiled (increasing almost regularly in both width and height), rather convex; last 1 / 8 whorls never ascending onto the penultimate whorl in left lateral view. Suture impressed. Constriction located above the parietal/columellar junction of the aperture, associated with prominent changes of the ribbing density and height of the spiny ribs, internally without palatal plica except a low axial-crest caused by the constriction. Tuba 7 / 8 whorls. Apical septum absent. Protoconch 1 ��� 1 + 1 / 8 whorls, almost smooth, very weakly and very finely punctated in high magnification. Axial ribs developed on the teleoconch, oblique to the coiling axis, strongly and highly protruded as spines on most whorls; ribs undeveloped on the first 1 / 3 ��� 1 / 2 whorls of the teleoconch; ribs on the following 1 / 8 whorls of initial teleoconch rather low, moderately spaced, almost straight (not folded at the peripheries of the whorls); ribs on the subsequent whorls (except above the aperture) very strong, highly protruded as spines, very widely spaced (8���11 ribs in a whorl), not synchronized with those on the previous whorls, somewhat folded at the peripheries of the whorls (hardly folded on the upper whorls, but becoming strongly folded on the lower whorls), elevated by axial protrusions of shell wall along the ribs; upper margin of the ribs above the spiny projection (between the suture and the spines) low, slightly flared; spiny projections of the ribs highly protruded, weakly pointed, folded, trough-shaped, weakly curved upward, protruded slightly upward on the upper whorls, becoming gradually protruded laterally or rather downward on the last whorl, rather narrow, arising at some distance below the suture (near the peripheries of the whorls), upper margin folded over the lower margin, lower margin gradually reduced in height and extended below to the suture of the next whorl, with fine spiral sculptures on the outer surface; ribs above the aperture (ribs on the last 1 to last 3 / 4 whorls around the constriction) abruptly becoming moderately spaced (more closely spaced than those on the preceding and following whorls), with distinctly shorter spiny projections than those on the upper whorls, but still retaining developed spiny projections, less folded than those on the upper whorls. Interspace between the spiny ribs with characteristic growth wrinkles; growth wrinkles strongly developed below the suture on the last 3���4 whorls, coarse, shortly but distinctly protruded, somewhat lamellar, short, disappearing above the peripheries of the whorls. Spiral sculpture present on teleoconch, prominent throughout whorls, fine, extended onto the outer surface of the spiny ribs and the outer peristome, sinuous in high magnification. Base not convex, with a characteristic basal keel. Basal keel developed at the base of the last 5 / 8 whorls, rather sharply edged, leaving a shallow and narrow ���pseudo-umbilicus��� in front and oblique umbilical views, shorter and weaker than other subspecies of H. goniobasis, located very close to (developed just around) the columella in umbilical view, arising at a short distance after the constriction, connected to the outer peristome, but is not fused with the outer peristome like other subspecies of H. goniobasis, with a series of fine growth lines at the columellar side, extended beyond the inner peristome (at the columellar side) in front view. Umbilicus closed in adult. Axial protrusions of shell wall (along the ribs) strongly developed on the last 3 / 4 whorls. Aperture of usual size, tilted downward against the coiling axis, located below the base of the upper body whorl, slightly protruded from the previous whorl, strongly protruded from the outer peristome; columellar axis within aperture almost vertical. Columellar tooth weakly developed, low, located near the base of the columella, visible within the aperture, but never extended onto the inner peristome. Peristome double. Outer peristome strongly developed, widely expanded beyond the inner peristome in front view, independent of the basal keel (not fused with the basal keel as other subspecies of H. goniobasis), distinctly expanded from the basal keel at the baso-columellar edge in oblique umbilical view, abruptly expanded from the preceding whorl, with a series of fine growth lines on the inner surface, with fine spiral sculptures on the outer surface. Inner peristome squarish circular, strongly protruded from the outer peristome, weakly expanded at the basal and columellar sides, hardly expanded at the parietal and upper palatal sides; parietal margin almost straight, located below the base of the upper body whorl, slightly protruded downward from the previous whorl. Interspace between the outer and inner peristomes wide, with a series of fine riblets or growth lines. FIGURE 1. Hungerfordia goniobasis exserta subsp. nov. (A 1���9) A topotypic paratype, Loc. Uo 8 - 5, B 1242 a: (A 9) sculpture on the penultimate whorl; (B 1���6) holotype, B 1242 T; (C) reddish color variant, a topotypic paratype, B 1242 b; (D 1���2) outer surface, and (D 3���4) inner surface of operculum, a topotypic paratype, B 1242 c. Scale bars, 1mm (A 1���8, B, C), 0.5 mm (A 9, D). All specimens, UMUTZ-MG. Dimensions. Shell height 2.9���3.1 mm, diameter 1.9 ���2.0 mm, suture width 1.1 mm, peristome height 0.7���0.8 mm. Operculum (Figs. 1 D 1���4). Operculum corneous, multispiral, circular, transparent, amber colored, thin, rather flat; outer surface smooth; inner surface with a low, arcuate ridge near the columellar margin. Penis. Penis absent. Radula. Not examined. Distribution and ecology. Endemic to Palau: Ulong islands. The subspecies inhabits limestone rocks. Remarks. H. goniobasis exserta is easily distinguished from other subspecies of H. goniobasis in the following points: the inner peristome is strongly protruded from the outer peristome, never recessed inside the outer peristome; the interspace between the inner and outer peristomes is wide, with a series of fine growth lines or riblets on its surface; the basal keel is weaker, shorter, less sharply edged, and located close to the columella; the outer peristome is distinctly expanded beyond the basal keel at the baso-columellar edge, not completely merged with the basal keel like other subspecies; the distribution of this subspecies is endemic to Ulong islands. Etymology. The subspecific name, derived from Latin exsertus, refers to the aperture which is strongly protruded from the outer peristome., Published as part of Yamazaki, Midori, Yamazaki, Kazunori, Rundell, J. & Ueshima, Rei, 2015, Systematic review of diplommatinid land snails (Caenogastropoda, Diplommatinidae) endemic to the Palau Islands. (3) Description of eight new species and two new subspecies of Hungerfordia, pp. 511-538 in Zootaxa 4057 (4) on pages 512-514, DOI: 10.11646/zootaxa.4057.4.3, http://zenodo.org/record/241686

Hungerfordia globosa M. Yamazaki and Ueshima sp. nov. (Figs. 11, 12A) Materials examined. Holotype. Northern part of Ngermalk island (Loc. Nm 5 - 1), Palau. Collected by R. U. on Nov. 13, 2006. UMUTZ-MG-B0521T. Paratype. Ngermalk: same data as holotype, 1 specimen, UMUTZ-MG- B0521. Diagnosis. Shell sinistral, very small for Hungerfordia species, subspherical. Shell color transparant, glossy. Axial ribs partially present on teleoconch; ribs on the apical 3 whorls present, closely spaced, low but distinct; ribs on the last 2 ��� 1 + 3 / 4 whorls absent, but reappearing just behind the thickened lip. Spiral sculpture very weakly developed only on the apical whorls, absent on most whorls. Dorsal grooves weakly developed on the last 3 / 4 whorls, axial, 4���5, widely spaced, shallow; interspace between the dorsal grooves wide, convex. Base convex, with a circum-columellar fold; circum-columellar fold prominently developed on the last 3 / 4 whorls, extended along the columellar side of the thickened lip, narrow, with fine growth lines on its suface. Outer peristome hardly developed. Inner peristome subcircular. Parietal tooth absent. Columellar tooth rather strongly developed. Thickened lip strongly developed behind the aperture, wide, gradually narrowed toward the aperture at the palatal and basal sides; dorsal margin distinct, abruptly expanded from the preceding whorl. FIGURE 11. Hungerfordia globosa sp. nov. (A 1���7, B 1���6) holotype, B0521T; (C) a topotypic paratype, Loc. Nm 5 - 1, B0521. Scale bars, 1mm. All specimens, UMUTZ-MG. Description. Shell (Figs. 11 A���C, 12 A). Shell sinistral, thick, very small for Hungerfordia species, subspherical, penultimate whorl widest when excluding the thickened lip; sides well convex (particulary on the last 3 whorls). Shell color transparant, rather glossy. Whorls 5 ��� 5 + 1 / 4, convex, broadest (in longitudinal width) on the penultimate whorls in dorsal view; last 3 / 4 whorls with axial dorsal grooves, with a circum-columellar fold; last 1 / 8 whorls (including the thickened lip) expanded upward, ascending onto the penultimate whorl in left lateral view. Suture impressed. Constriction located above the parietal/columellar junction of the aperture, less prominent in external view. Tuba 7 / 8 whorls. Protoconch 1 ��� 1 + 1 / 8 whorls, almost smooth, finely and very weakly punctated on lower whorl. Apical septum absent. Axial ribs poorly developed on teleoconch, partially disappearing, almost straight (never folded at the peripheries of the whorls), oblique to the coiling axis: ribs on apical 3 ��� 3 + 1 / 8 whorls of teleoconch present, closely spaced (14���17 ribs in a quarter whorl), weak but distinct, very low, becoming weaker on the penultimate whorl; ribs on the last 2 ��� 1 + 3 / 4 whorls disappearing, completely absent or reduced to faint growth lines, but reappearing just behind the thickened lip (very fine and somewhat distorted by dorsal grooves). Spiral sculpture hardly deveoloped on teleoconch, absent throughout whorls, or very weakly developed only only on the apical 2.5 whorls of teleoconch, very weak and fine when present, sinuous in high magnification, always absent on the last 2.5 whorls. Dorsal grooves developed on the last 3 / 4 whorls, axial, 4���5, moderately impressed, widely spaced, arising at a short distance below the suture, disappearing at the base above the cirum-columellar fold: interspace between the axial grooves wide, well convex. Base convex, with a prominent circum-columellar fold; circum-columellar fold developed on the last 7 / 8 whorls, delimitted by a prominent subspiral groove, initially developed along the columellar side of the thickened lip and almost radially straight in umbilical view, then winding around the columella just behind the thickened lip, with a series of fine and short growth wrinkles around the columella. Umbilicus closed in adult. Aperture hardly protruded, slightly tilted downward against the coiling axis in left lateral view; columellar axis within aperture somewhat tilted outward in front view. Columellar tooth rather strongly developed, emerged at the aperture, but never extended onto the inner peristome, deeply extended inward about 3 / 4 whorls. Parietal tooth absent within the aperture. Peristome single. Outer peristome undeveloped. Inner peristome subcircular of squarish circular, moderately expanded in all around; parietal margin located near or below the middle level of the upper body whorl. Thickened lip strongly developed behind the aperture, wide, expanded beyond the inner peristome in front view, white, lacking of periostracum, with fine growth lines on its outer surface, gradually narrowed toward the aperture (in particulr, at the palatal and basal sides), slightly constricted just behind the inner peristome; dorsal margin rather distinct, abruptly expanded from the preceding whorl at the palatal side; uppermost margin (= suture) ascending onto the penultimate whorl toward the aperture in left lateral view; base also ascending toward the aperture in left lateral view; palatal margin protruded beyond the penultimate whorl in apical view. Dimensions. Shell height 2.6���2.7 mm, shell diameter 1.8���1.9 mm, suture width 1.6 mm, peristome height 1.0 mm. Operculum. Not examined. Penis. Not examined. Radula. Not examined. Distribution and ecology. Endemic to Palau: Known only from the type locality of Ngermalk island. The species inhabits limestone rocks. Remarks. H. globosa is most similar to H. lutea hemilaevis Yamazaki & Ueshima, 2015 in having smooth penultimate whorl without axial ribs, but differs from the latter by the more spherical shell shape, the presence of axial dorsal grooves on the last 3 / 4 whorls, the absence of axial ribs above the aperture, and the dorsal margin of the thickened lip being abruptly expanded from the preceding whorl (Figs. 12 A, B). H. globosa is also similar to H. crassilabris crassilabris (Crosse, 1866), but distinguished from the latter in the following points: the shell shape is more spherical; the axial ribs are almost absent on the last 2 whorls; parietal tooth is never developed within the aperture; the dorsal grooves are not so deeply impressed, but occur in higher number (4���5); the spiral sculpture is absent on the lower whorls (Figs. 12 A, C)., Published as part of Yamazaki, Midori, Yamazaki, Kazunori, Rundell, J. & Ueshima, Rei, 2015, Systematic review of diplommatinid land snails (Caenogastropoda, Diplommatinidae) endemic to the Palau Islands. (3) Description of eight new species and two new subspecies of Hungerfordia, pp. 511-538 in Zootaxa 4057 (4) on pages 534-536, DOI: 10.11646/zootaxa.4057.4.3, http://zenodo.org/record/241686, {"references":["Yamazaki, M., Yamazaki, K. & Ueshima, R. (2015) Systematic review of diplommatinid land snails (Caenogastropoda, Diplommatinidae) endemic to the Palau Islands. (2) Taxonomic revision of Hungerfordia species with low axial ribs. Zootaxa, 3976 (1), 1 - 89.","Crosse, H. (1866) Note sur les Mollusques opercules terrestres des iles Pelew ou Palaos. Journal de Conchyliologie, 14, 346 - 350."]}

Hungerfordia irregularis M. Yamazaki and Ueshima sp. nov. (Figs. 9, 12F) Material examined. Holotype. A limestone outcrop, 1.5 km west of Oikull (Loc. BA 7 - 1), Airai State, Babeldaob island, Palau. Collected by R. U. and M. Y. on Feb. 1, 2010. UMUTZ-B 1093 T. Paratypes. Babeldaob : same data as the holotype, 15 specimens, UMUTZ-B 1093. Diagnosis. Shell sinistral, small for Hungerfordia species, slender conical. Shell color white or reddish brown. Axial ribs on most whorls strongly developed, shortly protruded and somewhat lamellar, irregularly spaced, with or without some riblets at the interspace; interstitial riblets frequently developed between the ribs, also irregularly spaced, variable in the degrees of height and strength; ribs above the aperture always closely spaced, low. Spiral sculpture present throughout teleoconch whorls. Base convex, with a weak circum-columellar fold. Columellar tooth moderately developed. Outer peristome weakly develped at the dorsal margin of the thickened lip or not. Thickened lip developed, rather wide. Description. Shell (Figs. 9 A���D, 12 F). Shell sinistral, thick, small for Hungerfordia species, slender conical, penultimate whorl widest when excluding the thickened lip; sides almost straight or slightly concave on the upper conical whorls, weakly convex on the last 2 whorls. Shell color white or reddish brown. Whorls 6.5���7, convex; last 3 / 4 whorls without dorsal groove, without dorsal fold; last 1 / 8 whorls (including the thickened lip) expanded upward, ascending onto the penultimate whorl in left lateral view. Suture impressed. Constriction located above the parietal/columellar junction of the aperture, associated with abrupt changes of the ribbing pattern, internally without palatal plica except a low and axial crest caused by the constriction, without parietal tooth. Tuba 7 / 8 whorls. Protoconch 1���1.25 whorls, almost smooth, weakly and finely punctated on the lower whorls. Apical septum absent. Axial ribs developed on teleoconch, distinct throughout whorls, almost straight (never folded at the peripheries of the whorls), oblique to the coiling axis; ribs on the first 1 / 4 whorls of the teleoconch absent; ribs on the following 1 / 3 ��� 1 / 4 whorls of initial teleoconch weak, closely spaced, very low; ribs on subsequent whorls (ribs on most whorls except above the aperture) strongly developed, distinctly (but shortly) protruded, somewhat lamellar, irregularly spaced, with or without some riblets at the interspace; interstitial riblets frequently developed between the ribs, also irregularly spaced, always somewhat lower (less protruded) than the ribs, variable in the degrees of height and strength (variable from highly protruded like the ribs to very weak and almost reduced to FIGURE 9. Hungerfordia irregularis sp. nov. (A 1���7) A topotypic paratype, Loc. BA 7 - 1, B 1093 a: (A 7) sculptures on the penultimate and the higher 2 whorls, showing the irregularly spaced axial ribs and riblets, note that the interstitial riblets are variable in the degrees of development; (B 1���5) holotype, B 1093 T; (C, D) topotypic paratypes, B0476b���c; (E 1���2) outer surface, and (E 3���4) inner surface of operculum, a topotypic paratype, B 1093. Scale bars, 1mm (A���D), 0.5 mm (E). All specimens, UMUTZ-MG. growth lines), occasionally undeveloped between some ribs; ribs above the aperture (ribs on the last 1 to last 3 / 4 whorls around the constriction) abruptly becoming closely spaced and low, never protruded. Spiral sculpture present on teleoconch, fine, prominent throughout whorls, sinuous in high magnification. Base rather convex, with a circum-columellar fold; circum-columellar fold weakly developed around the columella on the last 3 / 4 whorls, with a series of fine and short growth wrinkles in umbilical view. Umbilicus closed in adult. Aperture not protruded, tilted downward against the coiling axis in left lateral view; columellar axis within the aperture almost vertical. Columellar tooth moderately developed, emerging at the aperture, but never extended onto the inner peristome. Peristome single or double. Outer peristome weakly developed at the dorsal margin of the thickened lip or not. Inner peristome subcircular or squarish circular, moderately expanded, but less expanded at the upper palatal (=angular) side; parietal margin located below the middle level of the upper body whorl. Thickened lip developed behind the aperture, wide, expanded beyond the inner peristome at the upper palatal and columellar sides in front view, white, lacking of periostracum, with a series of fine growth lines on the outer surface, gradually narrowed toward the aperture in umbilical view; dorsal margin distinct, abruptly expanded from the preceding whorl; uppermost margin (=suture) ascending onto the penultimate whorl in left lateral view; base extended almost horizontally toward the aperture in left lateral view; palatal margin weakly protruded beyond the penultimate whorl in apical view. Dimensions. Shell height 2.8���3.2 mm, shell diameter 1.7 ���2.0 mm, suture width 1.4���1.6 mm, peristome height 0.9���1.1 mm. Operculum (Figs. 9 E 1���4). Operculum corneous, multispiral, circular, semi-transparent, amber-colored, thin, flat; outer surface smooth; inner surface with a low, thin, arcuate ridge near the columellar margin. Penis. Penis absent. Radula. Not examined. Distribution and ecology. Endemic to Palau: Known only from a limestone outcrop near Oikull, Babeldaob island. The species inhabits limestone rocks. Remarks. H. irregularis is distinguished from other low-ribbed Hungerfordia species by having irregularly spaced ribs and riblets, the slender shell shape, and the absence of dorsal fold and groove on the last 3 / 4 whorls. Etymology. The specific name, derived from Latin irregularis, refers to the irregularly spaced ribs and riblets., Published as part of Yamazaki, Midori, Yamazaki, Kazunori, Rundell, J. & Ueshima, Rei, 2015, Systematic review of diplommatinid land snails (Caenogastropoda, Diplommatinidae) endemic to the Palau Islands. (3) Description of eight new species and two new subspecies of Hungerfordia, pp. 511-538 in Zootaxa 4057 (4) on pages 530-532, DOI: 10.11646/zootaxa.4057.4.3, http://zenodo.org/record/241686

Hungerfordia ngereamensis M. Yamazaki and Ueshima sp. nov. (Figs. 2, 12 O) Material examined. Holotype. Chongelungel, south of Ngeream (Loc. BA 4 - 2), Airai state, Babeldaob island, Palau. Collected by R. U. and M. Y. on July 7, 2006. UMUTZ-MG-B0293T. Paratypes. Babeldaob: Loc. BA 3 -1, 10 specimens, 27 /VI/ 2005, UMUTZ-MG-B0172; Loc. BA 4 -2, 26 specimens, 7 /VII/ 2006, UMUTZ-MG-B0293. Diagnosis. Shell sinistral, large to medium-size for Hungerfordia species, slender conical. Shell color white, upper whorls frequently yellow or yellowish brown. Apical septum absent. Axial rib on most whorls highly protruded as spines, very widely spaced, strongly folded at the peripheries of the whorls; spiny projections of the ribs moderately high, rather narrow, arising at some distance below the suture; ribs above the aperture abruptly becoming closely spaced, with greatly reduced spiny projections, less folded. Interspace between the spiny ribs with some very weak and very short growth wrinkles below the suture on the lower whorls. Spiral sculptures present throughout teleoconch. Aperture weakly protruded. Columellar tooth very weakly emerged within the aperture in front view, but strongly developed deeply inward. Peristome triple, with doubled inner peristomes, stout. Outer peristome strongly developed, widely expanded. Inner peristome double, also expanded; distal inner peristome strongly developed just behind the proximal inner peristome, gradually narrowed toward the aperture; proximal inner peristome always narrower than the distal one in front view. Interspace between the outer and inner peristomes wide; outer surface with a series of fine and very low riblets at the abapertural half (near the outer peristome), almost smooth on the adapertural half (just behind the inner peristomes). Description. Shell (Figs. 2 A���F, 12 O). Shell sinistral, large to medium-size for Hungerfordia species, conical or fusiform, penultimate whorl widest (slightly wider than the last whorl) or last 2 whorls almost equally widest when removing the expanded ribs and peristomes; sides almost straight. Shell color white, upper whorls occasionally yellow. Whorls 6���7, slightly convex; last 1 / 8 whorls (including the interspace between the outer and inner peristomes) expanded upward and ascending onto the penultimate whorl in left lateral view. Suture weakly impressed. Constriction located above the parietal/columellar junction of the inner peristome, associated with abrupt changes of the ribbing pattern, internally without palatal plica except a low axial-crest caused by the constriction. Tuba 7 / 8 whorls. Protoconch 1 + 1 / 8 ��� 1 + 1 / 4 whorls, almost smooth, very weakly and finely punctated. Apical septum absent. Axial ribs strongly developed on the teleoconch, oblique to the coiling axis, highly protruded as spines on most whorls: ribs on the first 1 / 3 ��� 1 / 2 whorls of the initial teleoconch weak, almost straight (not folded at the peripheries of the whorls), initially closely spaced and very low, gradually becoming widely spaced and stronger; ribs on the subsequent whorls except above the aperture (ribs on most whorls except above the aperture) strong, highly protruded as spines, very widely spaced (9���11 ribs in a whorl), not synchronized with those on the previous whorls, strongly folded at the peripheries of the whorls, strongly elevated by axial protrusions of shell wall (along the ribs) on the last 3 / 4 whorls; upper margin of the ribs above the spiny projections (between the suture and spines) very low, never protruded, indistinct; spiny projections of the ribs high, pointed, folded at the peripheries of the whorls, trough-shaped, rather narrow, arising at some distance below the suture (arising near the peripheries of the whorls), upper margin folded over the lower margin, lower margin gradually decreased in height and extended below to the suture of the next whorl, with fine spiral sculptures on the outer surface, protruded upward on most whorls, becoming protruded laterally on the last 5 / 8 whorls; ribs above the aperture (ribs on the last 1 to last 3 / 4 whorls around the constriction) abruptly becoming low and closely spaced, with greatly reduced spiny projections (but still retaining very low spiny projections), less folded than those on the preceding whorl, becoming much less folded toward the aperture, partially hidden by the inner peristome. Interspace between the spiny ribs smooth on the upper whorls, with some growth wrinkles below the suture on the last 1.25���2.25 whorls; growth wrinkles developed below the suture on the lower whorls, closely spaced, short, very low, initially very weak, becoming rather prominent on the last 3 / 4 whorls. Spiral sculptures present throughout the teleoconch, fine, rather prominent, sinuous in high magnification, extending on the outer surface of the spiny ribs and outer peristome. Base rather convex, without basal fold; interspace between the ribs with a series of fine and short growth wrinkles around the columella. Axial protrusions of shell wall (along the ribs) on the last 3 / 4 whorls strongly developed, hidden by the spiny ribs in fresh specimens (becoming obvious after removal of the spiny projections of the ribs), associated with prominent axial depressions between them; abapertural side slightly or hardly convex in umbilical view; adapertural side well convex, wider than the abapertural side in umbilical view. Umbilicus closed in adult. Aperture hardly or slightly tilted against the coiling axis (in left lateral view), slightly protruded from the previous whorl; columellar axis within the aperture almost vertical or slightly oblique in front view. Columellar tooth present, very weakly appearing within the aperture in front view, but strongly developed deeply inside the aperture, never extended onto the inner peristome. Peristome triple, with an outer peristome and doubled inner peristomes. Outer peristome strongly developed, widely expanded, but hardly or only slightly expanded beyond the inner peristome in front view, obtusely angulate at the palatal side, frequently with a rounded baso-columellar edge, reflected backward, abruptly expanded from the preceding whorl, with concentric growth riblets or growth lines on the inner surface, with fine spiral sculptures on the outer surface. Inner peristome strongly protruded (separated) from the outer one, double with distal and proximal inner peristomes; distal inner peristome strongly developed just behind the proximal one, widely expanded, shortly expanded beyond the proximal one in front view; proximal inner peristome squarish circular, always narrower and weaker than the distal one, rather widely expanded, parietal margin almost straight and located near or below the middle level of the upper body whorl. Interspace between the outer and inner peristomes wide, with a series of fine riblets on the outer surface near the outer peristome, rather smooth near the inner peristome; uppermost margin (= suture) ascending onto the penultimate whorl toward the aperture in left lateral view; base extended almost horizontally in left lateral view. Dimensions. Shell height 4.5���5.3 mm, diameter 2.6���3.6 mm, suture width 1.9���2.1 mm, aperture height 1.8��� 2.2 mm. Operculum (Figs. 2 G 1���4). Operculum corneous, multispiral, circular, transparent, amber colored, thin, flat; outer surface smooth; inner surface with a very weak arcuate ridge near the columellar margin. Penis. Penis absent. Radula. Not examined. Distribution and ecology. Endemic to Palau: Southern limestone coast of Babeldaob island. This species inhabits limestone rocks. Remarks. H. ngereamensis is most similar to H. triplochilus Yamazaki & Ueshima, 2013 (Figs. 12 O, P), but differs in the following points: the axial protrusions of shell wall along the ribs are strongly developed on the last 3 / 4 whorls; the riblets between the inner and outer peristomes are less developed, much lower than the latter species; the distal inner peristome is always wider and more strongly developed than the proximal inner peristome; the shell is broader and less slender; the distribution is endemic to Babeldaob island. H. ngereamensis differs from a sympatric congener, H. echinata tubulispina Yamazaki & Ueshima, 2013, in the following points: the spiny projections of the ribs are wider, never tubular; the inner peristome is double; the interspace between the outer and inner peristomes is wider; the axial ribs above the aperture have very short spiny projections. H. ngereamensis also differs from H. expansilabris Yamazaki & Ueshima, 2013, which is distributed in the same area of Babeldaob island, in the following points: the outer and inner peristomes are not so widely expanded as the latter species, more robust; the columellar tooth is present; the distal inner peristome is always wider and more strongly developed than the proximal inner peristome; the axial ribs above the aperture have short spiny projections. Etymology. The specific name is derived from the type locality. FIGURE 2. Hungerfordia ngereamensis sp. nov. (A 1���6) A topotypic paratype, Loc. BA 4 - 2, B0293a; (B 1���2) a topotypic paratype after removal of the spiny projections of the axial ribs, B0293b: (B 1) umbilical view, only the last whorl shown, note that the axial protrusions of the shell wall are strongly developed along the ribs; (B 2) oblique dorsal view, showing the axial protrusions of the shell wall on the last 3 / 4 whorls; (C 1���2) a yellowish and subfusiform specimen, a topotypic paratype, B0293c: (C 2) sculptures on the upper body whorl, showing the reduced axial ribs above the aperture; (D) columellar tooth, dorsal view of a partially opened shell, a topotypic paratype, B0293d; (E 1���6) holotype, B0293T; (F) a worn specimen, only the last whorl shown, note the axial protrusions of the shell wall which are usually hidden by spiny ribs in fresh specimens (see Fig. E 6), a topotypic paratype, B0293e; (G 1���2) outer surface, and (G 3���4) inner surface of operculum, a topotypic paratype, B0293. Scale bars, 1mm (A���F), 0.5 mm (G). All specimens, UMUTZ-MG., Published as part of Yamazaki, Midori, Yamazaki, Kazunori, Rundell, J. & Ueshima, Rei, 2015, Systematic review of diplommatinid land snails (Caenogastropoda, Diplommatinidae) endemic to the Palau Islands. (3) Description of eight new species and two new subspecies of Hungerfordia, pp. 511-538 in Zootaxa 4057 (4) on pages 514-516, DOI: 10.11646/zootaxa.4057.4.3, http://zenodo.org/record/241686, {"references":["Yamazaki, K., Yamazaki, M. & Ueshima, R. (2013) Systematic review of diplommatinid land snails (Caenogastropoda, Diplommatinidae) endemic to the Palau Islands. (1) Generic classification and revision of Hungerfordia species with highly developed axial ribs. Zootaxa, 3743 (1), 1 - 71. http: // dx. doi. org / 10.11646 / zootaxa. 3743.1.1"]}

Hungerfordia elegantissima anomphala M. Yamazaki and Ueshima subsp. nov. (Figs. 7, 12T) Material examined. Holotype. Metkerel a Chudel (Loc. Mc 8 - 5), eastern part of Mecherchar island, Palau. Collected by R. U. and M. Y. on Sept. 28, 2006. UMUTZ-MG-B 1285 T. Paratypes: Mecherchar: Loc. Mc 2 -1, 36 specimens, 8 /XI/ 2004, UMUTZ-MG-B0080; ditto, 17 specimens, 1 /VII/ 2005, UMUTZ-MG-B0232; Loc. Mc 8 -5, 6 specimens, 28 /II/ 2011, UMUTZ-MG-B 1285. Diagnosis. Shell similar to the nominal subspecies, but differs in the following features. Umbilicus closed in adult. Last 1 / 16 ��� 1 / 8 whorls attached to the previous whorl, never decoiled downward. Shell more slender. Spiny ribs above the aperture somewhat shorter than those on the preceding and following whorls. Distribution endemic to Mecherchar island. Description. Shell ( Figs. 7 A���C, 12 T). Shell sinistral, small for Hungerfordia species, slender conical, penultimate whorl widest (slightly wider than the last whorl) when removing the spiny ribs and the peristomes. Shell diameter including the spiny ribs almost equal to or slightly smaller than the shell height. Shell color yellow or white. Whorls 5.5���6, convex, obtusely angulate at the periphery on the penultimate whorl; last 1 / 16 ��� 1 / 8 whorls (behind the outer peristome) attached to the previous whorl, never decoiled downwards like the nominal subspecies, never ascending onto the penultimate whorl in left lateral view. Suture impressed. Constriction located at a half whorl behind the aperture, hardly recognizable in external view, but associated with very weak reduction of the spiny ribs in its length (height), internally without palatal plica except a low axial-crest caused by the constriction. Tuba a half whorl. Protoconch 1 ��� 1 + 1 / 8 whorls, almost smooth, finely and very weakly punctated on FIGURE 7. Hungerfordia elegantissima anomphala subsp. nov. (A 1���9) A paratype, Loc. Mc 2 - 1, B0080a: (A 9) left lateral view of the last 2 whorls, note that the last 1 / 8 whorls are attaching to the penultimate whorl (never decoiled downward like the nominal subspecies); (B 1���6) holotype, arrowheads indicate the position of the constriction, B 1285 T; (C) a partially broken specimen after removal of the spiny ribs, apical view, showing the position of the constriction (arrowhead), a paratype from Loc. Mc 2 - 1, B0080b; (D 1���2) outer surface, and (D 3���4) inner surface of operculum, a paratype from Mc 3 - 3, B0232. Arrowheads indicate the positions of the constriction. Scale bars, 1mm (A���C), 0.5 mm (D). All specimens, UMUTZ-MG. lower whorl. Apical septum absent. Axial rib strongly developed on teleoconch, oblique to the coiling axis, strongly protruded as spines on most whorls: ribs on the initial 1 / 4 whorls of the teleoconch weak, low, moderately spaced, almost straight (not folded at the peripheries of the whorls); ribs on the subsequent whorls highly protruded as spines, strongly folded at the peripheries of the whorls, widely spaced (10���12 ribs in a whorl); upper margin of the ribs above the spiny projections (between the suture and spines) low, very slightly elevated or indistinct; lower (basal) margin of the ribs below the spines prominently protruded at the base of the last whorl, gradually decreased in the height towards the umbilicus; spiny projections of the ribs very long, weakly pointed, slender, strongly folded, obliquely depressed, narrow, arising at some distance below the suture (near the peripheries of the whorls), abruptly reduced in height just below the peripheries of the whorls (but still shortly protruded and extended to the suture of the next whorl), protruded slightly upward on the upper whorls, becoming protruded horizontally or downward on the last whorl, upper margin folded over the lower margin, with fine spiral sculptures on the outer surface; ribs above the aperture (ribs on the last 1 to last 2 / 3 whorls) showing a slight change of the ribbing pattern, though still retaining developed spiny projections, but with shorter spiny projections and slightly more closely spaced than those on the preceding and following whorls. Interspace between the spiny ribs almost smooth, without prominent axial sculpture except some faint growth lines on the last 1 / 2 whorls. Spiral sculptures present throughout the teleoconch, fine, weak, extended onto the outer surface of the spiny ribs and the outer peristome, sinuous in high magnification. Base slightly convex, without basal keel, with shortly protruded axial ribs. Umbilicus closed in adult. Axial protrusion of shell wall (along the spiny ribs) hardly developed. Aperture evidently tilted downward against the coiling axis (in left lateral view), shortly protruded downward from the upper body whorl. Columellar tooth very weak, reduced to a very low swelling of columella, which is gradually thickened from the parietal roof, recessed inside the aperture, short. Peristome double. Outer peristome widely expanded beyond the inner one at the palatal, basal and lower columellar sides, abruptly reduced at base of the columellar side to form a rounded baso-columellar edge, undeveloped on the parietal and upper columellar sides, thin, with fine and concentric growth lines on the inner surface. Inner peristome squarish circular, moderately expanded at the lower palatal, basal, and columellar sides, hardly expanded (in front view) but slightly extended forward at the parietal side, only slightly protruded from the outer peristome at the palatal and basal sides, elected and free from the base of the last whorl at the columellar side; parietal margin shortly separated downward from the upper body whorl. Interspace between the inner and outer peristomes very narrow. Dimensions. Shell height 3.5���4.1 mm, diameter 3.1 ���4.0 mm, suture width 1.5���1.6 mm, apertural height 1.1��� 1.2 mm. Operculum (Figs. 7 D 1���4). Operculum corneous, multispiral, circular, rather thin, transparent, amber colored, flat or slightly concave; outer surface smooth; inner surface with a strongly developed arcuate ridge near the columellar margin; arcuate ridge very thick, highly protruded, somewhat wavy at the edge, lateral processes much less developed than the nominal subspecies. Penis. Penis absent. Radula. Not examined.. Distribution and ecology. Endemic to Palau: Mecherchar island. The species inhabits limestone rocks. Remarks. H. elegantissima anomphala differs from the nominal subspecies in the following points (Figs. 12 T, U): the umbilicus is closed in adult; the last 1 / 8 whorls is attached to the pervious whorl, never decoiled downward, although the aperture itself is shortly protruded downward; the aperture is not so strongly turned downward; the shell is more slender; the spiny ribs above the aperture are shorter and more closely spaced than those on the preceding and following whorls. The distribution of H. elegantissima anomphala is endemic to Mecherchar island. Etymology. The subspecific name, derived from Latinized Greek, a + omphalus, refers to the closed umbilicus., Published as part of Yamazaki, Midori, Yamazaki, Kazunori, Rundell, J. & Ueshima, Rei, 2015, Systematic review of diplommatinid land snails (Caenogastropoda, Diplommatinidae) endemic to the Palau Islands. (3) Description of eight new species and two new subspecies of Hungerfordia, pp. 511-538 in Zootaxa 4057 (4) on pages 525-527, DOI: 10.11646/zootaxa.4057.4.3, http://zenodo.org/record/241686

YAMAZAKI, KAZUNORI, YAMAZAKI, MIDORI, UESHIMA, REI (2013): Systematic review of diplommatinid land snails (Caenogastropoda, Diplommatinidae) endemic to the Palau Islands. (1) Generic classification and revision of Hungerfordia species with highly developed axial ribs. Zootaxa 3743 (1): 1-71, DOI: http://dx.doi.org/10.11646/zootaxa.3743.1.1, URL: http://dx.doi.org/10.11646/zootaxa.3743.1.1

Hungerfordia expansilabris K. Yamazaki and Ueshima sp. nov. (Figs. 9C, 14, 29B) Material examined. Holotype. East of Pkulaklim islet (Loc. BA 5-5), Airai state, Babeldaob island, Palau. Collected by R. U. and K. Y. on Nov. 12, 2006. UMUTZ-MG-B0480 T. Paratypes. Babeldaob: Loc. BA 6-1, 13 specimens, 21/X/2007, UMUTZ-MG-B0667; Loc. BA 8-3, 6 specimens, 1/III/2011, UMUTZ-MG-B1309; Loc. BA 8-4, 3 specimens, 1/III/2011, UMUTZ-MG-B1322. Pkulaklim islet: Loc. BA 5-4, 1 specimen, 12/XI/2006, UMUTZ-MG-B0472; Loc. BA 5-5, 6 specimens, 12/XI/2006, UMUTZ-MG-B0480; ditto, 58 specimens, 1/III/ 2011, UMUTZ-MG-B1299; Loc. BA 6-2, 21 specimens, 21/X/2007, UMUTZ-MG-B0672. Diagnosis. Shell sinistral, medium-size for Hungerfordia species, slender conical. Shell color white or yellow. Constriction located above the aperture, followed by an abrupt swelling of whorl, associated with abrupt reduction of the spiny ribs. Axial ribs on most whorls highly protruded as spines, strongly folded at the periphery, widely spaced; upper margin of the ribs above the spiny projections (between the suture and spines) low, indistinct; spiny projections of the ribs very high, rather narrow, arising at some distance below the suture, folded at the periphery; ribs above the aperture very low and closely spaced, almost hidden by the expanded inner peristome. Spiral sculptures fine. Axial protrusions of shell wall (along the ribs) strongly developed on the last 3/4 whorls. Aperture strongly protruded, hardly tilted. Columellar tooth absent. Peristome usually double, very widely and extremely expanded, thin, fragile; outer peristome very widely expanded beyond the inner one at the palatal and basal sides, with rounded upper-palatal and baso-columellar edges, columellar margin strongly folded backward and covering the umbilicus; inner peristome very widely expanded all around, circular, parietal margin expanded beyond the middle level of the upper body whorl. Interspace between the outer and inner peristomes wide, abruptly expanded towards the aperture, almost smooth on its outer surface. Description. Shell (Figs. 14A–H, 29B). Shell sinistral, medium-size for Hungerfordia species, slender conical, last whorl widest when removing the expanded ribs and peristomes. Shell color semi-transparent, white or light yellow, apical whorls occasionally yellow with the color becoming deeper toward the apex. Whorls 6–7, slightly convex; last 1/8 whorls (including interspace between the outer and inner peristomes) expanded upward and ascending onto the penultimate whorl in left lateral view. Suture weakly impressed. Constriction located above the parietal/columellar junction of the aperture, followed by an abrupt swelling of whorl, associated with abrupt reduction of the spiny ribs, internally without palatal plicae except a low axial-crest caused by the constriction. Tuba 7/8 whorls. Protoconch 1.5–2 whorls, apical whorl smooth, lower whorl faintly punctated below the suture. Apical septum absent. Axial ribs developed on the teleoconch, oblique to the coiling axis, strongly protruded as spines on most whorls; ribs on the first 1/2 to 3/4 whorls of the teleoconch low, closely spaced, not folded, gradually becoming higher and widely spaced; ribs on the subsequent whorls (except above the aperture) highly protruded as spines, strongly folded at the periphery, widely spaced (10–12 ribs in a whorl), not synchronized with those on the previous whorls; upper margin of the ribs above the spiny projection (between the suture and spines) low, indistinct or faintly elevated; spiny projections of the ribs very high, weakly pointed, narrow, arising at some distance below the suture (arising near periphery of a whorl), folded at the periphery, trough-shaped, protruded slightly upward or laterally on upper whorls, becoming to protruded rather downward on the last half whorl, upper margin weakly folded over the lower margin, lower margin extended below to the suture of the next whorl, with fine spiral sculptures on the outer surface; ribs above the aperture (ribs on the last 1 to the last 3/4 whorls around the constriction) abruptly becoming closely spaced and very low, without spiny projections, gradually becoming less folded at the periphery, almost hidden by the expanded inner peristome. Interspace between the spiny ribs without prominent axial sculpture on upper whorls, with coarse growth wrinkles below the suture on the last 3/4 whorls. Spiral sculptures present throughout the teleoconch, fine, rather prominent, expanded on the outer surface of the spiny ribs and outer peristome, sinuous in high magnification. Base convex, without basal keel. Umbilicus closed in adult by columellar reflection of the outer peristome not by narrowing of umbilical area. Axial protrusions of shell wall (along the ribs) very strongly developed on the last 3/4 whorls, hidden by the spiny ribs, strongest just after the constriction, becoming weaker toward the aperture, with rather deep axial depressions between them, adapertural side steep, abapertural side strongly convex and much wider than the adapertural one in umbilical view. Aperture hardly tilted against the coiling axis, strongly protruded; columellar axis within the aperture almost vertical. Columellar tooth usually absent, occasionally appeared as a faint columellar swelling. Peristome usually double, rarely triple with doubled inner peristome, very widely and extremely expanded; outer peristome thin, fragile, very widely expanded beyond the inner one at the palatal and basal sides, abruptly narrowed towards angular and columellar side to form rounded upper-palatal and baso-columellar edges, reflected backward, columellar margin strongly folded backward and covering the umbilicus, smooth and polished on the inner surface; inner peristome strongly protruded from the outer one, extremely widely expanded all around, especially on columellar and parietal sides, thin, very fragile, circular, parietal margin frequently expanded beyond the middle level of the upper body whorl; distal inner peristome usually undeveloped, weak and narrower than the proximal one when present. Interspace between the outer and inner peristomes wide, strongly expanded towards the aperture, outer surface almost smooth; uppermost margin continuously ascending toward the aperture in left lateral view, frequently with a series of very short growth wrinkles; base extended almost horizontally and then abruptly expanded toward the aperture in left lateral view. Dimensions. Shell height 4.7–5.3 mm, diameter 3.2–3.8 mm, suture width 1.5–2.1 mm, peristome height 2.1– 2.6 mm. Operculum (Figs. 14I 1 –I 4). Operculum corneous, multispiral, circular, thin, amber colored, transparent, flat or weakly concave; outer surface smooth; inner surface with a low, C-letter shaped, arcuate ridge near the columellar margin. Penis. Penis absent. Radula (Figs. 9C). Radula of specialized type of taenioglossate; ribbon very long. Central tooth very large, with a very wide, large, and blunt central cusp, with a pair of small lateral cups, basal plate longitudinally elongated and strongly constricted at the base. Lateral teeth also large, with a large, very wide and blunt major cusp, with or without a vestigial inner cusp, without outer cusp, basal plate obliquely elongated. Inner marginal teeth shorter than the lateral teeth, elongated, with a large major cusp, with two slender inner cusps and a small outer cup. Outer marginal teeth with a large, wide and blunt major cusp, with two slender inner cusps, without outer cusp. Distribution and Ecology. Endemic to Palau: Babeldaob island (southern coastal limestone area) and its satellite islet, Pkulaklim. The species inhabits vertical walls of large limestone rocks or cliffs, occasionally on limestone rubbles. Remarks. H. expansilabris is most similar to H. triplochilus, but differs from the latter by the much wider and fragile peristomes, the absence of a columellar tooth, the smooth interspace between the outer and inner peristomes, the absence or poor development of the distal inner peristome, and the axial protrusions of shell wall on the last 3/4 whorls being strongly developed. H. expansilabris is also distinguished from a sympatric congener, H. echinata, by having more widely spaced ribs, more slender spire, much wider and fragile peristomes, and the absence of a columellar tooth. Etymology. The specific name, derived from Latin expansus (= expanded) and labrum (= lip), refers to the very widely expanded inner peristome., Published as part of YAMAZAKI, KAZUNORI, YAMAZAKI, MIDORI & UESHIMA, REI, 2013, Systematic review of diplommatinid land snails (Caenogastropoda, Diplommatinidae) endemic to the Palau Islands. (1) Generic classification and revision of Hungerfordia species with highly developed axial ribs , pp. 1-71 in Zootaxa 3743 (1) on pages 27-28, DOI: 10.11646/zootaxa.3743.1.1, http://zenodo.org/record/5271036

Hungerfordia echinata tubulispina K. Yamazaki and Ueshima subsp. nov. (Figs. 9F, 17, 29E) Materials examined. Holotype. Limestone outcrop along a small river, north of Ngeream (Loc. BA 3-1), Airai state, Babeldaob island, Palau. Collected by R. U. and K. Y. on June 27, 2005. UMUTZ-MG-B0169 T. Paratypes. Same data as holotype, 90 specimens, UMUTZ-MG-B0169. Diagnosis. Shell generally similar to H. echinata echinata but differs by the following features: axial ribs more widely spaced than the nominal subspecies, upper margin of ribs above the spiny projections (between the suture and spines) indistinct or less protruded; spiny projections of the ribs almost tubular, narrower; growth wrinkles between the spiny ribs below the suture weaker. Distribution restricted to a limestone outcrop which is isolated from distribution range of the nominal subspecies. Description. Shell (Figs. 17A–I, 29E). Shell sinistral, medium-size to large for Hungerfordia species, fusiform or conical, last whorl or penultimate whorl widest when removing the expanded ribs and peristomes. Shell color white, occasionally colored yellowish brown around the closed umbilicus, upper whorls usually colored yellow or light brown with the color becoming deeper toward the apex or the apical septum (apical whorls above the apical septum always uncolored). Whorls 6.5–7.5, convex or obtusely angulate at the periphery of the spiny ribs; last 1/8 whorls (including interspace between the outer and inner peristomes) expanded upward and ascending onto the penultimate whorl in left lateral view. Suture impressed. Constriction located above the parietal/columellar junction of the aperture, followed by a weak swelling of whorl, associated with abrupt reduction of the spiny ribs, internally without palatal plica except for a low axial-crest caused by the constriction. Tuba 7/8 whorls. Protoconch 1+3/4 to 2 whorls, apical whorl almost smooth, lower whorl faintly punctated below the suture. Apical septum present or absent. Axial ribs developed on the teleoconch, oblique to the coiling axis, strongly protruded as tubular spines on most whorls; ribs on first 1/3 whorls of teleoconch low, closely spaced, not folded; ribs on the subsequent whorls (except above the aperture) highly protruded as tubular spines, widely spaced (10–13 ribs in a whorl), not synchronized with those on previous whorls, strongly curled at the periphery; upper margin of the ribs above the tubular spines (between the suture and spines) low, indistinct or hardly protruded; spiny projections on the ribs almost tubular, with a blunt and opened tip, with a narrow longitudinal slit at the adapertural side, very high, slender, almost straight, narrow, arising near the periphery (at some distance below the suture), disappeared just below the periphery, lower margin overlapped with or close to the upper margin at the base, protruded rather upward on most whorls, becoming protruded laterally to rather downward on last 3/4 whorls, with fine spiral sculptures on the outer surface; ribs above the aperture (ribs on last 1 to last 3/4 whorls around the constriction) abruptly becoming very closely spaced and very low, without any spiny projections, becoming less folded at the periphery toward the constriction. Interspace between the spiny ribs with fine growth wrinkles; upper growth wrinkles developed below the suture on lower whorls, low, fine; lower growth wrinkles occasionally present above the suture of the last 1–2 whorls, very weak, fine, rather long (extending near the periphery), almost vertical. Spiral sculptures present throughout teleoconch, fine, weak, extending on outer surface of the tubular spines and outer peristome, sinuous in high magnification. Base convex, rather smooth, without basal keel, occasionally with slightly elevated axial ribs. Axial protrusions of shell wall (along the spiny ribs) weakly developed on last 3/4 whorls, hidden by the tubular spines (visible after removal of the tubular spines), abapertural side steep and narrow, adapertural side convex and much wider than the abapertural one in umbilical view. Umbilicus closed in adult. Aperture hardly tilted against the coiling axis, slightly or hardly protruded; columellar axis within aperture almost vertical. Columellar tooth reduced or very weakly developed deeply inside the aperture. Peristome double, expanded; outer peristome more or less expanded beyond the inner one at the palatal and basal sides, with rounded palatal and baso-columellar edges, weakly folded at the periphery or not, weakly reflected, with fine and concentric growth wrinkles on the inner surface; inner peristome pentagonal or squarish circular, occasionally with a rounded angular edge, expanded all around, hardly reflected, strongly protruded from outer one, angular and parietal margin located slightly below middle level of the upper body whorl. Interspace between the inner and outer peristomes rather wide, almost smooth or with a series of fine and very low growth lines on the outer surface, gradually constricted at the basal and palatal sides, then gradually expanded toward the aperture; uppermost margin ascending onto the penultimate whorl toward the aperture in left lateral view. Dimensions. Shell height 4.7–5.7 mm, diameter 3.8–4.4 mm, suture width 2.2–2.9 mm, peristome height 1.8– 2.1 mm. Operculum (Figs. 17J 1 –J 4). Operculum corneous, multispiral, circular, thin, transparent, amber colored, almost flat or weakly concave; outer surface smooth; inner surface with a low, weak, arcuate ridge neat the columellar margin. Penis. Penis absent. Radula (Fig. 9F). Radula of specialized type of taenioglossate; ribbon very long. Central tooth very large, with a very wide, large, and blunt central cusp, with or without a pair of vestigial lateral cups, basal plate longitudinally elongated and strongly constricted at the base. Lateral teeth also large, with a large, very wide and blunt major cusp, without inner nor outer cusp, basal plate obliquely elongated. Inner marginal teeth shorter than the lateral teeth, elongated, with a large major cusp, with two slender inner cusps, with a small outer cup. Outer marginal teeth shorter than the inner marginal, with a large, wide and blunt major cusp, with two slender inner cusps, without outer cusp. Distribution and ecology. Endemic to Palau: only known from the type locality, a narrow limestone outcrop isolated from distribution range of the nominal subspecies. The subspecies inhabits limestone rocks. Remarks. H. echinata tubulispina differs from the nominal subspecies by the spiny projection of the ribs being tubular, the upper margin of the ribs (between the suture and the tubular projections) being less flared or indistinct, the more widely spaced axial ribs, and the axial protrusions on the last 3/4 whorl being more strongly developed. Etymology. The subspecific name, derived from Latin tubulus (=tubular) and spina (=spine), refers to the tubular spines on the axial ribs., Published as part of YAMAZAKI, KAZUNORI, YAMAZAKI, MIDORI & UESHIMA, REI, 2013, Systematic review of diplommatinid land snails (Caenogastropoda, Diplommatinidae) endemic to the Palau Islands. (1) Generic classification and revision of Hungerfordia species with highly developed axial ribs , pp. 1-71 in Zootaxa 3743 (1) on pages 35-37, DOI: 10.11646/zootaxa.3743.1.1, http://zenodo.org/record/5271036