Corresponding author: Martin Haase (
Academic editor: Robert Hershler
During the course of a project aiming at the reconstruction of the colonization of the South Pacific islands by tateid gastropods based on molecular data we discovered five new species on New Caledonia belonging to the genera
Haase M, Zielske S (2015) Five new cryptic freshwater gastropod species from New Caledonia (Caenogastropoda, Truncatelloidea, Tateidae). ZooKeys 523: 63–87. doi:
New Caledonia is famous for being a biodiversity hotspot harboring a high number of endemic species (
Localities of new species and samples used for morphometric comparisons. Inset shows position of New Caledonia in the Southwest Pacific. Arrows indicate type localities of species represented by more than one sample (see also Table
Holotypes.
Locality data of all samples and GenBank accession numbers of specimens represented in phylogeny (see also Fig.
Species, sample | Locality | Latitutde, longitude |
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16S | IT2 |
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Bouloupari, Ouaméni valley |
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Bouloupari, Ouaméni, prop. Debels |
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Bouloupari, Ouitchambo |
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Moindou, road toward barrage |
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Farino, Sentier de la Cascade et des Sources |
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Sarraméa, track to “Trou d’Eau” |
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Hienghène, Tendo |
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Moindou, road toward barrage |
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Dumbéa, Koé, prop. Oesterlin |
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Sarraméa, road side of RPN 5 |
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Moindou, road side SW Katrikoin |
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Bourail, Oua Oué |
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Poya, road side between Nétéa and Goipin |
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Bouloupari, road side N Nassirah |
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Mont Dore, Rue des Roseaux, prop. Solier |
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Poya, side of road to Goipin |
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Poya, stream into Grotte d‘Adio |
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Houailou, Néoua |
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Moindou, Aremu valley |
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Moindou, road side SW Katrikoin |
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Poya, Grotte d’Adio |
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Bouloupari, road side of RP 4 |
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Sarraméa, prop. Bonnard |
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Bouloupari, Oua Tom |
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Kouaoua, road side N Koh |
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Kouaoua, road side N Koh |
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Poya, Mt. Krapé |
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Poya, Montfaoué |
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Poya, beginning of road into Vallée d’Adio |
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Poya, near Nétéa |
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Hienghène, Tendo |
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Koumac, seepage in N of town |
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Voh, Boyen, overflow of reservoir |
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Hienghène, Tendo |
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Snails were fixed in 70% ethanol in the field, transferred to propylene glycol for shipping by courier, and returned to ethanol, this time 96%, after arrival in our lab. For measurements, up to 20 snails per sample were photographed under a Zeiss SteREO Discovery.V20 dissecting microscope with a Zeiss Axio Cam MR3. Five dimensions – shell height, shell width, aperture height, aperture width, body whorl width – were measured using the program AxioVision 40 V4.8. (Zeiss) and whorls counted to the nearest eighth (
Morphometric analyses of shell measurements including canonical variates analyses (CVA) maximizing the differentiation of groups in multivariate space, multivariate analyses of variance
Phylogenetic analyses were based on a selection of sequences generated by cytochrome oxidase subunit I internal transcribed spacer 2 maximum likelihood
Type and non-type material is deposited at the Museum National d’Histoire Naturelle in Paris Naturhistorisches Museum Wien
Diagnoses and descriptions of
Morphometry. Measurements in mm. Shell measures: , aperture hight , aperture width , width of body whorl , shell height , shell width , number of whorls , coefficient of variation corrected for unequal sample sizes , maximum , minimum , standard deviation
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2.70 | 1.25 | 0.90 | 0.87 | 1.08 | 2.17 | 5.4 | |
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2.40 | 1.10 | 0.80 | 0.75 | 0.97 | 2.00 | 4.50 |
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2.78 | 1.28 | 0.93 | 0.91 | 1.08 | 2.35 | 5.50 |
mean | 2.60 | 1.18 | 0.85 | 0.82 | 1.02 | 2.20 | 5.14 |
median | 2.60 | 1.17 | 0.85 | 0.82 | 1.01 | 2.23 | 5.25 |
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0.11 | 0.05 | 0.04 | 0.04 | 0.03 | 0.11 | 0.28 |
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4.40 | 3.93 | 4.23 | 4.54 | 2.77 | 4.94 | 5.49 |
2.29 | 1.24 | 0.88 | 0.87 | 1.09 | 1.84 | 4.50 | |
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2.10 | 1.16 | 0.83 | 0.83 | 1.04 | 1.71 | 4.13 |
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2.42 | 1.31 | 0.96 | 0.96 | 1.15 | 1.90 | 4.75 |
mean | 2.25 | 1.25 | 0.88 | 0.89 | 1.10 | 1.80 | 4.36 |
median | 2.24 | 1.24 | 0.88 | 0.89 | 1.10 | 1.80 | 4.25 |
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0.09 | 0.04 | 0.04 | 0.03 | 0.03 | 0.05 | 0.18 |
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4.21 | 2.96 | 4.15 | 3.92 | 2.94 | 2.91 | 4.24 |
2.19 | 1.35 | 0.97 | 0.91 | 1.16 | 1.62 | 4.25 | |
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2.03 | 1.29 | 0.87 | 0.86 | 1.10 | 1.53 | 3.75 |
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2.43 | 1.46 | 1.03 | 1.00 | 1.25 | 1.69 | 4.25 |
mean | 2.19 | 1.35 | 0.94 | 0.92 | 1.16 | 1.62 | 4.03 |
median | 2.15 | 1.35 | 0.93 | 0.92 | 1.17 | 1.62 | 4.00 |
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0.11 | 0.05 | 0.04 | 0.04 | 0.04 | 0.04 | 0.15 |
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4.92 | 4.06 | 4.76 | 4.54 | 3.77 | 2.71 | 3.78 |
2.49 | 1.32 | 0.94 | 0.95 | 1.16 | 1.91 | 4.50 | |
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2.23 | 1.26 | 0.89 | 0.88 | 1.07 | 1.77 | 4.25 |
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2.49 | 1.32 | 0.94 | 0.95 | 1.16 | 1.91 | 4.50 |
mean | 2.38 | 1.28 | 0.91 | 0.92 | 1.10 | 1.86 | 4.41 |
median | 2.41 | 1.27 | 0.90 | 0.93 | 1.09 | 1.89 | 4.44 |
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0.11 | 0.03 | 0.02 | 0.03 | 0.04 | 0.06 | 0.12 |
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4.83 | 2.30 | 2.68 | 3.44 | 4.00 | 3.52 | 2.89 |
2.07 | 1.12 | 0.75 | 0.77 | 0.91 | 1.84 | 4.50 | |
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1.85 | 0.97 | 0.65 | 0.70 | 0.82 | 1.76 | 4.25 |
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2.23 | 1.17 | 0.79 | 0.80 | 0.95 | 2.01 | 5.00 |
mean | 2.05 | 1.09 | 0.73 | 0.75 | 0.88 | 1.88 | 4.46 |
median | 2.04 | 1.10 | 0.73 | 0.75 | 0.87 | 1.88 | 4.38 |
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0.12 | 0.05 | 0.03 | 0.03 | 0.03 | 0.07 | 0.19 |
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6.05 | 4.82 | 4.71 | 4.69 | 3.99 | 3.72 | 4.25 |
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2.58 | 1.18 | 0.88 | 0.83 | 1.03 | 2.05 | |
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3.21 | 1.39 | 1.03 | 0.97 | 1.16 | 2.40 | |
mean | 2.79 | 1.27 | 0.94 | 0.91 | 1.09 | 2.19 | |
median | 2.74 | 1.25 | 0.93 | 0.90 | 1.09 | 2.18 | |
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0.17 | 0.06 | 0.04 | 0.04 | 0.05 | 0.09 | |
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6.20 | 4.91 | 4.31 | 4.52 | 4.23 | 4.36 | |
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2.20 | 1.06 | 0.77 | 0.73 | 0.94 | 1.93 | |
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2.48 | 1.25 | 0.87 | 0.91 | 1.04 | 2.28 | |
mean | 2.33 | 1.13 | 0.81 | 0.81 | 0.97 | 2.06 | |
median | 2.32 | 1.12 | 0.80 | 0.80 | 0.96 | 2.03 | |
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0.08 | 0.04 | 0.03 | 0.04 | 0.02 | 0.10 | |
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3.49 | 3.70 | 3.36 | 4.60 | 2.36 | 4.75 | |
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2.35 | 1.16 | 0.83 | 0.83 | 1.04 | 1.96 | |
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2.62 | 1.28 | 0.92 | 0.93 | 1.14 | 2.19 | |
mean | 2.50 | 1.21 | 0.87 | 0.87 | 1.08 | 2.07 | |
median | 2.51 | 1.21 | 0.87 | 0.88 | 1.08 | 2.07 | |
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0.07 | 0.04 | 0.03 | 0.03 | 0.02 | 0.07 | |
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2.90 | 3.20 | 3.16 | 3.25 | 2.32 | 3.43 | |
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2.26 | 1.09 | 0.74 | 0.77 | 0.96 | 2.03 | |
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2.74 | 1.23 | 0.89 | 0.87 | 1.08 | 2.38 | |
mean | 2.49 | 1.17 | 0.84 | 0.83 | 1.03 | 2.12 | |
median | 2.49 | 1.17 | 0.85 | 0.83 | 1.05 | 2.09 | |
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0.14 | 0.05 | 0.04 | 0.03 | 0.04 | 0.11 | |
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5.87 | 4.33 | 5.41 | 3.89 | 3.66 | 5.50 | |
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2.32 | 1.14 | 0.79 | 0.82 | 1.03 | 1.97 | |
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2.64 | 1.23 | 0.91 | 0.91 | 1.12 | 2.14 | |
mean | 2.43 | 1.18 | 0.85 | 0.85 | 1.06 | 2.05 | |
median | 2.42 | 1.19 | 0.86 | 0.85 | 1.06 | 2.04 | |
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0.10 | 0.03 | 0.03 | 0.03 | 0.03 | 0.05 | |
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4.25 | 2.42 | 3.79 | 3.22 | 2.64 | 2.64 | |
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2.28 | 1.16 | 0.78 | 0.82 | 1.04 | 1.86 | |
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2.63 | 1.31 | 0.96 | 0.93 | 1.14 | 2.14 | |
mean | 2.47 | 1.23 | 0.87 | 0.88 | 1.09 | 2.00 | |
median | 2.47 | 1.23 | 0.86 | 0.87 | 1.10 | 2.02 | |
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0.10 | 0.04 | 0.04 | 0.03 | 0.03 | 0.07 | |
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4.18 | 3.08 | 4.18 | 3.28 | 2.62 | 3.62 | |
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2.43 | 1.25 | 0.88 | 0.89 | 1.09 | 1.93 | |
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2.75 | 1.34 | 0.96 | 0.96 | 1.15 | 2.08 | |
mean | 2.62 | 1.30 | 0.92 | 0.92 | 1.12 | 2.01 | |
median | 2.69 | 1.30 | 0.91 | 0.91 | 1.11 | 2.03 | |
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0.12 | 0.04 | 0.03 | 0.03 | 0.02 | 0.06 | |
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4.80 | 2.84 | 3.30 | 3.10 | 2.00 | 3.00 | |
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2.35 | 1.31 | 0.93 | 0.94 | 1.12 | 1.61 | |
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2.74 | 1.62 | 1.10 | 1.06 | 1.34 | 1.80 | |
mean | 2.50 | 1.46 | 1.01 | 1.00 | 1.25 | 1.72 | |
median | 2.43 | 1.46 | 1.01 | 1.00 | 1.27 | 1.70 | |
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0.16 | 0.12 | 0.07 | 0.05 | 0.08 | 0.07 | |
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6.50 | 8.31 | 6.95 | 4.88 | 6.32 | 4.12 | |
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1.87 | 0.98 | 0.68 | 0.68 | 0.86 | 1.84 | |
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2.50 | 1.16 | 0.84 | 0.79 | 0.97 | 2.16 | |
mean | 2.06 | 1.05 | 0.73 | 0.71 | 0.91 | 1.96 | |
median | 2.02 | 1.06 | 0.72 | 0.71 | 0.91 | 1.95 | |
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0.14 | 0.04 | 0.04 | 0.02 | 0.03 | 0.08 | |
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7.04 | 4.02 | 5.11 | 3.51 | 3.21 | 4.23 | |
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2.17 | 1.17 | 0.78 | 0.82 | 1.02 | 1.77 | |
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2.42 | 1.32 | 0.94 | 0.93 | 1.13 | 1.93 | |
mean | 2.31 | 1.26 | 0.88 | 0.88 | 1.07 | 1.84 | |
median | 2.33 | 1.25 | 0.89 | 0.88 | 1.07 | 1.85 | |
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0.07 | 0.04 | 0.04 | 0.03 | 0.03 | 0.05 | |
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3.24 | 3.16 | 4.52 | 3.15 | 2.87 | 2.58 | |
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2.20 | 1.21 | 0.84 | 0.88 | 1.07 | 1.76 | |
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2.46 | 1.31 | 0.94 | 0.98 | 1.20 | 1.97 | |
mean | 2.34 | 1.27 | 0.90 | 0.92 | 1.12 | 1.84 | |
median | 2.35 | 1.28 | 0.91 | 0.92 | 1.12 | 1.83 | |
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0.07 | 0.03 | 0.03 | 0.03 | 0.03 | 0.06 | |
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3.14 | 2.30 | 3.31 | 3.00 | 2.54 | 3.14 | |
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2.17 | 1.20 | 0.85 | 0.85 | 1.06 | 1.76 | |
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2.54 | 1.36 | 1.00 | 0.99 | 1.17 | 1.97 | |
mean | 2.35 | 1.28 | 0.91 | 0.93 | 1.12 | 1.83 | |
median | 2.34 | 1.27 | 0.90 | 0.93 | 1.13 | 1.82 | |
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0.10 | 0.04 | 0.04 | 0.04 | 0.03 | 0.05 | |
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4.27 | 3.12 | 4.50 | 3.93 | 2.91 | 2.62 | |
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2.07 | 1.04 | 0.72 | 0.69 | 0.88 | 1.88 | |
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2.18 | 1.10 | 0.82 | 0.78 | 0.97 | 2.00 | |
mean | 2.11 | 1.08 | 0.76 | 0.74 | 0.92 | 1.96 | |
median | 2.07 | 1.09 | 0.76 | 0.75 | 0.92 | 1.99 | |
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0.07 | 0.03 | 0.05 | 0.05 | 0.05 | 0.07 | |
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3.36 | 3.19 | 7.25 | 6.89 | 5.51 | 3.67 | |
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1.80 | 1.03 | 0.68 | 0.64 | 0.83 | 1.76 | |
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2.30 | 1.16 | 0.81 | 0.77 | 0.99 | 1.99 | |
mean | 2.01 | 1.08 | 0.73 | 0.70 | 0.90 | 1.87 | |
median | 2.02 | 1.05 | 0.72 | 0.68 | 0.89 | 1.86 | |
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0.15 | 0.05 | 0.05 | 0.04 | 0.06 | 0.09 | |
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7.73 | 4.89 | 6.40 | 6.29 | 6.87 | 4.79 |
Holotype
NeCa 12, Bouloupari: Ouaméni-valley, small stream on W-side of road in secondary forest,
The new species is dedicated to the senior author’s daughter on the occasion of her ‘quinceañera’, the 15th birthday.
Conical, 2.2 times higher than wide, 4.5-5.5 whorls, without colour, transparent; protoconch faintly pitted with 1-1.25 whorls; palatal denticle large, elongate, c. 1/3 whorl behind outer lip; with columellar fold in the body whorl; aperture slightly higher than wide (Figs
Shells (all paratypes).
Protoconchs (left) and close-up views of apical microstructure (right).
Elongate-ellipsoidal, paucisprial, nucleus submarginal, orange, one large and one small non-calcareous white peg, well separated from each other (
Operculum.
Epidermis without pigment, eyes black.
Ctenidium with 24–26 (2 males) or 25–28 (3 females) filaments; osphradium kidney-shaped, behind middle of ctenidium.
Radula formula (
Radula.
Ovary without lobes, proximal end c. 1.25 whorls below apex, comprising 0.25–0.5 whorls, eventually reaching stomach; anterior capsule gland yellow-orange, posterior capsule gland opaque-white, albumen gland milky-white; proximal loop of renal oviduct upright comprising 180°, distal loop short; bursa copulatrix pear-shaped, reaching only slightly behind albumen gland; bursal duct long, entering anterior; seminal receptacle on ventral edge of and as long as bursa (
Female genitalia.
Proximal end of lobate testis 1–1.25 whorls below apex, comprising 0.75 whorls, covering proximal end of stomach; vesicula seminalis arising from anterior third of testis; penis fairly delicate with blunt end (
Penis.
This is
Holotype
NeCa 49, Poya: Massif d’Adio, stream flowing into Grotte d’Adio, open secondary forest,
NeCa 43, Poya: small stream on W-side of road between Nétéa and Goipin, on forest edge,
Adioincola is composed of the name of the area of Adio and the Latin noun incola meaning inhabitant, and thus refers to the type locality of the new species.
Pupiform, 1.8 times higher than wide, 4.125-4.75 whorls, without colour, transparent; protoconch faintly pitted with c. 1 whorl; palatal denticle a small droplet 1/8 whorl behind outer lip; aperture as high as wide (Figs
Elongate-ellipsoidal, paucisprial, nucleus submarginal, orange, usually two non-calcareous white pegs, eventually accompanied by a small third one (
Epidermis without pigment, eyes black.
Ctenidium with 18-19 (3 males) or 21–24 (2 females) filaments; osphradium kidney-shaped, behind middle of ctenidium.
Radula formula (
Ovary without lobes, proximal end 1.25 whorls below apex, comprising 0.25-0.5 whorls, eventually reaching stomach; anterior capsule gland yellow-orange, posterior capsule gland opaque-white, albumen gland milky-white; proximal loop of renal oviduct bent forward, distal loop short; bursa copulatrix almost cubical, reaching behind albumen gland; bursal duct long, entering anterior; no seminal receptacle (
Proximal end of lobate testis 1.25–1.5 whorls below apex, comprising 0.5-0.75 whorls, covering proximal end of stomach; vesicula seminalis arising from anterior half of testis; penis slender, terminal end occasionally forming short filament (
This is
Holotype
NeCa 33, Moindou: spring-fed stream close to road in Aremu valley, under shrub,
The new species is named after the Aremu valley, where it has been discovered.
Broadly pupiform, 1.62 times higher than wide, 3.75-4.25 whorls, without colour, transparent; protoconch faintly pitted with 0.75-0.9 whorls; palatal denticle a small droplet 1/8 whorl behind outer lip; aperture practically as high as wide (Figs
Elongate-ellipsoidal, paucisprial, nucleus submarginal, orange, two non-calcareous white pegs, eventually accompanied by a small third one (
Epidermis without pigment, eyes black.
Ctenidium with 15-16 (2 males) or 19-20 (2 females) filaments; osphradium elongate, slightly behind middle of ctenidium.
Radula formula (
Ovary without lobes, proximal end 1.25-1.75 whorls below apex, comprising 0.25-0.5 whorls, reaching stomach; capsule gland with long and slender, opaque-white vestibulum, anterior capsule gland yellow-orange, toward posterior capsule gland covered with brown spots, posterior capsule gland opaque-white with a central milky section, albumen gland milky-white; proximal loop of renal oviduct bent forward, distal loop long; bursa copulatrix higher than long, reaching behind albumen gland; bursal duct long, entering anterior; no seminal receptacle (
Proximal end of lobate testis 1 whorl below apex, comprising c. 0.75 whorls, covering proximal end of stomach; vesicula seminalis arising from distal third of testis; penis very long and slender (
This is
Holotype
NeCa 30, Moindou: spring along road SW of Katrikoin, under shrub,
The Latin adjective clandestinus means clandestine and refers to the new species’ external identity with
Pupiform, 1.86 times higher than wide, 4.25-5 whorls, without colour, transparent; protoconch very faintly pitted with c. 1 whorl; palatal denticle a small droplet 1/8 whorl behind outer lip; aperture as high as wide (Figs
Epidermis without pigment, eyes black.
This is
Holotype
NeCa 44, Poya: stream at side of small road branching off road between Nétéa and Goipin toward the Vallée d’Adio, under shrub close to overgrown garden,
The new species is named after the village of Nétéa, which is closely proximal to our collecting locality.
Elongate-pupiform, 1.88 times higher than wide, 4.25–5 whorls, without colour, transparent; protoconch faintly pitted with c. 1 whorl; palatal denticle an elongate droplet c. 1/4 whorl behind outer lip; aperture slightly wider than high (Figs
Elongate-ellipsoidal, paucisprial, nucleus submarginal, orange, one non-calcareous white peg (
Epidermis without pigment, eyes black.
Ctenidium with 15 (1 male) or 19-22 (5 females) filaments; osphradium short-elongate, behind middle of ctenidium.
Radula formula (
Ovary without lobes, proximal end 1.25-1.5 whorls below apex, comprising 0.25-0.5 whorls, not reaching stomach; anterior capsule gland yellow-orange, posterior capsule gland opaque-white, albumen gland milky-white; proximal loop of renal oviduct bent forward, distal loop short; bursa copulatrix globular, reaching slightly behind albumen gland; bursal duct long, entering anterior; no seminal receptacle (Fig.
Proximal end of lobate testis 1 whorl below apex, comprising slightly more than 0.5 whorls, covering proximal end of stomach; vesicula seminalis arising approximately in middle of testis; penis very long and slender (
This is
The CVA plot (Fig.
CVA plot for
Pairwise morphometric comparisons of , not significant
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The CVA (Fig.
CVA plot for
Pairwise morphometric comparisons of , not significant
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In the phylogenetic analysis (Fig.
Maximum likelihood phylogram showing bootstrap support when > 50%. Outgroup pruned from tree; new species highlighted by bold face type.
Average pairwise uncorrected (p) distances between selected species based on the
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9.7 | 8.1 | 8.5 | 7.6 |
Our phylogenetic analyses based on DNA sequence data confirmed the suspicion of
The new species provide an additional truncatelloid example stressing the importance of an integrative taxonomic approach combining morphological, anatomical and genetic methods (e.g.
Genetic differentiation was an important indicator of species status. Pairwise p-distances > 7.4% are far above any threshold suggested by advocates of barcoding (e.g.,
While conducting our morphometric analyses we appreciated that the measuring methods applied for the material described previously (
Another methodological problem almost expectedly occurred in the field. All collections made for our previous monograph (
Four of the five new species were found in single sites and the fifth was found at only two sites. Considering the vulnerability of small habitats like springs and the rapid anthropogenic development and changes on New Caledonia just outlined immediately raises concern regarding the chances of long-term survival of these species (see also
This publication would not have been possible without the assistance of numerous local field guides and landowners who provided access to their properties. We thank Christel Meibauer for assistance in the molecular lab. Rabea Schlüter is acknowledged for support at the SEM. We are grateful to the authorities of New Caledonia for issuing the relevant permits. Christine Pöllabauer has been a generous host in Noumea during our field trip. The manuscript benefited from suggestions of two anonymous reviewers and Robert Hershler. Financial support was received from the German Research Foundation (HA 4752/2-1).