Kleokyphus cowiei, Sartori, André F., Gargominy, Olivier & Fontaine, Benoît, 2014

Sartori, André F., Gargominy, Olivier & Fontaine, Benoît, 2014, Radiation and decline of endodontid land snails in Makatea, French Polynesia, Zootaxa 3772 (1), pp. 1-68 : 33-38

Kleokyphus cowiei sp. nov.

Figures 20 D; 22; 35M; 37E; 39.

Examined material (188 specimens). Holotype: MNHN 25580, Mk12. Paratypes: MNHN 25581, 8 shells, Mk12. Additional material: 173 shells, Mk12; 6 shells, Mk08.

Type locality. Approximately 1 km north of Temao port, Makatea (15.81638°S; 148.27639°W). Large fault in the cliff, shaded. Ferns. Limestone, alt. 10 m; station Mk12.

Diagnosis. Shell less than 5 mm in diameter, robust, depressed, usually without flammulations, seldom flammulated; teleoconch sculptured by broad, low, relatively crowded primary axial ribs (> 90 ribs on body whorl); umbilicus U-shaped; palatal wall with 4 barriers and commonly 1 trace; parietal wall with 3 barriers.

Description. Shell depressed, white to fawn, usually without flammulations, seldom with faint, regularly spaced, amber flammulations. Shell wall moderately robust, opaque; periostracum adherent, shiny. Apex and spire elevated; later whorls descending more rapidly. Apical and umbilical sutures impressed; whorls and periphery rounded; basal and columellar walls transitioning smoothly. Transition between protoconch and teleoconch indistinct. Protoconch sculptured by fine axial riblets, initially with interspaces 10–15 times their width, undifferentiated; axial riblets progressively differentiating from the second half of the first whorl onwards, some increasing in prominence and transitioning into the primary ribs of the teleoconch, others becoming close-set and wavy, transitioning into the secondary riblets of the teleoconch. Spiral sculpture of the protoconch composed of approximately equidistant lirae with interspaces 3–5 times their width, persisting on the surface of the teleoconch, forming tiny nodules at intersections with the secondary axial sculpture. Primary axial sculpture of the teleoconch composed of broad, prominent ribs, with interspaces c. 1–2 times their width, overlaid by a secondary axial sculpture of fine, crowded, wavy riblets, with interspaces c. 1–2 times their width. Umbilicus deep, U-shaped. Peristome crescent-shaped; columellar lip slightly reflected. Palatal wall with 4 barriers, all basal in position, extending c. 1/8 whorl, descending gradually anteriorly and posteriorly, regularly spaced, recessed within the aperture; barriers 1 and 4 more deeply recessed and slightly less prominent than the others. Deeply recessed palatal trace commonly present, columellar in position. Parietal wall with 3 barriers, extending c. 3/16 whorl, descending somewhat abruptly anteriorly and posteriorly, regularly spaced, not recessed within the aperture; barrier 1 often slightly less prominent than barrier 2; barrier 2 often slightly less prominent than barrier 3. Other shell features that can be expressed numerically are shown in Table 2 .

Remarks. K. cowiei shares with K. hypsus a U-shaped umbilicus, a similar arrangement of apertural barriers, and a robust shell, heavier than those of species of Mautodontha (Garrettoconcha) Solem, 1976 . It is distinguished from K. hypsus by its smaller shell size and lower peristome. Specimens with a higher spire approach the shell shape of M. (G.) passosi , but are easily distinguished by their larger, heavier shell, and by exhibiting three rather than two parietal barriers.

Etymology. This species is dedicated to Robert Cowie, in recognition of his efforts to understand and preserve the land snail fauna of Pacific islands.

M. (M.) domaneschii sp. nov.

holotype 3.14 5.00 61 1.36 0.78 1.08 0.18 1.24 0.43 0.72 0.39 0.13 paratype 1 3.05 5.07 75 1.46 0.78 1.03 0.24 1.05 0.48 0.76 0.35 0.16 paratype 2 3.18 4.95 74 1.42 0.78 1.07 0.25 1.15 0.45 0.73 0.36 0.18 paratype 3 3.10 5.27 70 1.58 0.81 1.07 0.41 1.02 0.51 0.76 0.33 0.26 paratype 4 3.11 5.23 64 1.37 0.78 1.03 0.23 1.16 0.44 0.76 0.37 0.17 paratype 5 2.84 5.13 67 1.25 0.70 0.92 0.16 0.99 0.44 0.76 0.35 0.13 paratype 6 3.20 5.67 70 1.64 0.80 1.01 0.41 1.13 0.51 0.79 0.35 0.25 paratype 7 3.03 5.21 75 1.45 0.79 1.07 0.29 1.01 0.48 0.74 0.33 0.20 ......continued on the next page Specimen d wn rn h ah aw sp u h/d ah/aw u/d sp/h paratype 8 3.06 4.55 71 1.29 0.84 1.00 0.12 1.03 0.42 0.84 0.34 0.09 mean 3.08 5.12 69.67 1.42 0.79 1.03 0.25 1.09 0.46 0.76 0.35 0.17 standard deviation 0.11 0.30 4.90 0.13 0.04 0.05 0.10 0.08 0.03 0.04 0.02 0.06 M. (G.) aurora sp. nov.

holotype 3.05 5.60 176 1.80 1.02 1.31 0.43 0.55 0.59 0.78 0.18 0.24 paratype 1 2.87 4.99 123 1.58 1.00 1.21 0.30 0.61 0.55 0.83 0.21 0.19 paratype 2 2.94 5.04 136 1.65 1.01 1.24 0.33 0.65 0.56 0.81 0.22 0.20 paratype 3 3.23 5.56 163 1.98 0.99 1.27 0.63 0.66 0.61 0.78 0.20 0.32 paratype 4 3.05 5.52 143 1.87 0.94 1.22 0.53 0.63 0.61 0.77 0.21 0.28 paratype 5 3.01 5.27 144 1.77 0.96 1.20 0.51 0.73 0.59 0.80 0.24 0.29 paratype 6 3.02 5.18 138 1.73 1.03 1.22 0.44 0.62 0.57 0.85 0.20 0.25 paratype 7 2.95 5.13 134 1.72 1.02 1.15 0.37 0.69 0.58 0.89 0.23 0.21 paratype 8 2.96 5.17 145 1.72 0.96 1.23 0.38 0.69 0.58 0.78 0.23 0.22 mean 3.01 5.27 144.67 1.76 0.99 1.23 0.43 0.65 0.58 0.81 0.22 0.25 standard deviation 0.10 0.23 15.91 0.12 0.03 0.04 0.11 0.05 0.02 0.04 0.02 0.04 ......continued on the next page M. (G.) passosi sp. nov.

holotype 3.65 6.78 105 2.45 1.05 1.43 1.01 0.81 0.67 0.73 0.22 0.41 paratype 1 3.75 6.29 118 2.58 0.99 1.46 1.06 0.87 0.69 0.68 0.23 0.41 ......continued on the next page K. hypsus

holotype 6.46 7.75? 4.44 2.49 2.63 1.18 1.33 0.69 0.95 0.21 0.27 specimen 1 5.67 7.09 125 3.83 2.36 2.37 0.84 1.27 0.68 1.00 0.22 0.22 specimen 2 5.64 7.21 135 3.76 2.31 2.17 0.79 1.43 0.67 1.06 0.25 0.21 specimen 3 5.79 7.23 144 3.79 2.19 2.27 0.83 1.38 0.65 0.96 0.24 0.22 specimen 4 6.05 7.29 ~132 3.91 2.53 2.34 0.73 1.60 0.65 1.08 0.26 0.19 ......continued on the next page Specimen d wn rn h ah aw sp u h/d ah/aw u/d sp/h specimen 5 5.25 6.99 130 3.26 2.12 2.07 0.62 1.22 0.62 1.03 0.23 0.19 specimen 6 5.42 6.68 140 3.51 2.21 2.21 0.66 1.21 0.65 1.00 0.22 0.19 specimen 7 5.07 6.77 123 3.27 2.02 2.06 0.73 1.28 0.65 0.98 0.25 0.22 specimen 8 3.06 5.18 109 1.75 1.35 1.24 0.25 0.82 0.57 1.09 0.27 0.14 mean 5.38 6.91 129.75 3.50 2.18 2.15 0.74 1.28 0.65 1.02 0.24 0.21 standard deviation 0.96 0.72 10.95 0.75 0.35 0.38 0.24 0.21 0.03 0.05 0.02 0.03

Figures

FIGURE 20. A – C. Kleokyphus hypsus. A. Holotype (MNHN 25571); B. Specimen 1 (MNHN 25572); C. Specimen 8 (MNHN 25572); D. Kleokyphus cowiei sp. nov., holotype (MNHN 25580). Scale bars: A – B = 4 mm; C – D = 2 mm.

Tables

TABLE 2. Dimensions (in mm) and ratios of dimensions of specimens of Mautodontha (Mautodontha), Mautodontha (Garrettoconcha) and Kleokyphus. See Figure 3 for the placement of measurements. Abbreviations: ah, aperture height; aw, aperture width; d, shell diameter; h, shell height; rn, number of ribs on body whorl; sp, spire protrusion; u, diameter of umbilicus; wn, number of whorls.

Specimen d	wn	rn h	ah	aw	sp	u	h/d	ah/aw u/d	sp/h
M. (M.) daedalea
specimen 1 (Mk13) 3.69	5.55	154 1.49	1.00	1.27	0.29	1.31	0.40	0.79 0.36	0.20
specimen 2 (Mk13) 3.56	5.19	139 1.33	1.00	1.22	0.17	1.34	0.37	0.81 0.38	0.13
specimen 3 (Mk13) 3.66	5.68	152 1.50	0.98	1.21	0.34	1.36	0.41	0.81 0.37	0.23
specimen 4 (Mk13) 3.47	5.42	168 1.23	0.93	1.24	0.11	1.20	0.36	0.75 0.35	0.09
specimen 5 (Mk13) 3.47	5.49	115 1.30	1.05	1.24	0.11	1.23	0.37	0.84 0.35	0.09
specimen 6 (Mk13) 3.21	5.43	136 1.26	0.88	1.13	0.22	1.11	0.39	0.78 0.35	0.17
specimen 7 (Mk13) 3.20	5.35	128 1.26	0.90	1.13	0.19	1.08	0.39	0.79 0.34	0.15
specimen 8 (Mk13) 3.28	5.39	140 1.15	0.91	1.06	0.15	1.30	0.35	0.86 0.40	0.13
specimen 9 (Mk13) 3.10	6.35	165 1.36	0.99	0.99	0.20	1.16	0.44	0.99 0.37	0.14
specimen 10 (Mk13) 3.18	6.36	155 1.40	0.96	1.04	0.27	1.13	0.44	0.92 0.36	0.19
specimen 11 (Mk13) 3.17	6.04	147 1.28	0.99	0.97	0.14	1.24	0.41	1.02 0.39	0.11
specimen 12 (Mk13) 3.01	6.61	139 1.29	0.92	1.04	0.17	1.08	0.43	0.88 0.36	0.13
specimen 13 (Mk13) 3.10	6.12	155 1.27	1.03	1.00	0.19	1.18	0.41	1.03 0.38	0.15
specimen 14 (Mk13) 3.14	6.07	143 1.33	0.99	1.06	0.15	1.20	0.42	0.94 0.38	0.11
specimen 15 (Mk13) 2.91	6.22	156 1.25	0.96	1.01	0.14	1.10	0.43	0.94 0.38	0.11
specimen 16 (Mk13) 2.80	5.79	154 1.19	0.88	0.91	0.15	1.07	0.42	0.97 0.38	0.13
specimen 17 (Mk13) 2.92	5.93	114 1.30	0.89	1.04	0.18	1.06	0.45	0.85 0.36	0.14
specimen 18 (Mk22) 3.35	6.58	~160 1.55	1.15	1.26	0.23	0.97	0.46	0.91 0.29	0.15
specimen 19 (Mk22) 3.27	6.73	152 1.51	1.05	1.19	0.23	1.05	0.46	0.89 0.32	0.15
specimen 20 (Mk22) 3.35	6.30	~176 1.49	1.13	1.24	0.19	1.01	0.44	0.91 0.30	0.13
specimen 21 (Mk22) 3.19	6.52	~144 1.52	1.07	1.20	0.29	0.88	0.48	0.89 0.28	0.19
specimen 22 (Mk22) 2.98	6.15	147 1.27	1.01	1.08	0.13	1.00	0.43	0.94 0.34	0.10
specimen 23 (Mk22) 3.03	6.34	161 1.39	1.01	1.14	0.19	0.87	0.46	0.89 0.29	0.14
specimen 24 (Mk22) 3.11	6.21	~182 1.40	1.00	1.08	0.17	1.08	0.45	0.92 0.35	0.12
specimen 25 (Mk22) 2.94	6.15	133 1.33	0.98	1.10	0.20	0.87	0.45	0.89 0.30	0.15
specimen 26 (Mk22) 2.91	6.15	~148 1.30	0.98	1.08	0.18	0.87	0.45	0.91 0.30	0.14
mean 3.19	6.00	148.58 1.34	0.98	1.11	0.19	1.11	0.42	0.89 0.35	0.14
standard deviation 0.24	0.44	16.10 0.11	0.07	0.10	0.06	0.14	0.03	0.07 0.04	0.03