taxonID	type	description	language	source
03A69848FF9E1E56FF30FC9CFE00FC38.taxon	materials_examined	TYPE SPECIES. — Terebellum convolutum Lamarck, 1802 (= Bulla volutata Solander in Brander, 1766) by original designation. Lutetian, Paris Basin.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF9E1E54FEFBFBD2FBA5FD7A.taxon	materials_examined	TYPE LOCALITY. — Hordwell (England), Bartonian (Middle Eocene). TYPE MATERIAL. — Bulla volutata Solander in Brander, 1766: the single preserved syntype (BMNH GG 21010) has been selected unambiguously by Jung (1974: 18) as lectotype: “ the type specimen of Bulla volutata Solander in Brander (1766, p. 34, pl. 6, fig. 75) ”. According to article 74.5 of the ICZN (1999) this mention, made before 2000, constitutes a valid lectotype designation.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF9E1E54FEFBFBD2FBA5FD7A.taxon	description	DESCRIPTION Shells Shell up to 75 mm in height and 22 mm in diameter, slightly inflated with short and blunt to somewhat acute apex (Jung 1974). Aperture long and narrow. Callus of the inner lip very thin and rarely preserved except on some “ gerontic ” shells. Columella almost straight near the base, or rarely bent in some big shells. Outer lip very thin, not thickened, sinuous for the whole height and extending to the apex (Fig. 6 B). Siphonal notch moderately deep on the dorsal side of the shell. No sculpture on the surface of the shell. Colour pattern According to Jung (1974), perfectly preserved specimens from the Lutetian of Paris Basin show, in normal light, many small and irregular dots which compose the residual colour pattern. Under UV light, the colour pattern consists of numerous small dots on a lighter background (Fig. 7). These dots, variable in size, show two levels of residual pigmentation (Fig. 7 G). The first one consists of triangle-like spots, darker than the background. The apex of these spots is directed towards the growing edge. The second one consists of spots with a fuzzy border, white, fluorescent under UV light and directed towards the inner lip. They are adjacent to the base of the dark spots. Light spots are variable in shape and size, but they are usually narrower than the dark spots (Fig. 7 A-D, G-I). These two components form the dots. Dot density is generally more important on the apical part of the shell. They often form sinuous, axial rows nearly parallel to the edge of growth (Fig. 7 A, B). The space between these rows is narrow on the apex and broader on the median and basal parts of the shell. VARIABILITY OF THE COLOUR PATTERN The background is generally light, but sometimes shows large and diffuse darker areas (Fig. 7 H, I). The density of dots is very variable. Some shells have a few large dots and their axial rows are not clearly defined. On other shells, the density of dots is so great that their coalescence may hide the original background. Some specimens strongly coalescent show very few isolated dots. Juvenile specimens often show proportionally bigger dots (Fig. 7 E, F). Sometimes the relative size of the light spot is greater than the dark triangular spot (Fig. 7 E, F). STRATIGRAPHIC EXTENSION Several specimens (Fig. 8) collected in Sapicourt (Marne, France) constitute the new oldest occurence of the species (Ypresian [Cuisian]).	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF9C1E52FD25FD0FFB90FB5E.taxon	materials_examined	TYPE LOCALITY. — Hordwell (England), Bartonian (Middle Eocene). TYPE MATERIAL. — Lectotype (BMNH GG 21010) designated by Jung (1974). OTHER MATERIAL EXAMINED. — See Appendix 1.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF9C1E52FD25FD0FFB90FB5E.taxon	description	DESCRIPTION Shell Shell up to 40 mm in height and 14 mm in diameter, inflated with short and blunt apex (Fig. 6 F, G). Aperture long and narrow. Callus of the inner lip thin and rarely preserved, except on the largest shell. Columella mainly straight but bent back, near the base. Outer lip very thin, not thickened and sinuous for the whole height in labral view (Fig. 6 G). Siphonal notch moderately deep on the dorsal side of the shell. No sculpture on the surface of the shell. Colour pattern The colour pattern consists of numerous small and irregular dots on a lighter background (Fig. 9). These dots show two levels of residual pigmentation (Fig. 9 C, D, H, J, K): triangular spots, darker than the background, with the apex orientated towards the outer lip and diffuse, fluorescent spots lighter than the background and orientated towards the inner lip. These white fluorescent spots, adjacent to the base of the triangles, are variable in shape. They can be narrower than the dark triangles (Fig. 9 C, D, J, K) or bigger and ovoid to almost circular (Fig. 9 H). These two components form the dots. The dots are of various sizes. Th ey are more distorted and axially elongated on the shell extremities (Fig. 9 A, C). When they coalesce, these dots form sinuous axial nearly collabral rows (Fig. 9 A, E, F).	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF9A1E4EFD0CFB30FE40FEA6.taxon	materials_examined	TYPE LOCALITY. — Chaussy (Val d’Oise, France), Lutetian (Middle Eocene). TYPE MATERIAL. — The illustrated syntype of Cossmann (1889) has been selected unambiguously by Jung (1974: 21) as lectotype, using wrongly the term holotype: “ Cossmann’s figure […]. Th is specimen is the holotype ”. According to article 74.5 of the ICZN (1999), this mention, made before 2000, constitutes a valid lectotype designation. Unfortunately, the type material of this species has not been found in the Bourdot collection, deposited at University of Lyon 1 (UCBL). A neotype designation is needed to clarify the taxonomic status of this species. A neotype from the type locality (MNHN A 28885, leg. Pacaud, Fig. 10 F-H) is accordingly chosen here. Th is initiative is in conformity with the rules specified by the ICZN (1999: art. 75), as regards the designation of a neotype. OTHER MATERIAL EXAMINED. — See Appendix 1.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF9A1E4EFD0CFB30FE40FEA6.taxon	description	DESCRIPTION Shell Shell up to 28 mm in height and 7 mm in diameter, moderately slender with elongated and pointed apex (Fig. 6 H). Aperture long and narrow. Callus of the inner lip very thin, inconspicuous and rarely preserved. Straight columella near the base. Outer lip not thickened, reaching the apex and sinuous for the whole height in labral view, i. e. outer lip prosocyrt on the anterior part and opisthocyrt on the posterior part of the shell (Fig. 6 J). Siphonal notch moderately deep on the dorsal side of the shell. No sculpture on the surface of the shell. Colour pattern The shells show small and irregular dots on a lighter background. Th ese dots are composed of two levels of residual pigmentation (Fig. 10 E, H). The first one consists of triangular spots, darker than the background. These small triangles are pointed towards the growing edge. The second one consists of white fluorescent fuzzy-bordered spots directed towards the inner lip. They are adjacent to the base of the dark triangles. White fluorescent spots are generally narrower than the dark spots (Fig. 10 E, H). Th ese two components form the dots. Th ese dots, variable in size and sometimes coalescent (Fig. 10 H), form sinuous axial nearly collabral rows (Fig. 10 C, D, F, G). Th e space between these rows is narrow on the apex and broader on the median and basal parts of the shell. VARIABILITY OF THE COLOUR PATTERN The background is generally uniform and can bear large and diffuse darker areas (Fig. 10 F-H). The density of the dots varies. When the density is high the axial rows are clearly defined, whereas they are less clearly or not defined when it decreases. COMPARISONS OF THE THREE SPECIES OF SERAPHS: S. VOLUTATUS, S. SOPITUS AND S. OLIVACEUS Shell Among the representatives of Seraphs, the adult shell shape of S. volutatus, S. sopitus and S. olivaceus is very similar (Fig. 6). Moreover, these shells share a lack of sculpture. Seraphs volutatus generally has a straight columella, while it is bent in the two other species. Jung (1974) regarded Bulla volutata as a synonym of Bulla sopitus. However, according to Le Renard (1992), S. sopitus differs from S. volutatus by its short and blunt apex. For Jung (1974), S. olivaceus is distinguished from S. volutatus (= S. volutatus + S. sopitus sensu Jung [1974]) by an elongated and acute apical part (Fig. 6 H). According to Jung (1974), S. volutatus shows a gradual change in the shell shape during ontogeny, and we observed some very large specimens similar to S. olivaceus and some small specimens similar to S. sopitus. Residual colour pattern Dots on a light background are common to S. volutatus (Fig. 8), S. sopitus (Fig. 9) and S. olivaceus (Fig. 10). These species show the same type of dots with two levels of residual pigmentation: 1) dark triangular spots orientated towards the growing edge; and 2) white fluorescent spots towards the inner lip. Furthermore, they have sinuous rows, almost parallel to the line of growth. In the studied material (22 ex.) of S. olivaceus, the dots are always small and less numerous than in the two other species.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF9A1E4EFD0CFB30FE40FEA6.taxon	discussion	DISCUSSION After study of 15 specimens of S. sopitus, 22 of S. olivaceus, 562 of S. volutatus, the colour patterns of each species appear variable, but very similar and no distinction can be made. According to Jung (1974) and Le Renard (1992), the main differences result from shell characters (the morphology of the apex and the columella). However, these differential characters, which vary during ontogeny and even in adulthood, were not defined considering variability. In conclusion, our personal opinion is that, following the colour patterns, the discrimination of these three species is not justified. In order to know if it is necessary to continue to distinguish them, it would be rewarding to study the shell using modern morphometric methods.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF861E4EFED0FE48FBAEFEC6.taxon	materials_examined	MATERIAL EXAMINED. — Lutetian, Chaussy (Les Garennes), Val d’Oise, 1 ex. (MNHN A 25011, Faullummel coll.).	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF861E4EFED0FE48FBAEFEC6.taxon	description	DESCRIPTION Shell Shell 13 mm in height and 4 mm in diameter, slightly inflated with short and blunt apex. Aperture long and narrow. Callus of the inner lip not preserved. Straight columella near the base. Outer lip very thin, not thickened and extending to the apex (Fig. 11 A, B). Siphonal notch moderately deep on the dorsal side of the shell. No sculpture on the surface of the shell. Colour pattern The residual pattern is only composed of two levels of residual pigmentation: fluorescent axial stripes on a darker background. Th e stripes, sinuous overall, include some zigzags on short segments (Fig. 12 A- C). They are rarely continuous on the whole height of the shell. Th eir width is variable. COMPARISONS The shell shape is indistinguishable from that of young S. volutatus and they share the lack of sculpture. However, the residual pattern of Seraphs sp. 1 is very different from that of all other Seraphs in showing axial sinuous stripes. Moreover, it is the only pattern showing light elements on a darker background.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF861E4EFED0FE48FBAEFEC6.taxon	discussion	DISCUSSION Although the residual colour pattern of Seraphs sp. 1 is clearly distinct (no intermediate pattern) from that of S. volutatus and even from that of the other Seraphs, we believe that it is preferable to avoid the description of a new species, because of the strong intrageneric variability and the occurence of only one specimen having this pattern.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF861E4EFC8FFEABFB1DF970.taxon	materials_examined	MATERIAL EXAMINED. — Lutetian, Fresville (Coteau de Vauville), Manche, 2 ex. (MNHN A 28577, de Morgan coll.).	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF861E4EFC8FFEABFB1DF970.taxon	description	DESCRIPTION Shell Shell 18 mm in height and 6 mm in diameter, slightly inflated with short and blunt apex. Aperture long and narrow. Callus of the inner lip not preserved. Straight columella near the base. Outer lip extending to the apex but broken in labral view (Fig. 11 D). Siphonal notch not observable (Fig. 10 E). No sculpture on the surface of the shell. Colour pattern On the best preserved specimen, the residual pattern is only composed of two levels of residual pigmentation: dark spiral stripes on a lighter background. The stripes are broad and have marked borders (Fig. 12 D-F). Th ey are continuous and each stripe has the same width on the whole of the last whorl. The width of the different stripes is variable. On the less well-preserved specimen, a dark spiral stripe is present with dots, like in S. volutatus. COMPARISONS The shell shape of Seraphs sp. 2 is similar to that of young S. volutatus (Fig. 6 A-C) and both species lack sculpture. Under UV light, the best preserved specimen is very different from the other Seraphs in showing thick and dark spiral stripes. The less well-preserved specimen is slightly similar, but bears dots as in S. volutatus.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF861E4EFC8FFEABFB1DF970.taxon	discussion	DISCUSSION If the best preserved specimen bears this peculiar pattern, the less well-preserved specimen suggests continuity between this pattern and that of S. volutatus, because of the occurrence of dots. Th us, as for Seraphs sp. 1, we prefer to avoid describing a new species, based on a small number of specimens.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF821E4AFF32FF2AFBFEFDE4.taxon	materials_examined	Type locality. — Chaussy, Les Garennes (Val d’Oise, France), Lutetian (Middle Eocene). TYPE MATERIAL. — Holotype (Deshayes coll.); 1 paratype (Deshayes coll., Les Garennes, Chaussy). Unfortunately, the type material of this species has not been found in the Deshayes collection, deposited at the UCBL. A neotype designation is needed to clarify the taxonomic status of this species. A neotype from the type locality (MNHN A 28781, Ledon leg.; Figs 11 F, H, I; 13 A-D) is thus chosen here. Th is initiative conforms to the rules specified by the ICZN (1999: art. 75), as regards the designation of a neotype. OTHER MATERIAL EXAMINED. — See Appendix 1.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF821E4AFF32FF2AFBFEFDE4.taxon	description	DESCRIPTION Shell Shell up to 36 mm in height and 8 mm in diameter, very slender and elongated, with straight and almost parallel sides (Fig. 11 F, H). Aperture long and narrow. Callus of the inner lip very thin, but well delineated, continuing adapically on the spire and forming a narrow band parallel to the outer lip. Columella slightly bent backwards near the base. Outer lip thickened, almost straight in labral view, opisthocline and extending to the apex (Fig. 11 I). Shallow siphonal notch on the dorsal side of the shell (Fig. 11 H). Few oblique faint grooves near the base (Fig. 11 G). Colour pattern The residual pattern is only composed of 2 levels of residual pigmentation: dark dots on a lighter background (Fig. 13). Th e dark dots are small and spirally elongated forming narrow triangles (Fig. 13 B-C, I). These triangles, orientated towards the outer lip, are irregularly spread over the whole surface. COMPARISONS Seraphs leukoleptus is the only species having a shell with straight and almost parallel sides. Moreover, it is easily distinguished from the other species by a shallow siphonal notch. Th e pattern of S. leukoleptus is distinguishable from the other patterns by its dots. Th ey do not form rows, as observed in S. volutatus and S. olivaceus. Moreover, these dots are composed of only one level of residual pigmentation, while the other species have dots composed of two levels of residual pigmentation.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF821E48FD36FD8EFE0BF970.taxon	materials_examined	TYPE LOCALITY. — Gaas (Landes, France), Rupelian (Oligocene). TYPE MATERIAL. — Seraphs subconvolutus: the illustrated syntype (UBT 65 - 2 - 120, Grateloup. coll.) of Grateloup (1834) has been selected unambiguously by Jung (1974: 26) as lectotype, using wrongly the term of holotype: “ They therefore represent the holotypes of S. subconvolutus ... ”. But in the original work of d’Orbigny (1852), no holotype was designated for the taxon. In accordance with article 74.6 of the ICZN (1999), Jung’s stipulation constitutes a lectotype fixation.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF821E48FD36FD8EFE0BF970.taxon	description	DESCRIPTION Shell Shell up to 31 mm in height and 8 mm in diameter moderately slender with acute apical part. Aperture long and narrow. Callus of the inner lip thin, but well delineated, extending towards the apex and forming a narrow band parallel to the outer lip. Columella slightly bent backwards near the base of the shell. Outer lip slightly thickened and somewhat prosocyrt, in labral view, below the adapical end of the aperture (Jung 1974). Outer lip extending onto the posterior part of the shell, but bent towards the dorsal side (Fig. 14 B, C, E) and “ reaching the apex after about half a volution ” (Jung 1974). Siphonal notch moderately deep. Surface sculpture of narrow oblique grooves observable on the ventral side near the base (Fig. 14 F). Colour pattern No residual pattern is observed under UV light. Nevertheless, the studied specimens seem slightly corroded and we cannot preclude the possibility that the species originally had a colour pattern. COMPARISONS OF THE SHELL MORPHOLOGY Seraphs subconvolutus is similar to S. chilophorus and S. peterjungi n. sp. (Fig. 15) in sharing an abapical part of the columella slightly bent backwards, narrow oblique grooves on the ventral side near the base and a well delineated callus (Fig. 14 D) of the inner lip forming adapically a narrow band parallel to the outer lip. However, this species is the only Seraphs having an opisthocline extension of the outer lip on the posterior part of the shell. The outer lip does not continue directly to the apex, but is bent towards the dorsal side reaching the apex after about half a volution (Fig. 14 B, C, E). This feature of S. subconvolutus resembles that of members of Paraseraphs (Jung 1974), but the shell is involute as in other Seraphs.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF801E46FD3AFF2AFBB0F970.taxon	materials_examined	TYPE LOCALITY. — Fontenay-en-Vexin, Bois du But (Eure, France), Lutetian (Middle Eocene). TYPE MATERIAL. — Th e illustrated syntype (Bourdot coll., sample Bernay) of Cossmann (1889) has been selected unambiguously by Jung (1974: 22) as lectotype, using wrongly the term holotype: “ Cossmann’s original figure […]. This specimen is the holotype ”. In accordance with article 74.5 of the ICZN (1999) this mention, made before 2000, constitutes a valid lectotype designation. Unfortunately, the type material of this species has not been found in the Bourdot collection, now deposited at UCBL. A neotype designation is needed to clarify the taxinomic status of this species. A neotype from the locality of Fontenay-en-Vexin (MNHN A 25009, Fig. 16 C-E) is accordingly chosen here. This initiative conforms to the rules specified by the ICZN (1999: art. 75), as regards the designation of a neotype. OTHER MATERIAL EXAMINED. — See Appendix 1.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF801E46FD3AFF2AFBB0F970.taxon	description	DESCRIPTION Shell Shell up to 28 mm in height and 8 mm in diameter, moderately slender with acute apical part. Aperture long and narrow. Callus of the inner lip well developed and well delineated, continuing adapically on the spire and forming a narrow band parallel to the outer lip (Fig. 15 C). Columella bent backwards near the base. Outer lip, slightly thickened and sinuous in labral view, near the adapical end of the aperture. Outer lip extending to the apex and continuing on the opposite side of the apex for a short distance (Fig. 15 G, I). Jung called this apical part of the outer lip, the apical canal. Siphonal notch moderately deep on the dorsal side. Surface sculpture of narrow oblique grooves near the base (Fig. 15 H). Colour pattern The colour pattern is composed of two different dark elements on a lighter background: thick rows of patches nearly perpendicular to the line of growth and tiny dots located between these rows. The rows are composed of dark triangular patches (7 - 8). Th e apex of each of is orientated towards the outer lip. Th e patches are usually disconnected and separated by a small white fluorescent patch. This morphology is called a fishbone pattern (Fig. 16 I, J) by Meinhardt (1998: fig. 4.10). The distance between the rows and their width are variable. Sometimes, several rows are adjacent to one another (Fig. 16 F). Th e dots are small and more or less triangular (Fig. 16 E). As for the patches, the apex is always directed towards the outer lip. Th eir size, distribution and shape are heterogenous and irregular. Above the most adapical row, the density and the size of dots are often greater as is the degree of coalescence (Fig. 16 H). COMPARISONS OF THE SHELL MORPHOLOGY The morphologies of S. chilophorus and S. peterjungi n. sp (Fig. 15 J-N) are very similar with their thickened callus of the inner lip and their narrow band on the adapical part of the shell. Moreover, they share an apical canal (Fig. 15 G, I, L), lacking in the other Seraphs. The shell of S. chilophorus is more slender than those of S. volutatus (Fig. 6 A-E) and S. sopitus (Fig. 6 F, G). Th is species is generally more slen- der than S. olivaceus, although both can have an acute apex. Also, S. chilophorus clearly differs from S. olivaceus, S. volutatus or S. sopitus by the morphology of its outer lip and its callus of the inner lip. VARIABILITY OF THE COLOUR PATTERN Among 163 specimens from various localities (11 from the Lutetian and 1 from the Bartonian), 160 (98.16 %) show a residual colour pattern under UV light (Table 1). Accordingly, these observations provide a good general survey of the intraspecific variability. The residual pattern of S. chilophorus is quite variable (Fig. 17) and is often subject to significant distortion of the elements on the shell extremities, especially the apical one (Fig. 17 C-F). The huge variability is visible on every element of the residual pattern. Th e number of spiral rows varies from 0 to 6. Several shells show partially or totally connected rows (Fig. 17 B, D). Th e shape of the dark patches can show important variations from triangular (Fig. 17 A, C) to completely distorted or shapeless (Fig. 17 F, J). Although they are usually separated by a small fluorescent patch, they may be connected with each other (Fig. 17 E). The density of dots fluctuates from low values (Fig. 17 A, D-F) to very high ones (Fig. 17 K). In this case, the dots form axial rows more or less coalescent (Fig. 17 K- N). Th is coalescence sometimes increases until the dots disappear and only axial segments are observable (Fig. 17 H, M, N). Th ere are often some large triangular and coalescent patches near the edge of the outer lip. Finally, there are sometimes small axial segments between the dark patches of the rows (Fig. 17 J). In spite of this variability, we can distinguish three morphological types. Th e most abundant an representative type 1 is composed of spiral rows of dark triangles directed towards the outer lip and small dots between them (Fig. 17 A-C, I). Th e type 2 shows some thin and tightened axial segments with, often, remnants of spiral rows of triangular patches (Fig. 17 H, N, O). Th e type 3 shows only large and more or less coalescent dark triangles (Fig. 17 D, E). Several shells bear a morphology intermediate (Fig. 18) between types 1 and 2 (Fig. 17 J-M), between types 1 and 3 (Fig. 17 F), between types 2 and 3 (Fig. 17 G). Th e relative abundance of these morphologies is given in Figure 18. Figure 17 illustrates this variability with a certain disparity, but the observation of numerous intermediate specimens suggest that it is impossible to separate one type from another, as different species. COMPARISONS OF THE COLOUR PATTERN Seraphs chilophorus (Fig. 16) bears dots, as S. volutatus, S. sopitus, S. olivaceus and S. leukoleptus (Figs 7 - 10). Th e pattern of S. chilophorus is, however, easily distinguishable as the dots are composed of only one residual colour and by the occurrence of broad spiral rows of dark patches.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF8B1E42FF1EFF2AFE02FB9E.taxon	materials_examined	TYPE LOCALITY. — Chaussy, Les Garennes (Val d’Oise), France, Lutetian (Middle Eocene). TYPE MATERIAL. — Holotype: MNHN A 28578, leg. Pacaud. Paratypes: Chaussy (Les Garennes), Val d’Oise, 2 ex., MNHN A 27613 (1965 - 11 coll.); 1 ex., MNHN A 28506 (Morlet coll.); 1 ex., MNHN A 28931 (leg. Pacaud); 1 ex., MNHN A 27273 (leg. Pacaud); 1 ex. MNHN A 28537 (Faullummel coll.); 1 ex., MNHN A 28538 (Faullummel coll.); 1 ex., MNHN A 27550 (Schtrock coll.); 1 ex., MNHN A 28550 (leg. Pacaud); 1 ex., MNHN A 28559 (Ledon coll.); Beynes, Yvelines, 1 ex., MNHN A 28789; Fercourt, Oise, 5 ex., MNHN A 27634 (leg. Pacaud); Grignon, Yvelines, 1 ex., MNHN A 27632 (ex Galerie de Zoologie coll.); 1 ex., MNHN A 28512 (leg. Caze); 1 ex., MNHN A 28530 (Ballot coll.); Parnes, Oise, 1 ex., MNHN A 11186 (d’Orbigny coll.); 1 ex., MNHN A 28511 (Roissy coll.); Parnes (Grande Cronière), Oise, 1 ex., MNHN A 28521 (leg. Pacaud); Saulx-Marchaix, Yvelines, 1 ex., MNHN A 28520 (1965 - 11 coll.).	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF8B1E42FF1EFF2AFE02FB9E.taxon	etymology	ETYMOLOGY. — Dedicated to Peter Jung for his work on the Seraphsidae.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF8B1E42FF1EFF2AFE02FB9E.taxon	description	DESCRIPTION OF THE HOLOTYPE Shell 23 mm in height and 7 mm in diameter, slender with acute apical part. Aperture long and narrow. Callus of the inner lip well developed and delineated, continuing adapically on the spire and forming a narrow band parallel to the outer lip. Columella bent backwards near the base (Fig. 15 K). Outer lip slightly thickened and, in labral view, sinuous near the adapical end of the aperture. Outer lip extending to the apex and continuing on the opposite side for a short distance (Fig. 15 K, L, N). Siphonal notch moderately deep on the dorsal side of the shell. Surface sculpture of narrow oblique grooves near the base (Fig. 15 M). COLOUR PATTERN DESCRIPTION The colour pattern shows two different and superimposed levels of residual pigmentation: a dark one and a white and fluorescent one. The background displays an intermediate colour between these two levels. Th e darker elements correspond to several thick spiral rows of patches. Th e lighter fluorescent elements, overlaid on the darker, consist of narrow spiral lines. Th e morphology of the dark elements is similar to the pattern of S. chilophorus: several spiral rows of dark triangles of which the apex is directed towards the outer lip. Th eir width, depending on the size of the patches, is variable (Fig. 19 A, B). In addition, some tiny and isolated triangles, orientated towards the outer lip, are visible between the rows (Fig. 19 G, H). Th eir size, distribution and shape are heterogenous and irregular (Fig. 19 G, H). On the apical part of the shell, the elements are more dense and coalescent, tending to be shapeless. VARIABILITY The variability is only observable on the darker elements of the residual pattern. Th e rows can be more or less distant, from widely spaced to contiguous (Fig. 19 A, E, F). Dark patches can show considerable shape variation from triangular to completely distorted (Fig. 19 E, F). Th ey can be well separated or connected. The fluorescent elements, narrow spiral lines, do not show perceptible variation except in their number: between 16 and 20. SHELL AND COLOUR PATTERN COMPARISONS The shell of Seraphs peterjungi n. sp. is indistinguishable from that of Seraphs chilophorus. Accordingly, for shell comparison with the other species, see S. chilophorus. The dark underlying elements of S. peterjungi n. sp. resemble the colour pattern of S. chilophorus, but it appears to be less variable. Seraphs chilophorus never has fluorescent narrow lines.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF8B1E42FF1EFF2AFE02FB9E.taxon	discussion	DISCUSSION The comparisons between S. peterjungi n. sp. and S. chilophorus are respectively based on 24 specimens and 163 specimens (Table 1). Using the shell shape, no distinction can be made between the two populations. Using the residual pattern, we can observe a strong similarity in the dark component, suggesting a close relationship. However, S. peterjungi n. sp. shows an additional element with some light and fluorescent spiral lines (Fig. 19), lacking in all examined S. chilophorus (Fig. 16) and suggesting a morphological discontinuity. Considering the number of specimens studied and the lack of intermediaries between the two colour patterns, we cannot regard both populations as a single species. In this case, the residual pattern does seem to be a unique discriminating criterion enabling the differentiation of fossil species.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF8A1E42FF28FB73FBACFDBB.taxon	materials_examined	TYPE SPECIES. — Ovula media Deshayes & Milne-Edwards, 1844 (= Ovula fragilis Defrance, 1825) by monotypy. Lutetian, Paris Basin. GENERIC DISCUSSION Jung (1974) considered Diameza and Miniseraphs Jung, 1974, as two subgenera of Seraphs. Nevertheless, S. (Diameza) fragilis (Defrance, 1825), S. (Miniseraphs) eratoides (Cossmann, 1889) and S. (Miniseraphs) isabella (Bernay in Deshayes, 1865) have very small and inflated shells with a large apical angle (Fig. 20) which are obviously different from those of Seraphs (s. s.) (Figs 6; 11; 15) which are larger, slender and more elongated. Th us, these species are, here, no longer considered to belong to the genus Seraphs and are placed together within Diameza, which is here raised to the generic level. The name Diameza, older than the name Miniseraphs, is selected for this genus including two subgenera: Diameza and Miniseraphs. Under UV light, these three species show very similar patterns (Figs 21 - 23). They bear sinuous or wavy axial lines which are lacking in Seraphs (Figs 7 - 10; 12; 13; 16; 18), in Paraseraphs (Figs 26; 28; 30) and Terebellum (Figs 1; 2). These observations strengthen the initial interpretation based on the shell morphology: Diameza (s. s.) fragilis, D. (Miniseraphs) eratoides and D. (Miniseraphs) isabella are closely related and very different from all other fossil and Recent species.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF8A1E40FD3CFD0FFF6CF970.taxon	materials_examined	TYPE LOCALITY. — Villiers-Saint-Frédéric (Yvelines, France), Lutetian (Middle Eocene). TYPE MATERIAL. — Ovula fragilis: syntype (s) from Grignon (Yvelines, France), not found. Ovula media: lectotype designated by Jung (1974) from the Deshayes collection and now deposited at the UCBL, not found; paralectotypes (UCBL, Deshayes coll.), not found. A neotype designation is needed to clarify the taxonomic status of this species. A neotype from the locality Villiers-Saint-Frédéric (MNHN A 28939, leg. Pacaud, Fig. 21 A-F), near Grignon (Yvelines) is accordingly chosen here. Th is initiative conforms to the rules specified by the ICZN (1999: art. 75), as regards the designation of a neotype. OTHER MATERIAL EXAMINED. — See Appendix 1.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF8A1E40FD3CFD0FFF6CF970.taxon	description	DESCRIPTION Shell Inflated shell up to 11 mm in height and 4 mm in diameter. Aperture long and narrow. Callus of inner lip clearly delineated, but weakly developed. Columella slightly bent backwards near the base. Outer lip thickened and extended above the apex. Outer lip forming an acute structure (Fig. 20 B) hiding the actual apex (Jung 1974). Outer lip sinuous, prosocyrt on the abapical part and opisthocyrt on the adapical part of the shell (Fig. 20 C). Siphonal notch moderately deep. Surface sculpture of narrow oblique grooves near the base (Fig. 20 D). Colour pattern The residual pattern is composed of two levels of residual pigmentation: dark lines on a lighter background. Th ese lines are axial and from wavy to zigzagging (Fig. 21 A, G, H, L-N). They usually cover the outer lip and the extremities of the shell. On the median part, the shells show a gradual transition from wavy lines to a chessboard pattern (Meinhardt 1998) (Fig. 21 A, B, D, E). VARIABILITY The lines can be more or less wavy. Some specimens have only axial straight lines on the whole shell surface (Fig. 21 C, J, K), while other specimens have a chessboard pattern on the median part of the shell.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF881E40FD1CFF25FB27FE97.taxon	materials_examined	TYPE SPECIES. — Terebellum (Seraphs) eratoides Cossmann, 1889 by original designation. Lutetian, Paris Basin.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF881E7EFD1AFE8BFEACFCD9.taxon	materials_examined	TYPE LOCALITY. — Chaussy, Les Garennes (Val d’Oise, France), Lutetian (Middle Eocene). TYPE MATERIAL. — Lectotype designated by Jung (1974) from the Bernay collection; paralectotype (Boutillier coll.) from Vaudancourt (Oise), not found. The type material of this species has not been found in the Bernay collection, now deposited at the UCBL. A neotype designation is needed to clarify the taxonomic status of this species. A neotype from the type locality (MNHN A 28932, Faullummel coll., Fig. 22 A-D) is accordingly chosen here. This initiative conforms to the rules specified by the ICZN (1999: art. 75), as regards the designation of a neotype. OTHER MATERIAL EXAMINED. — See Appendix 1.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FF881E7EFD1AFE8BFEACFCD9.taxon	description	SHELL DESCRIPTION Shell up to 8 mm in height and up to 5 mm in diameter, very stout with wide apical angle (Fig. 20 F-H). Aperture long and narrow. Callus of the inner lip clearly delineated, but weakly developed. Columella slightly bent backwards near the base. Outer lip thickened and forming a small protuberance at the apex (Fig. 20 J). Outer lip slightly sinuous for the whole height in labral view (Fig. 20 G). Siphonal notch moderately deep on the dorsal side. Faint oblique to spiral grooves near the base (Fig. 20 I). COLOUR PATTERN DESCRIPTION The residual pattern is only composed of 2 levels of residual pigmentation: dark lines on a lighter background. Th ese axial lines are wavy, but do not cover the whole shell surface. Most specimens show a combination of two morphologies: 1) dark wavy lines, sometimes forming zigzags (Fig. 22 E), which are visible on the extremities of the shell and near the outer lip; 2) on the median part of the shell, a chessboard morphology (Fig. 22 A, C, E, G, I, J), as defined by Meinhardt (1998: fig. 5.6). Th e chessboard pattern can appear as “ light dots ” on a darker background on poorly preserved shells (Fig. 22 G, J). VARIABILITY The intraspecific variability is weak. Some specimens show numerous continuous wavy lines near the outer lip (Fig. 22 E, F), while others show a chessboard pattern on the whole shell surface, except for the abapical and adapical extremities (Fig. 22 G, H).	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FFB61E7AFF36FCACFDA2FCB8.taxon	materials_examined	TYPE LOCALITY. — Chaussy, Les Garennes (Val d’Oise, France), Lutetian (Middle Eocene). TYPE MATERIAL. — Lectotype designated by Jung (1974) from the Bernay collection, and now deposited at UCBL, not found; paralectotype (UCBL, Deshayes coll.), not found. Accordingly, a neotype designation is needed to clarify the taxonomic status of this species. A neotype from the type locality (MNHN A 28935, leg. Pacaud, Fig. 23 A-D) is chosen here. Th is initiative conforms to the rules specified by the ICZN (1999: art. 75), as regards the designation of a neotype. OTHER MATERIAL EXAMINED. — See Appendix 1.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FFB61E7AFF36FCACFDA2FCB8.taxon	description	DESCRIPTION Shell Small shell up to 16 mm in height and up to 8 mm in diameter, inflated with wide apical angle. Aperture long and narrow. Callus of the inner lip clearly delineated, but weakly developed. Columella slightly bent backwards near the base. Outer lip thickened and extending to the apex (Fig. 20 K). Outer lip prosocyrt on the abapical part and opisthocyrt on the adapical part in labral view (Fig. 20 N). Siphonal notch moderately deep. Oblique grooves near the base of the shell. Colour pattern The colour pattern of the species is composed of 2 levels of pigmentation: dark axial wavy lines on a lighter background. Th e axial lines form some zigzags (Fig. 23). The zigzags cover mainly the median part of the shell. On the base and the apex, the lines remain straight or slightly sinuous, showing very weak curves (Fig. 23 I-M, O, P). Th e width of the lines is almost homogenous. Th e space between them is equivalent to their width. VARIABILITY The specimens bear zigzags of greater (Fig. 23 D) or lesser amplitude (Fig. 23 I). Some specimens bear a gradual transition from wavy lines to a zigzag pattern (Fig. 23 H, I), while others show an abrupt transition, probably due to a growth interruption (Fig. 23 F, G). Wavy axial lines crossing and forming a localized grid, called meshwork (Fig. 23 M, N) as defined by Meinhardt (1998: fig. 5.3 c, d), have been observed too. COMPARISONS OF THE THREE SPECIES OF DIAMEZA Shell	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FFB61E7AFF36FCACFDA2FCB8.taxon	description	All species share a pattern composed of two levels of pigmentation. Th e colour pattern of Diameza (s. s.) fragilis (Fig. 21) is closely related to that of D. (Miniseraphs) eratoides (Fig. 22). It shares with D. (M.) eratoides a combination of two morphologies: 1) dark wavy lines visible on the extremities of the shell and near the outer lip; and 2) a chessboard pattern on the median part of the shell. Nevertheless, some shells of Diameza (s. s.) fragilis bear a residual pattern only composed of axial lines (Fig. 21 J-K), lacking in D. (Miniseraphs) eratoides. Diameza (M.) eratoides differs from D. (M.) isabella by a chessboard pattern covering a large part of the shell. Moreover, the amplitude of the wave of each line is lower than that observed on Diameza (Miniseraphs) isabella (Fig. 23).	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FFB21E7AFF3CFC4DFDA0FBC8.taxon	materials_examined	TYPE SPECIES. — Paraseraphs tetanus Jung, 1974 by original designation. Ypresian (Cuisian), Paris Basin.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FFB21E78FF34FB93FE80FCB8.taxon	materials_examined	TYPE LOCALITY. — Cuise-la-Motte (Oise, France), Ypresian (Cuisian, Lower Eocene). TYPE MATERIAL. — Holotype (NMB H 15409); 3 paratypes (NMB H 15408, H 15405, H 16502). OTHER MATERIAL EXAMINED. — See Appendix 1.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FFB21E78FF34FB93FE80FCB8.taxon	description	DESCRIPTION Shell Shell up to 38 mm in height and 8 mm in diameter, evolute with straight and almost parallel sides (Fig. 24 A, C, D, F). Suture distinct on the adapical part (Fig. 24 A, C, D, F). Aperture long and narrow. Callus of the inner lip well developed and well delineated. Columella bent backwards near the base of the shell (Fig. 24 B, E). Outer lip slightly opisthocline and thickened on the adapical part. Outer lip not reaching directly to the apex, but bent towards the dorsal side of the shell (Fig. 24 B, C, E, F). Callus of the inner lip, thin, extending on the spire towards the apex and forming a narrow band parallel to the outer lip. Siphonal notch moderately deep. No sculpture observed. Colour pattern The pattern is composed of 2 levels of colouration: dark elements on a lighter background (Fig. 25). The darker elements are usually very thin and form axial rows of dots (Fig. 25 C, F, I) and segments (Fig. 25 K). On the subsutural part of the last whorl and on the anal canal, a peculiar pattern can be distinguished. It corresponds to an oblique succession of darker and lighter “ patches ” (Fig. 25 H, J, L, P, R). These “ patches ” are not true patches as defined in the part terminology, but are the result of the coalescence of axial segments. VARIABILITY Among 319 specimens from 5 localities (Cuisian), 304 (95.3 %) show a residual pattern under UV light (Table 1). Th us, these observations provide a good general survey of intraspecific variability of the pattern of this species. The residual pattern of Paraseraphs tetanus shows strong variability (Fig. 26). Th e dots are variable in size and shape from axially elongated to circular or triangular shape or indeed completely distorted (Fig. 26 I, J). Th eir density on the shell, and the degree of coalescence, can be weak to very considerable. Some shells bear only a few dark dots disseminated on the surface (Fig. 26 F, G, K). When the density of dots increases, they form some thin axial rows (Fig. 26 D, E, H) or, by coalescence, compressed segments (Fig. 26 A-C) or sinuous stripes (Fig. 26 O- R). In spite of this variability, two representative morphological types can be distinguished (types 1 and 2, Fig. 27). Type 1 corresponds to the most abundant portion of the analysed specimens. The pattern of the type 1 is composed of axial rows of more or less coalescent dark dots (Fig. 26 A-F). Type 2 bears some sinuous and more or less discontinuous axial stripes (Fig. 26 O-R). Several shells show a morphology intermediate between both types (Figs 26 G-K, L-N; 27). Th e relative abundance of these morphologies is given in Figure 27. The Figure 26 illustrates each type described above with several intermediate specimens, showing the impossibility of separating one type from another, as different species.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FFB01E78FF29FC4DFCFFFA12.taxon	materials_examined	TYPE LOCALITY. — Les Garennes, Chaussy (Val d’Oise, France), Lutetian (Middle Eocene). TYPE MATERIAL. — Holotype (NMB H 14491); paratype (NMB H 10539), Les Garennes, Chaussy (Val d’Oise, France); paratypes (UCBL coll. Deshayes), Mouchy-le- Châtel (Oise) and Parnes (Oise), not found. OTHER MATERIAL EXAMINED. — See Appendix 1.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FFB01E78FF29FC4DFCFFFA12.taxon	description	DESCRIPTION Shell Shell up to 30 mm in height and 7 mm in diameter, moderately slender and evolute with distinct suture. Aperture long and narrow. Callus of the inner lip well developed, well delineated, extending to the apex and forming a narrow band parallel to the outer lip (Fig. 24 I). Basal part of the columella bent backwards. Outer lip, almost straight and slightly opisthocline (Fig. 24 H, M), not thickened on the basal part of the shell. Adapical part of the outer lip thickened and extending from the adapical end of the aperture, towards the apex (Fig. 24 H), or bent towards the dorsal side (Fig. 24 M). Siphonal notch moderately deep. Sculpture with a few oblique grooves near the base of the shell (Fig. 24 K). Colour pattern Paraseraphs placitus has numerous and irregular dots with 2 levels of residual coloration on a lighter background (Fig. 28). The first level consists of triangular spots, darker than the background. Th ese spots, more or less distorted, are orientated towards the growing edge (Fig. 28 C, F, L). The second level consists of fluorescent spots with fuzzy borders, lighter than the background. These light spots, adjacent to the base of dark spots, are narrow and directed towards the inner lip in ventral view (Fig. 28 C, F, L). They are variable in size. Th ese two components form the dots. The density of dots is generally low. Also, a peculiar pattern can be distinguished on the adapical part of the outer lip: an oblique succession of darker and lighter “ patches ” (Fig. 28 B, D, E, M, N). These “ patches ” are probably the result of the coalescence of the dark axial segments (Fig. 28 D). VARIABILITY Sometimes the darker spots of the dots are larger near the edge of the outer lip (Fig. 28 B, C, P) and are coalescent.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FFB01E74FD38F9F7FE2BFAFC.taxon	materials_examined	TYPE LOCALITY. — Saffré, Bois-Gouët (Loire-Atlantique, France), Bartonian (Middle Eocene). TYPE MATERIAL. — Th e type material of Vasseur (1882) was lost (Cossmann 1898). Accordingly, Cossmann (1898: pl. 8, figs 10, 15) designated a neotype from the Dumas collection deposited in MNHN, but this specimen has not been found either. Later, Jung (1974: 36) wrongly considered the neotype of Cossmann to be the holotype of this species. OTHER MATERIAL EXAMINED. — See Appendix 1.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FFB01E74FD38F9F7FE2BFAFC.taxon	description	SHELL DESCRIPTION Shell 73 mm in height and 13 mm in diameter, slender and elongated. Shell evolute with distinct suture. Aperture long and narrow. Callus of the inner lip well developed and clearly delineated (Fig. 29 A, B). Abapical part of the columella bent backwards. Outer lip, almost straight (Fig. 29 C), not thickened except near the adapical end of the aperture. Outer lip running towards the apex and slightly bent on the dorsal side of the shell (following Jung [1974], our material being broken). Siphonal notch moderately deep on the dorsal side. No sculpture on the surface of the shell. COLOUR PATTERN DESCRIPTION Paraseraphs armoricus bears numerous and variablesized dots on a lighter background (Fig. 30). These dots, generally circular, spread over the whole surface of the shell. Th ey show 2 levels of residual pigmentation (Fig. 30 B, C, E): 1) round dark spots orientated toward the inner lip; and 2) white fluorescent, crescent-like spots orientated towards the growing edge. Th ese 2 components form the dots. The density of dots is low, with little coalescence (Fig. 30 B). COMPARISONS OF THE THREE SPECIES OF PARASERAPHS: P. TETANUS, P. PLACITUS AND P. ARMORICUS Shell These species of Paraseraphs are easily distinguishable from the species of Seraphs by their evolute spire (Figs 24; 29). Moreover, the shells are usually more slender. According to our observations, several differential characters attributed by Jung (1974) to distinguish Paraseraphs tetanus and P. placitus seem to be erroneous. Jung (1974) believed that P. tetanus is more slender than P. placitus. In fact, the shape of P. tetanus is variable and some specimens can be as inflated as P. placitus (Fig. 24 F, L). Furthermore, Jung (1974) suggested that, on the posterior part of the shell, the outer lip of P. tetanus is bent towards the dorsal side, while that of P. placitus runs directly to the apex. For P. tetanus, his observations seem correct, but in P. placitus, the outer lip can continue directly to the apex or can be bent towards the dorsal side (Fig. 24 H, M). Accordingly, this character is not consistent to distinguish the two species. On the other hand, contrary to Jung assumptions, the shell of P. tetanus does not show oblique grooves near the base. The Bartonian species P. armoricus is very similar to the other species, but much bigger (Fig. 29 A-D). Residual colour pattern The dots constituting the pattern of P. placitus (Fig. 28) appear very similar to those of S. volutatus, S. sopitus and S. olivaceus (Figs 7 - 10). However, the dots are clearly ordered in these Seraphs, while no axial rows of dots have been observed in P. placitus. Regarding P. armoricus (Fig. 30), the orientation of the two components of the dots is inversed in comparison to P. placitus and even Seraphs. The darker spots are orientated towards the outer lip in P. armoricus and towards the inner lip in P. placitus. Furthermore, the shape of the dots is obviously different between P. placitus and P. armoricus. It is generally triangular for P. placitus (Fig. 28 C, F, L), while it is circular for P. armoricus (Fig. 30 B, E). The pattern of P. tetanus is quite variable, but differs sharply by the occurrence of axial rows and by lacking of white fluorescent spots (Fig. 25). Using shell characters, it is difficult to separate these three species of Paraseraphs, while using residual colour pattern we can obviously observe differences justifying their distinction.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FFBC1E74FF38FA96FB2CFB3D.taxon	materials_examined	TYPE LOCALITY. — Abbecourt (Oise), France, Thanetian (Upper Paleocene), biozone NP 9. TYPE MATERIAL. — Holotype (MNHN A 05707, leg. Pacaud)	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FFBC1E74FF38FA96FB2CFB3D.taxon	etymology	ETYMOLOGY. — As the precursor of the Eocene Paraseraphs.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FFBC1E74FF38FA96FB2CFB3D.taxon	description	DESCRIPTION OF THE HOLOTYPE (FIG. 29 E, F) Shell 19 mm in height and 7 mm in diameter, inflated and evolute with distinct suture. Spire short and blunt. Aperture long and narrow. Callus of the inner lip not preserved. Basal part of the columella and outer lip not preserved. No sculpture on the surface of the shell. No residual pattern observed under UV light (shell too poorly preserved). COMPARISONS OF THE SHELLS This species is easily distinguishable from the members of Seraphs by its evolute spire. Within Paraseraphs, although the shell of Paraseraphs praecedens n. sp. is incompletely preserved, it seems clearly stouter than the other species (Fig. 29 D-F). Also, it is the smallest species of the genus.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
03A69848FFBC1E74FF38FA96FB2CFB3D.taxon	discussion	DISCUSSION From the Thanetian, Paraseraphs praecedens n. sp. is the oldest record of Paraseraphs for the world and the oldest record for the Seraphsidae in Europe. Previously, the oldest European occurrence of the Seraphsidae was Paraseraphs tetanus from the Ypresian (Cuisian, biozone NP 12) of the Paris Basin. Another species in this family is recorded in the? Late Paleocene-Early Eocene, Seraphs minus (Vincent, 1913) from Angola (Landana beds, Cabinda, West Africa). Jung (1974) regarded it as a nomen dubium, because the two syntypes “ represent the immature stage of some species of Seraphs ”. However, although this species is only based on young specimens, it should be considered as one of the oldest records of the Seraphsidae, along with P. praecedens n. sp.	en	Caze, Bruno, Merle, Didier, Pacaud, Jean-Michel, Saint Martin, Jean-Paul (2010): First Systematic Study using the Variability of the Residual Colour Patterns: The Case of the Paleogene Seraphsidae (Mollusca, Gastropoda, Stromboidea). Geodiversitas 32 (3): 417-477, DOI: 10.5252/g2010n3a4, URL: http://www.bioone.org/doi/abs/10.5252/g2010n3a4
