A new species of Casmaria H. Adams & A. Adams, 1853 (Gastropoda, Cassidae) from the Philippines identified by molecular data Author Fedosov, Alexander Author Olivera, Baldomero M. 6378C0BB-05EF-4772-9CB9-0DD10BD9233E Department of Biology, University of Utah, Salt Lake City, UT 84112, USA & urn: lsid: zoobank. org: author: 6378 C 0 BB- 05 EF- 4772 - 9 CB 9 - 0 DD 10 BD 9233 E Author Watkins, Maren D8C297B4-2A7F-4E60-9295-A22E2EE60581 Department of Biology, University of Utah, Salt Lake City, UT 84112, USA & urn: lsid: zoobank. org: author: D 8 C 297 B 4 - 2 A 7 F- 4 E 60 - 9295 - A 22 E 2 EE 60581 Author Barkalova, Varvara 96884FF5-5586-47B3-AE82-B0BC9E74DD76 Moscow State University, Faculty of Biology, 1 - 12, Leninskie Gory, Moscow, 119991 Russia & urn: lsid: zoobank. org: author: 96884 FF 5 - 5586 - 47 B 3 - AE 82 - B 0 BC 9 E 74 DD 76 text European Journal of Taxonomy 2014 2014-03-28 78 1 13 journal article 22016 10.5852/ejt.2014.78 a9fa297c-2ef8-4599-ac6e-4ede538b0500 2118-9773 3835042 3C2942F0-9C88-4382-B28C-0F02D7200EA5 Casmaria boblehmani sp. nov. urn:lsid:zoobank.org:act: FA0142B4-BEF4-4F08-8BFD-C0870F39DC88 Fig. 2 Casmaria ponderosa – Robin 2008: 145 , fig. 5 — Buijse et al. 2013: 13 , fig. 25-2 [middle specimen], pl. 6, figs 6, 8, 13, pl. 7, figs 1, 5, 8-10, 12-15, 17, pl. 8, fig. 1,?pl. 10, fig. 11. Etymology This species is named to honor I. Robert (Bob) Lehman, a great scientist, mentor and teacher. One of the authors (BMO) has been the grateful direct beneficiary of his remarkable generosity. Type material Holotype Molecular voucher Casm.5, 36.3 mm , Danajon Banks , off Olango Island , Cebu , Philippines , 10°17.8’ N , 124°05.3’ E , 10-25 m , coll. local diver, Jun. 2012 , ANSP-453749 ( Fig. 2A-C ). Paratypes 1 ( 35.1 mm ) and 2 ( 34.8 mm , molecular voucher Casm. 7), same locality as holotype , ANSP-453750 ( Fig. 2D-E ). The holotype and paratype 2 provided the two DNA sequences labeled Casm. 7 and Casm. 5, respectively, in Fig. 1 . Fig. 2. Casmaria boblehmani sp. nov. A-C . Holotype, 36.3 x 19.5 mm, Danajon Banks, Olango Is, Philippines. D-E . Paratype 2, 35.1 x 21.7 mm, same locality as holotype. F . 27.7 x 18.8 mm, Tip of Cape York, Queensland, Australia. G . 38.6 x 25.4 mm, off Woka Island, Molucca Sea, Indonesia. H-I . ANSP 276463, 41.5 x 24.2 mm, Akuilan Is, Fiji. K-L . ANSP 266122, 36.5 x 24.6 mm, Atiue, Tahiti, Society Islands. Other material examined PHILIPPINES : 34.6 mm , off Masbate Island, Central Philippines, coll. local diver, BO collection; 34.9 mm , off Calituban Island, Bohol , 9 m , on mud, BO collection; 44.9 mm , off Cuyo Island, Palawan , 10-25 m , BO collection; 32.2 mm , Palawan Island, unknown locality, 10-15 m , BO collection. AUSTRALIA : 27.7 x 18.8 mm , tip of Cape Yorke, Queensland , BO collection ( Fig. 2F ). INDONESIA : 38.6 x 25.4 mm , off Woka Island, Molucca Sea, BO collection ( Fig. 2G ). VANUATU : juv., Expedition “ SANTO 2006”, Stn. FR10, Belmoul Lagoon, 15°36.7’ S , 167°05.8’ E , 5-25 m , MNHN IM- 2007-33625; juv., Expedition “ SANTO 2006”, Stn. FR03, Belmoul Lagoon entrance, 15°36.2’ S , 167°06.3’ E , 3-32 m , MNHN IM- 2007-33628 ( Fig. 1A ). FIJI : 41.5 x 24.2 mm , off Akuilan Island, ANSP 276463 ( Fig. 2H-I ). SOCIETY ISLANDS: 36.5 x 24.6 mm , Atiue, Tahiti, ANSP 266122 ( Fig. 2K-L ). Description Adult shell small to medium-sized, solid, glossy, ovate-elongate with very prominent terminal varix, studied specimens 34.8-36.3 mm high. Proportions of shell vary considerably, ratio of shell breadth to height 0.58-0.7. Protoconch of about 4 smooth, evenly convex whorls. Teleoconch of 4-5 whorls; early spire whorls evenly convex, with no sculpture. Last adult whorl and in most specimens penultimate whorl with row of glossy nodules (6 to 12 on last whorl), weakly developed in some specimens ( Fig. 2D-E ), prominent in others ( Fig. 2 G ). Due to nodules, late whorls distinctly shouldered; shoulder situated at mid-height of penultimate whorl and adapical quarter of last adult whorl. Aperture ovate, outer lip strongly thickened, drawn upwards, with wide, shallow sinus in upper side. Interior of outer lip with distinct dentition. Apertural varix very prominent, thick, with ten brownish-black blotches on abapertural face; adapertural face with seven or eight well-developed spines directed transversally to apertural plane. Their positions correspond approximately with black blotches on abapertural face. Columellar shield solid, white, heavily callused, with numerous spiral wrinkles in some specimens. Last adult whorl of pinkish, tan, or pale brown background color, with indistinct brown spiral stripes and a row of brown to blackish-brown spots below suture. Siphonal canal with black blotch. Interior of aperture pale brown. Tip of protoconch brown. Fig. 3. Distributions of Casmaria turgida (short-dashed line and triangles), C . unicolor (dashed line and filled circles) and C . boblehmani sp. nov. (solid line and squares) in the Indo-Pacific. Table 2. Summary of morphological differences among Casmaria species. Abbreviations: OL – outer lip; TV – terminal varix; n.f. – nodulose form; r – rarely. Number / direction of spines on OL: f – forward; tr – transverse to apertural plane. Position of spines on OL: a 1/3 – anterior third. Number of blotches on TV: N+n – N major blotches + n strokes; N se – number of subequal blotches .

Species	Smooth form	Nodulose form	Number / direction of spines on OL	Position of spines on OL	Number of blotches on TV	Dentition within	Row of blotches at suture	Row of blotches on shell base	Color pattern	Individual characters
C. erinaceus	x	x	3-5 / f	a 1/3	7-14 se	-	-	-	no, zigzags, spiral rows of dots
C. kayae	x	x	3-5 / f	a 1/3	9-12 se	-	-	-	zigzags
C. ponderosa	x	x	7-9 / tr	a 1/2 - 3/4	5 + 1-2	+	+	+	no
C. turgida	x	-	8-11 / tr	a 1/2 - 3/4	5 +1-2 or 10-11 se	-	+	-	zigzags	shell length usually exceeds 60 mm
C. boblehmani	?	x	8-9 / tr	a 1/2 - 3/4	8-10 se	+	+	r	no, spiral bands
C. unicolor	x	x	10-12 / tr	a 2/3 - 4/5	11-12 se	-	+	+ (n.f.)	no, zigzags, spiral rows of dots	inflated upper whorl profile giving the shell a wider, more stepped spire outline
C. cernica	x	-	6-7 / tr	a 1/2	5 + 1-2	-	+	+	spiral rows of blotches	narrow and narrowly varicate shell
C. atlantica	x	-	6-7 / tr	a 1/2	5 + 1-2	-	+	-	spiral bands	narrow and narrowly varicate shell
C. perryi	x	-	6-7 / tr	a 1/2	5 + 1-2	-	+	-	spiral bands	narrow and narrowly varicate shell
C. beui	x	-	>9	a 2/3	10-11 se	r	+	+	spiral rows of blotches, zigzags	shell very small, wide and thin
C. kalosmodix	x	-	?-	-	5, 10-11 se	-	+	r	axial lines + spiral bands	narrow axial lines over entire shell surface

Remarks Specimens that are most appropriately assigned to Casmaria boblehmani sp. nov. are found at sites in eastern Indonesia , northern Australia ( Northern Territory and Queensland ), Vanuatu and the South Pacific. The largest specimen ( 51.8 mm ) that we can confidently assign to C. boblehmani sp. nov. is known from the Philippines ( Buijse et al. 2013 : pl. 6, fig. 13). In the Philippines C. boblehmani sp. nov. occurs syntopically with C. ponderosa , C. erinaceus and C. turgida . All four species co-occur in sandy patches at depths of 1 to 30-40 meters. Characters allowing the unequivocal recognition of Casmaria boblehmani sp. nov. and the delimitation of the Casmaria species described to date are summarized in Table 2 . Casmaria ponderosa and C. boblehmani sp. nov. can be distinguished from each other by the number and arrangement of brown blotches on the dorsal side (abapertural face) of the terminal varix: C. ponderosa has five major blotches and two to three intermediate strokes, while C. boblehmani sp. nov. has nine to ten blotches of equal or subequal size, evenly distributed along the varix of most specimens. Another character, although less stable, is a row of spots on the shell base, posterior to the siphonal canal. These are always distinct in C. ponderosa , but are missing in most specimens, or in rare cases present but indistinct, in C. boblehmani sp. nov. Specimens of Casmaria erinaceus can easily be distinguished from both C. ponderosa and C. boblehmani sp. nov. by the absence of dentition on the inner surface of the outer lip and by the presence of only three or four (very rarely five) spines restricted to the anterior edge of the outer lip, directed forward. The anterior edge of the outer lip is drawn slightly forward and abaxially. Both C. ponderosa and C. boblehmani sp. nov. have distinct dentition on the inner surface of the outer lip and have six or more spines directed transversely to the apertural plane, located along the anterior and medial edges of the outer lip. Most of these characters were mentioned by Abbott (1968) and Buijse et al. (2013) as allowing discrimination between C. erinaceus and C. ponderosa . Casmaria boblehmani sp. nov. can be readily distinguished from Casmaria turgida (Reeve, 1848) . The latter species is rather conservative in morphology throughout its range ( Buijse et al. 2013 ) and has a shell with evenly convex whorls, notable because of the characteristic zigzag color pattern. Besides that, C. turgida exceeds all co-occurring Casmaria forms in size. Unfortunately, C. turgida is missing in our molecular dataset. However, the conservative morphology of this species and its sympatry on a small scale with C. erinaceus , C. ponderosa and C. boblehmani sp. nov. at a number of Indo-Pacific localities prove that it is a distinct species. Despite the fact that C. ponderosa , C. boblehmani and C. turgida in most cases can be distinguished from each other confidently by the combination of characters discussed above, these characters may intergrade to a certain extent, creating somewhat intermediate forms that may be difficult to interpret. Buijse et al. (2013 : pl. 6, figs 6, 8, pl. 9, fig. 15) depicted a few specimens of intermediate appearance that we are not certain where to assign. All these specimens have smooth shells intermediate between C. ponderosa and C. turgida in size ( 48-51 mm ). Among the other Casmaria species not occurring sympatrically with C. boblehmani sp. nov. , C. unicolor (Pallary, 1926) is probably the most similar. As in other Casmaria species, C . unicolor varies significantly in sculpture and encompasses both smooth and nodulose forms, and the nodulose form closely resembles C. boblehmani sp. nov. (see Buijse et al. 2013 : pl. 22). These two forms seem to differ in the number of spines on the outer lip. C. boblehmani sp. nov. possesses seven to eight spines, while C . unicolor has 10-12, and they are located along the entire length of the lip in many specimens, reaching the adapical angle of the aperture. Apart from that, C. unicolor can be distinguished by its more inflated upper (adapical) whorl profile and more clearly distinguished, weakly flattened sutural ramp, giving it a wider, more stepped spire outline. Casmaria unicolor is restricted to the Red Sea ( Fig. 3 ), and the widely distant distributions of this species and C. boblehmani sp. nov. suggest that these two forms are sufficiently divergent to be considered as separate species. The three species Casmaria cernica (G. B. Sowerby III, 1888), C. perryi (Iredale, 1912) and C. atlantica Clench, 1944 are extremely similar to each other. They are characterized by being narrower in outline and with a narrower terminal varix than C. boblehmani sp. nov. Besides that, all three mentioned species share the same arrangement of dark blotches on the dorsal side of the apertural varix as in C. ponderosa : five major blotches with one or two intermediate strokes between each major pair. In comparison with Casmaria boblehmani sp. nov. , C. kalosmodix (Melvill, 1883) has a much larger, far more elongate and more narrowly varicate shell with narrow axial color lines that are not present in other Casmaria species, and with no spines on the outer lip. As in C. boblehmani sp. nov. , the blotches on the abapertural face of the varix in C. kalosmodix are narrow, but in the latter species they are grouped in 3-4 clusters, separated by wide interspaces. Another species, Casmaria kayae Buijse, Dekker & Verbinnen, 2013 , in general resembles C. erinaceus . C. boblehmani sp. nov. can easily be distinguished from C. kayae by the number and position of spines on the outer lip and by the color pattern. C. kayae has three or four (five in a few specimens) spines on the anterior portion of the lip, and these are directed forward. Many specimens have a distinctive zigzag color pattern, quite different from the usual coloration of C. boblehmani sp. nov. Finally, C. beui Buijse, Dekker & Verbinnen, 2013 , from Western Indian Ocean, can easily be distinguished from C. boblehmani sp. nov. by its small size, its smooth, thin, relatively very wide shell with distinctively inflated whorls, and by its characteristic color pattern, resembling that of C. turgida . Distribution Casmaria boblehmani sp. nov. is widely distributed across the Pacific Ocean: from the Philippines , eastern Indonesia , northern Australia , to Fiji and the Society Islands, and (based on specimens illustrated by Buijse et al. 2013 ) to Hawaii ( Fig. 3 ). Possibly reaches India (based on the specimen illustrated by Robin 2008 ).