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Colour pattern preservation in Fuersichella n. gen. (Gastropoda: Neritopsoidea), bivalves, and echinid spines from ....

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Heft 37 2007 Miscellanea Wirceburgensis - Franz T. Fürsich zum 60. Geburtstag

Colour pattern preservation in Fuersichella n. gen. (Gastropoda: Neritopsoidea), bivalves, and echinid spines from the Upper Jurassic of Portugal SIMON SCHNEIDER & WINFRIED WERNER 143-160

Würzburg 2007

Colour pattern preservation in Fuersichella n. gen. (Gastropoda: Neritopsoidea), bivalves, and echinid spines from the Upper Jurassic of Portugal

SIMON SCHNEIDER & WINFRIED WERNER

SCHNEIDER, S. & WERNER, W. 2007. Colour pattern preservation in Fuersichella n. gen. (Gastropoda: Neritopsoidea), bivalves, and echinid spines from the Upper Jurassic of Portugal. – Beringeria 37: 143-160, 10 text-figs.; Würzburg.

Abstract. Colour preservation has been observed in shells and spines of five different invertebrate taxa from the Upper Jurassic of the Lusitanian Basin, Portugal. In this paper the colour patterns of these animals are detailed; moreover, the neritimorph gastropod „Neritina“ bicornis is placed in the new genus Fuersichella. Coloured bands in the shell of the bivalve Isognomon rugosus are documented for the first time. Newly discovered material provides additional information on the morphology and colouration of Coelastarte discus. In addition, the first detailed characterization of the colour patterns preserved in fossil echinoid spines assignable to Pseudocidaris lusitanica and P. spinosa is given. Finally, chemical composition, formation, and possible functions of the colour patterns in these animals are discussed.

Colour patterns, Bivalvia, Gastropoda, Echinoidea, Late Jurassic, Portugal

Addresses of the authors: SIMON SCHNEIDER, WINFRIED W ERNER, Bayerische Staatssammlung für Paläontologie und Geologie, Richard-Wagner-Str. 10, D–80333 München, Germany; , .

Introduction

Our most sophisticated sense organs are the eyes. Through (e.g. KOBLUK & MAPES 1989, TICHY 1980). The second the eyes, we are visually attracted to two fundamental largest group of coloured shell-bearing fossils are the parameters of the objects: shape and colour. While shape brachiopods (e.g. FOERSTE 1930, KOBLUK & MAPES 1989). is critical in the analysis of virtually all fossils, colour is Colour preservation in echinoderms is known to occur in rarely preserved, and thus has attracted the attention of blastoids (e.g. REIMANN 1961, BEAVER & FABIAN 1998), palaeontologists since the late 18th century (BRUGUIÉRE cystoids (PAUL 1967), crinoids (e.g. BLUMER 1960, 1965,

1792). Today, there exists an extensive body of literature WOLKENSTEIN et al. 2006), and echinids (COTTEAU 1881, addressing colour preservation in a wide variety of fossil KROH 2003, HOSTETTLER 2006). Coloured carapaces of invertebrates (e.g. HOARE 1978, MAPES & HOARE 1987). trilobites (see KOBLUK & MAPES 1989 for a review),

Secretion of inorganic and/or organic hard-parts while phyllocarids (STUMM & CHILMAN 1969), and cirripeds (e.g. the animal is alive is usually a prerequisite for colour YAMAGUCHI 1980) are examples for colour preservation preservation in fossils. Most coloured fossils represent in fossil crustaceans. Last but not least, the chitinous calcareous skeletons, predominantly those of molluscs. carapaces of insects may retain colour, if preserved in

A variety of colour patterns and modes of preservation conservation-type Lagerstätten (see HOARE 1978 and have been documented for polyplacophorans (e.g. HOARE MAPES & HOARE 1987 for examples).

& SMITH 1984, HOARE et al. 2002), bivalves (e.g. NUTTALL Trilobites from the Cambrian are the oldest organisms

1969, KOBLUK & MAPES 1989), nautilids (e.g. FOERSTE known to date to display colour patterns (RAYMOND 1922).

1930, KOBLUK & MAPES 1989), ammonites (e.g. MAPES Early coloured nautiloids (e.g. MARSH 1869, KOBLUK &

& SNECK 1987, MAPES & DAVIS 1996), belemnites (JORDAN HALL 1976), brachiopods (SINGH 1979), and gastropods et al. 1975, SPÄTH 1983), and, most important, gastropods (WHITE 1896, RAYMOND 1906) have been reported from 144 SIMON SCHNEIDER & WINFRIED WERNER

Ordovician deposits. The earliest fossil bivalves display- Here, we present additional, better preserved material of ing colour come from the Devonian (OEHLERT 1881, these two taxa that allows both a refined taxonomic 1888). Colour preservation in fossils occurs throughout characterization and more detailed description of the the entire Phanerozoic, but is more abundant in the colour patterns. Moreover, the first information about younger strata. colour pattern preservation in the pteriacean bivalve With regard to the Late Jurassic of Portugal, colour Isognomon rugosus and the club-shaped spines of the preservation was briefly mentioned within the original cidaroid echinids Pseudocidaris lusitanicus and P. description of the gastropod Fuersichella bicornis and spinosa is presented. the heterodont bivalve Coelastarte discus (SHARPE 1850).

Material and methods

The specimens were collected from various sites located in the Lusitanian basin in central Portugal (Text-fig. 1) during several field campaigns organized by F.T. Fürsich and the authors. The Lusitanian basin belongs to a series of syn-rift basins bordering the North Atlantic; the development of the basin, which begins in the Late Triassic, is strongly connected to the opening of this ocean (WILSON

1985, WILSON et al. 1989). Following a first transgressive- regressive megasequence that ended with a subaerial exposure (uppermost Callovian to lower Oxfordian), the Late Jurassic megasequence 2 is characterized by a strike- slip dominated rift-phase of the basin that encompasses the development of sub-basins and activation of salt diapirs. Rift- and halocinetic tectonics result in highly differentiated facies types and environments, including siliciclastic and carbonate deep-water to very shallow, marginal marine to lacustrine sediments, as well as the formation of massive packages of terrestrial red beds. While the Oxfordian is largely dominated by carbonate lacustrine to ammonite-bearing marine facies, siliciclastic marginal-marine and fluviatile-terrestrial environments characterize the Kimmeridgian in various segments of the basin. The influx of siliciclastics starts in the northern part of the basin during the Oxfordian, and successively progrades to the south during the Kimmeridgian and Tithonian. It results in a nearly complete aggradation of the basin at the end of the Jurassic (LEINFELDER 1987,

LEINFELDER & WILSON 1998, WILSON et al. 1989).

Text-fig. 1. Geographical-geological overview of the Lusitanian basin. The material described in this study was collected from the following localities: (1) Cabo Mondego; (2) Vestiaria, west of Alcobaça; (3) Salgados; (4) São Martinho do Porto; (5) Sobral da Lagoa; (6) Conso- lação; (7) Santa Cruz; (8) Arranhó, southwest of Arruda dos Vinhos; (9) Chão da Cruz, southwest of Arruda dos Vinhos; (10) Lameiro das Antas, southwest of Arruda dos Vinhos; (11) Cabo Espichel. Color pattern preservation in Fürsichella n. gen., bivalves, and echinid spines (Upper Jurassic, Portugal) 145

Most of the specimens, included in this study come FELDER 1986, LEINFELDER & WILSON 1998, MANUPELLA from the siliciclastic marginal-marine facies of the upper 1999, WERNER 1986). Oxfordian section at Cabo Mondego, and the early to Many of the fossils were picked from ploughed fields. middle Kimmerdigian Alcobaça Formation and their The Isognomon shells were naturally exposed by weather- lateral equivalents (e.g. the sections of Consolação, São ing. Some of the neritids and echinid spines, and the Coel- Martinho do Porto, Salgados, Vestiaria, and Sobral da astarte specimens from Consolação were mechanically Lagoa). Other specimens come from the upper Kimmer- removed from sediment layers exposed along the Atlantic idgian Sobral Formation (locality Chão da Cruz) and the coastline. The fossils were cleaned using chemical (H2O2, younger upper Kimmeridgian to lower Tithonian Arranhó Rewoquat) and/or mechanical techniques. Formation and their equivalents (localities Lameiro das The material is deposited in the Geological Museum Antas, Santa Cruz), and a single specimen from the Port- (GML) in Lisbon, Portugal, the Bayerische Staatssamm- landian facies (= Tithonian) of Cabo Espichel (for details lung für Paläontologie und Geologie (BSPG) in Munich, on the geological setting and stratigraphy see AZEREDO et Germany, and the British Museum of Natural History al. 2005, FÜRSICH 1981, FÜRSICH & WERNER 1991, LEIN- (BMNH), London, UK. Systematic paleontology

The higher taxonomy of Gastropoda follows BOUCHET & Fuersichella bicornis (SHARPE, 1850)

ROCROI (2005). The systematic treatment of the bivalves Text-fig. 2A-C, Text-fig. 7A-H follows AMLER et al. (2000). The echinids are classified * 1850 Neritina bicornis, n. s. – SHARPE: 192, pl. 24, fig. 7. according to BARRACLOUGH FELL & PAWSON (1966). 1986 Neritina bicornis. – WERNER: 19. Measurements are given for the smallest (min) and largest 1986 Nerita bicornis. – WERNER: 35, 47, 49, 53, 55, 58. specimen (max) of a taxon; h = height, w = width, l = Holotype. Specimen illustrated in Text-fig. 7G (originally figured by length, t = thickness. SHARPE 1850, pl. 24, fig. 7), deposited in the British Museum of Natural History under accession number BM(NH) G 76588.

Locus typicus. „two miles south-west of Alenquer“ (SHARPE 1850); Class Gastropoda CUVIER, 1797 central Portugal.

Subclass Neritimorpha KOKEN, 1896 Stratum typicum. „subcretaceous series“ (SHARPE 1850). The term „subcretaceous series“ comprises a regionally and stratigraphically Order Cycloneritimorpha BANDEL & FRÝDA, broad spectrum of Jurassic and Lower Cretaceous sediments. The region of the type locality, south-west of Alenquer according to SHARPE, 1999 is characterized by sediments of the Arranhó Formation (the former „Pteroceriano“ facies), which is of late Kimmeridgian to early Superfamily Neritopsoidea RAFINESQUE, 1815 Tithonian age (LEINFELDER 1986, ZBYSZEWSKI 1965).

Family Neritopsidae GRAY, 1847 Material. With the exception of a single specimen that comes from Arranhó (Upper Kimmeridgian), the best preserved coloured Fuersi- Subfamily Naticopsinae WAAGEN, 1880 chella shells (shown in Text-fig. 7) are from Consolação (Lower Kim- Genus Fuersichella n. gen. meridgian). Nevertheless, specimens showing traces of colouration also occur in various other localities, e.g. Sobral da Lagoa (Lower Type species. Neritina bicornis SHARPE, 1850. Kimmeridgian) (GML 4233, 4234, 4235, 4236; BSPG 1984 XXI 546, 547, 548). Diagnosis. Neritopsid gastropod with Crepidula-like Measurements. min: h = 8.5 mm, w = 10 mm; max: h = 33 mm, w = outer shell morphology; last whorl large and oblique; 39 mm. parietal lip with extended and strongly convex callus, Original diagnosis. „Semi-oval, slightly and regularly resulting in reduced D-shaped aperture. Outer lip distinct- convex, with the margins produced into two nearly equal ly auriculate at both ends. pointed ears; spire produced and slightly oblique: colour Derivatio nominis. In honour of Prof. Dr. Franz T. Fürsich. His work significantly improved our knowledge of the fossil invertebrates from, tawny, with thin dark lines radiating from the apex to the and paleoecology of, the Late Jurassic in the Lusitanian Basin, margin, some of which bifurcate; these are crossed by Portugal. 146 SIMON SCHNEIDER & WINFRIED WERNER well-marked lines of growth. Aperture semi-lunar; body- is obviously original. If preserved, the colour seems to lip expanded into a gibbose callosity; outer lip sharp and be fairly durable. Fading-effects, similar to those seen in extended to the two ears.“ (SHARPE 1850). Isognomon (see section below), have not been observed in Fuersichella shells. Absence or faintness of the colour- Additional features. Shell globular, crepiduliform; spire ation is most probably the result of erosion, since incorpo- slightly protruding, with a rounded shoulder; last whorl ration of colour into the shell exclusively is restricted to very large, oblique, almost enclosing the earlier whorls; the outermost shell layer. surface smooth, with marked growth lines; inner lip extended, forming a thick, strongly convex callus pad that Discussion. The overall morphology of the Fuersichella is set off from the spire by a distinct sulcus; aperture D- shell strongly suggests that this form belongs to the Neriti- shaped; outer lip forming sharp-angled, pointed, auricu- morpha, which produce a mixed calcitic-aragonitic shell, late extensions at both ends of the parietal lip. usually with a thin outer calcitic and massive, crossed- lamellar inner aragonitic layer. The original mineralogy Colouration. Generally, the arrangement of the colour of the shell is masked by sparitic recrystallization, and marks in Fuersichella bicornis is fairly variable. The de- protoconchs are not preserved in the Portuguese material. scription of the colour pattern included in SHARPE’s (1850) From the morphologically similar aragonitic Calyptraea- original diagnosis (see above), corresponds well to most cea, Fuersichella clearly differs by the marked convex of our specimens. The colour typically occurs in narrow callus. Moreover, colour patterns are frequently preserved lines, less than 1 mm wide. The lines usually originate in in neritimorphs (COX 1960), and hence may also be used the apex of the shell and run to the aperture without inter- as a feature in classification. ruption. Bifurcating lines occur in some of the specimens. SHARPE (1850) assigned the species to the genus In other specimens, the lines start and end randomly at Neritina. This genus belongs to the superfamily Nerito- the whorls, and new lines are frequently intercalated idea, which is characterized by resorption of the inner between the existing lines in the ventral half of the last walls of the protoconch and teleoconch (e.g. WENZ 1938, whorl. On the other hand, at least one slightly weathered BANDEL 2000, 2001). Sections through two specimens specimen from Consolação shows a single dark conti- from Portugal show that the inner whorls of the teleoconch nuous band, up to 4 mm wide and faint marks of up to are present at least in part, which argues against assigning three additional bands. the Portuguese species to the Neritoidea, and consequent- The shells are typically beige to yellowish-brown in ly Neritina. Moreover, the convex callus and the auricu- colour, while the lines are dark violet-brown. It cannot late extensions of the outer lip clearly distinguish Fuersi- be established as to whether this combination of hues chella from the latter genus, and from the morphologically represents the original colouration or is a diagenetic arte- similar Septaria. fact. However, the pattern formed by the colour stripes

Text-fig. 2. Fuersichella bicornis (SHARPE, 1850). A. Adult specimen, Alcobaça Formation, Lower Kimmeridgian, Sobral da Lagoa; GML 4233; scale bar = 10 mm. B, C. Juvenile specimen with clearly visible spire, Alcobaça Formation, Kimmeridgian, Consolação; BSPG 1984 XXI 546. Scale bars = 5 mm. Color pattern preservation in Fürsichella n. gen., bivalves, and echinid spines (Upper Jurassic, Portugal) 147

In the Neritopsoidea the inner whorls do generally not Remarks. The specimens with preserved colour pattern become dissolved during ontogeny (e.g. BANDEL 2000, are conspicuously small. Full-grown Isognomon rugosus 2001). Crepidula- or limpet-like outer morphologies, may be more than 200 mm long. similar to that of the species described here, are common Colouration. The colour pattern preserved in Isognomon within the family Neritopsidae. Phylogenetically signifi- rugosus consists of up to twenty dark-coloured radiating cant or not, species with smooth outer shell surface are bands of irregular thickness. It is difficult to indicate the generally referred to the subfamily Naticopsinae. Within exact number of bands because some of the bands are this subfamily, the Triassic genera Haliotimorpha and only partly and/or faintly visible. Only two of the speci- Marmolatella, along with the Cretaceous Damesia, have mens display well-preserved colouration of the complete a Crepidula-like outer shell form, but clearly differ from shell. In the other shells, the colour pattern is visible in the new genus Fuersichella because they do not develop the umbonal area, but then fades out towards the ventral a broadly extended convex callus and ear-like extensions part. The colour ribbons usually start at the umbo and of the outer lip. On the other hand, Fuersichella does not run continuously to the ventral margin without forking produce hollow spines like those seen in Haliotimorpha. or anastomosing. In one specimen, a relatively broad band In addition, the new genus differs significantly from is subdivided into two bands near the ventral margin of Marmolatella and Damesia because the spire of the latter the shell. Typically, the bands are not entirely straight, taxa is much more globose than that of Fuersichella. but rather undulate in mm-scale. Undulation seems to be caused by the more or less pronounced concentric growth Class Bivalvia LINNÉ, 1758 lines because it occurs correspondingly in all colour bands. Occasionally, the bands are interrupted at promi- Superorder Pteriomorphia BEURLEN, 1944 nent concentric growth lines, and afterwards continue with Order Pterioida NEWELL, 1965 a lateral shift. Only a single specimen shows well- preserved colour marks in both valves. The colour bands Superfamily Pterioidea GRAY, 1847 do not coincide. Rather, the shell displays distinctly diffe- Family Isognomonidae WOODRING, 1925 rent colour patterns in the opposite valves.

Genus Isognomon SOLANDER in LIGHTFOOT, 1786 The bands are dark reddish-brown to violet-brown. Since the shell is yellowish-brown to relatively dark Subgenus Isognomon SOLANDER in LIGHTFOOT, 1786 greyish-brown, the contrast is sometimes fairly weak. It Type species. Ostrea perna LINNÉ, 1767. is likely that the original colouration of the Portuguese Isognomon shells was considerably brighter based on the Isognomon (Isognomon) rugosus (MÜNSTER, 1835) colouration of extant relatives (see Text-fig. 8D). The Text-fig. 8A-C colour is located in the sub-mm thin, originally calcitic

* 1835 Perna rugosa sp. nov. – MÜNSTER in GOLDFUSS: 105, pl. outer shell layer. The colour is obviously not situated in 108, fig. 2b. a specific shell structure since the pattern disappears 1989 Isognomon (Isognomon) rugosus (MÜNSTER, 1835) – during a simple bleaching-process. FÜRSICH & WERNER: 123, figs 12-13, pl. 8, figs. 1-2; pl. 12, fig. 7; pl. 18, fig. 2; pl. 20, fig. 3.; for a more extensive synonymy and detailed taxonomic treatment see FÜRSICH Superorder Heterodonta NEUMAYR, 1848 & WERNER (1989). Material. 7 coloured specimens from Santa Cruz (3 double-valved, 3 Order Veneroida ADAMS & ADAMS, 1857 right valves, a single left valve); 3 coloured specimens from Cabo Mondego (2 right valves and a single left valve). Several other speci- Superfamily Crassatelloidea FÉRRUSSAC, 1822 mens from both localities show traces of colour (GML 4237, BSPG Family Astartidae D’ORBIGNY, 1844 2005 V 1).

Measurements. min: l = 19 mm, h = 22 mm, t = 4 mm; max: l = Genus Coelastarte BÖHM, 1893 75 mm; h = 76 mm; t = 24 mm. Type species. Astarte excavata SOWERBY, 1819. 148 SIMON SCHNEIDER & WINFRIED WERNER

Original diagnosis. „The species cited above are charac- margin slightly excavated by a small, deep lunette: ventral terized by extraordinary flatness and slightly protruding and posterior margins rounded: ligament concealed in a umbos. Moreover, all of them are strongly oblique, with deep hollow bounded by projecting sharp edges. Surface the umbos situated distinctly in the anterior part; all in rough and uneven with regular concentric wrinkles, and all they should also possess a deeply impressed lunule originally ornamented with dark irregular radiating lines. and ligament area.“ (BÖHM 1893). Posterior muscle placed away from the margin on a broad flat elevation; anterior muscle close to the margin

Coelastarte discus (SHARPE, 1850) and near the umbo; third muscle well-developed.“

Text-fig. 3A-B, 4, 5, 6A-B, 9A-B (SHARPE 1850).

* 1850 Astarte ? discus, n. s. – SHARPE: 177, pl. 21, figs 4a, b, 5. Additional features. Dorsal margin slightly convex; umbo 1893 Astarte ? discus SHARPE. – BÖHM: 6. situated approximately one third of shell length from 1986 Coelastarte discus – LEINFELDER: 34. anterior margin; escutcheon formed by a deep sharp Locus typicus. „between Sobral and Torres Vedras“ (SHARPE 1850); incision, stretching from the umbo towards the posterior Central Portugal. margin, ending above the posterior adductor muscle scar; Stratum typicum. „subcretaceous limestones“ (SHARPE 1850). As noted ligament protected by strongly protruding dorsal shell above, SHARPE used this term for many Jurassic and Cretaceous sediments. Between Sobral de Monte Agraço and Torres Vedras, lime- margin; ontogenetic growth displacement at posterior end stones are common within the Sobral (Upper Kimmeridgian) and of dorsal margin resulting in constriction of inner shell especially the overlying Arranhó Formation (Upper Kimmeridgian - surface margin; muscle scars distinctly depressed (in con- Lower Tithonian) (LEINFELDER 1986, 1987). The type material probably comes from these formations since it is very similar to own collections trast to the original description by SHARPE 1850). Pallial from south of Sobral. line distinct, integripalliate, parallel to shell margin; inner

Material. 4 double-valved coloured specimens from Consolação; a shell margin smooth. Outer shell surface smooth, with single fragmented specimen showing radial striae from Chão da Cruz irregular growth lines. (GML 4230, 4231, 4232; BSPG 2005 V 2, BSPG 1984 XXI 545). Hinge: Right valve with strongly developed wedge- Measurements. min: l = 72 mm, h = 59.5 mm; max: l = 74 mm, h = shaped 3b, corresponding to marked cavity in left valve; 63.5 mm. 3a weakly developed, arcuate; cardinals (2 and 4b) in Original diagnosis. „Shell transverse, broadly obovate, left valve well developed; weak PII situated at dorsal irregular, nearly flat. Dorsal margin straight: umbos very end of nymph; nymph in both valves expressed by a broad small and placed near the anterior extremity: anterior flat area, tapering towards posterior end of ligament.

Text-fig. 3. Coelastarte discus (SHARPE, 1850), Arranhó Formation, Upper Kimmeridgian, Lameiro das Antas. A. Inner side of left valve; GML 4230. B. Inner side of right valve; GML 4231. Scale bar = 10 mm. Color pattern preservation in Fürsichella n. gen., bivalves, and echinid spines (Upper Jurassic, Portugal) 149

often fuse to form broad bands towards the ventral margin. In addition, partial diagenetic encrustations that cannot be removed render a complete counting of the bands impossible. The bands start at the umbo and run to the ventral margin without interruption. Some of the bands remain narrow and delicate, while others fuse and form broad bands, > 5 mm wide. The distribution of broad and narrow elements across the shell is completely irregular. The broad bands may be concentrated in the anterior, central, or posterior part of the valve, or thus are scattered across the shell. It is notable that the colour marks of the left Text-fig. 4. Coelastarte discus (SHARPE, 1850), Arranhó Formation, and right valve do not coincide, but rather differ with Upper Kimmeridgian, Lameiro das Antas. Drawing of specimen Text- regard to pattern in all four specimens. fig. 3B, illustrating the adductor muscle scars, pallial line, and hinge morphology. Scale bar = 10 mm. The shell surface is light to dark beige, while the bands are dark reddish-brown in colour. It is not clear whether

Colouration. SHARPE (1850) recognized „dark irregular this represents the original colour or is the result of fossili- radiating lines“ on the shell of his Coelastarte specimen. zation and diagenesis. An additional fragmented specimen His drawing of a right valve shows a set of nine faintly from Chão da Cruz sheds light on the structure of the visible, narrow radial bands, stretching from the umbo colouration. In that shell, the dark-coloured bands are over approximately half of the shell. Our four coloured eroded, leaving shallow radial grooves in the shell surface. specimens from Consolação exhibit between 20 and 30 Regardless of whether they are original or diagenetically radial bands. It is hardly possible to determine the exact caused, the coloured shell portion must have been less number of bands, because the individual narrow stripes

Text-fig. 6. Coelastarte discus (SHARPE, 1850), Alcobaça Formation, Lower Kimmeridgian, Consolação. Reconstruction of adverse colour patterns in specimen Text-fig. 9A. Scale bar = 10 mm.

erosion resistant, and therefore different in consistency, than the surrounding material.

Discussion. SHARPE (1850: 177) stated that the external and internal features of the single shell-preserved specimen and another single cast in his collection distinguished this taxon from the genus Astarte. On the other hand, he Text-fig. 5. Coelastarte discus (SHARPE, 1850), Sobral Formation; was not able to erect a new genus based on a specimen Upper Kimmeridgian, Chão da Cruz. Fragmented left valve, showing without observable hinge. The characters of the here radial groves where coloured shell material was lost by weathering; BSPG 2005 V 2. Scale bar = 10 mm. presented shells correspond well with those listed for the 150 SIMON SCHNEIDER & WINFRIED WERNER

genus Coelastarte (BÖHM 1893). The Portuguese speci- LORIOL (1890), the main difference between the spines mens differ from all other species of Coelastarte in that of these two species is with regard to ornamentation. In they are more oblique in outline, have a smooth outer P. spinosa granules are restricted to the distal portion of shell surface, extremely flattened shell, and display a the spine, whereas in P. lusitanica granules are distributed peculiar, broadly extended nymph. more regularly on the whole surface of the spine. We observed considerable variability with regard to this feature Class Echinoidea LESKE, 1778 in the spines from Portugal. Therefore, a clear separation of these two species by spine morphology remains impos- Order Hemicidaroida BEURLEN, 1937 sible. As a taxonomic revision is beyond the scope of

Family Hemicidaridae WRIGHT, 1857 this article, and because our material partly stems from localities or formations mentioned by LORIOL (1890) (e.g. Genus Pseudocidaris ÉTALLON, 1859 Santa Cruz, Alcobaça), we consequently use the species Pseudocidaris lusitanicus LORIOL, 1890 names proposed for material from these outcrops.

Pseudocidaris spinosa LORIOL, 1890 Colouration. A wide variety of spine morphotypes occur, Pseudocidaris cf. spinosa including globular, club-shaped, subcylindrical, and elongated types, some of which with transversal constrictions. Pseudocidaris sp. Colour patterns occur in several of these morphotypes, Text-fig. 10A-J but are most abundant in the club-shaped type. The Material. A total of approximately 45 coloured spines have been dis- colouration is composed of rounded, elongate, or com- covered, 10 of which from five different localities have been illustrated pletely irregular spots and blotches of variable size. These in Text-fig. 10 (Lower Kimmeridgian: São Martinho do Porto, Vestia- ria, Salgados, Sobral da Lagoa; Upper Kimmeridgian: Santa Cruz; blotches may occur isolated or fused, resulting in irregular Portlandian: Cabo Espichel) From most of these localities, relatively lines or large flat blotches. As a whole, the pattern is small numbers of coloured specimens have been collected together suggestive of military camouflage. In some specimens, with large quantities of spines lacking colouration. Only in one specific spots and elongated blotches are arranged in axial rows layer at Salgados, all spines show colouration. (GML 4115, 4238, 4239; BSPG 1978 XIX 8, 1984 XXII 351-354, 2005 V 3-4). that may almost reach from the base of the shaft to the tip (Text-fig. 10J). Near the tapering base, a few specimens Remarks. In a monograph treatment of the Jurassic echi- exhibit rings of small blotches around the spine (Text- noderms of Portugal LORIOL (1890) exclusively used fig. 10A). As far as we are aware, there is no correlation coronal features to discriminate individual cidaroid (s.l.) between colour pattern and spine morphotype. All the species, because articulated specimens were rarely more, separation of the two species of Pseudocidaris available. Assignment of isolated spines to certain species based on colour patterns is impossible. is based largely on the co-occurrence of single coronae The blotches are reddish-brown to violet-brown and and isolated spines in the same layer or the same for- contrast well with the lighter, yellowish to orange-brown mation. Here, we present only isolated spines that are background colour of the spines. Cleavage planes of consequently difficult to determine to genus or species broken spines indicate that the coloured area extends up level. The material most closely resembles Pseudocidaris to 1.5 mm into the hard spine material. lusitanica and Pseudocidaris spinosa. According to Colour patterns in related organisms

In this section, we briefly survey colour patterns in selec- LUKES 1974). From the Devonian to Permian, both platy- ted fossil and extant relatives of the five taxa presented ceratid and naticopsid shells from nearly a dozen localities in this study. have been recorded that preserve colour patterns (see

KOBLUK & MAPES 1989 for review). Coloured neriti- Fuersichella. The occurrence of colour patterns in fossil morphs frequently occur in Triassic (e.g. HAAS 1953, neritimorph gastropods is an endless story (COX 1960) TICHY 1980), Jurassic (e.g. STRUCKMANN 1878, FISCHER that begins with Platyceras from the Silurian (KRÍZ & Color pattern preservation in Fürsichella n. gen., bivalves, and echinid spines (Upper Jurassic, Portugal) 151

1908, 1925, COX & ARKELL 1948-1950), and Cretaceous patterns have also been observed in Neocrassina (Coel- strata (MEEK & HAYDEN 1865, WHITE 1883). Finally, in astarte) cotteausia (D’ ORBIGNY) and Neocrassina sp. s.

Cenozoic sediments neritimorphs are commonly str. from the Oxfordian of Cuba (PUGACZEWSKA 1978). preserved with colour. Neritimorph colour pattern types The specimens from Portugal presented here are only include dark spots, spiral, commarginal, or oblique lines, slightly younger than the forms from Cuba. Coloured zigzag lines, and combinations of these types. In several astartids have also been described from Oligocene cases, even different colours and hues are involved (e.g. (NEUFFER 1971: Crassatella) and Miocene strata (SACCO

PRINZ-GRIMM & THALHEIM 2003). Thus, it comes as no 1899: Astarte, own observations: Goodallia). With the surprise that colour patterns are preserved in the neriti- exception of the Middle Jurassic Pressastarte species, morph Fuersichella. colouration in all of these forms is composed of dark radial bands, a pattern that also is expressed in several Isognomon. With a few exceptions, all Paleozoic bivalves extant Indopacific species of Crassatina and Eucrassa- displaying colour belong to the Pterioida (KOBLUK & tella. On the other hand, the overall brownish colour that MAPES 1989). Three of these bivalves, Leptodesma is characteristic for most extant astartids may also have (Leiopteria) pseudolaevis and L. (L.) picta (OEHLERT characterized fossil representatives in this group, but this 1881, 1888) (the oldest coloured bivalves known to date), is generally difficult to evaluate based on fossil material. and „Avicula“ sublobata (PHILLIPS 1836) are placed in Pseudocidaris. Documentation of colour preservation in the superfamily Pteriacea. In the same superfamily, fossil echinids is rare. From the Oxfordian of France, coloured Triassic bakevellids (Hoernesia, Bakevellia) COTTEAU (1881: 222) describes and illustrates traces of have been reported from the German Muschelkalk „une ou deux larges bandes de couleur brune“ in spines (OPPENHEIM 1918, SCHMIDT 1928, HAGDORN 1995). Among of Acrocidaris nobilis. The bands are located in the proxi- the Isognomonidae, however, the Portuguese specimens mal half of the spine. HOSTETTLER (2006) figured spines represent the first showing colour preservation. Coloured of the same species from the Upper Jurassic of Switzer- fossil pteriaceans generally display a pattern composed land that show a faint pattern of wide colour bands in the of irregular radial bands and stripes. A similar colour proximal half of the spine and irregularly distributed small pattern is found in modern Isognomon, e.g. Isognomon spots in the distal part. The exact position of these bicolor (C.B. ADAMS) (Text-fig. 8D). elements may vary among spines. From the same locality, OLDFUSS Coelastarte. Astartellopsis nuda (G ) from the tests of Phymosoma corallinum (COTTEAU) and Stomechi-

Middle Triassic German Muschelkalk is the earliest nus perlatus (DESMAREST) displaying radial colour AGDORN astartid to date showing colour preservation (H patterns are reported (HOSTETTLER 2006). The only other 1995). Middle Jurassic erratic boulders from north- coloured echinid remains we could detect in literature eastern Germany have yielded specimens of Pressastarte are banded spines of Eucidaris zeamais (SISMONDA) from that display the natural overall brownish pigmentation the Upper Miocene of Austria (KROH 2003). (personal communication J. KOPPKA, Greifswald). Colour

Composition of pigments

Knowledge about the substances causing colouration in Tetrapyroles. Cyclic tetrapyroles, the porphyrines, have skeletal remains continues to be incomplete. Chemical been recorded for various gastropods (including certain isolation and decoding of the substances seems to be diffi- neritids) and bivalves (COMFORT 1950), and seem to be cult. Generally, pigments hitherto have mostly been detec- the most widespread pigments in molluscs (HOLLING- ted by spectrometric analysis, rather than extraction. WORTH & BAKER 1991). They are detected by red fluores- Nevertheless, a number of different organic pigments have cence in ultraviolet light. Even in fossils, up to Eocene in been identified from invertebrate skeletons. Here, we age, colour patterns that are no longer recognizable by focus on those pigments recorded for molluscs and echi- the naked eye can be visualized by that method (COMFORT noderms. 1950, NEUFFER 1971, 1972). Linear tetrapyroles, or bili- 152 SIMON SCHNEIDER & WINFRIED WERNER

Text-fig. 7. Fuersichella bicornis (SHARPE, 1850). A-D, F. Alcobaça Formation, Lower Kimmeridgian, Consolação. A. GML 4234. B, C. GML 4235. D. BSPG 1984 XXI 547. F. BSPG XXI 548. E. Juvenile specimen, Sobral Formation, Upper Kimmeridgian, Arranhó, GML 4236. G, H. Holotype, ?Arranhó Formation, Upper Kimmeridgian-Lower Tithonian, BM(NH) G 76588. Scale bars = 5 mm.

proteins, have also been recorded for molluscs (KENNEDY Indigoids. Although present in several gastropods (COM-

1979). FORT 1950), indigoids are normally solved during putre-

faction (TICHY 1980). Melanins. This group of highly durable pigments is composed of protein complexes, also named melanoproteins. Quinones. Representatives of these organic compounds Generally, these substances stain black, brown, or reddish. (fringelite, hypericin, and related substances) have been

Melanins are widely distributed in molluscs (COMFORT identified in fossil and extant crinoids (BLUMER 1960,

1950, 1951) and echinoderms (KENNEDY 1979). 1965, WOLKENSTEIN et al. 2006) where they cause violet to reddish colouration. Another group of quinones, the Polyenes. For a variety of extant gastropod and bivalve spinochromes, are common in spines and tests of extant taxa and Nautilus HEDEGAARD et al. (2006) identified echinids, among other unidentified pigments (ANDERSON polyenes in shell pigmentation. Polyenes are reported to et al. 1969), whereby up to six different spinochromes occur in the neritids Nerita albicilla LINNÉ and Smaragdia may co-occur in a single species. viridis LINNÉ among other gastropods. Polyenes have been detected in coloured parts of the shells, but are absent In conclusion, it is obvious that in both extant molluscs from colour-less regions. All molluscs studied by HEDE- and echinoderms - and most likely also in the Jurassic

GAARD et al. (2006) contain polyenes in their coloured fossils from Portugal - colouration is caused by a mixture shell parts. of different pigments, rather than by a single substance. Whether these pigments occur in the fossils in unaltered Carotenoids. Carotenoid pigments have been detected form, or are (partially) decayed or substituted, remains (HEDEGAARD et al. 2006) in the shells of certain cypraeids. to be analyzed. Generally, these substances are subject to rapid decay, unless bound by proteins (TICHY 1980).

Development of specific colour patterns

Colour secretion in molluscs occurs in specialized secre- STURGEON 1978, TICHY 1980), but, especially in neriti- tory cells along the mantle edge (OBERLING 1968, TICHY morphs, including Fuersichella bicornis, patterns may

1980), and is controlled by neural activity (ERMENTROUT vary within a single species. et al. 1986). The exact position of these cells and the Current knowledge about colour secretion in echinid functional details of colour secretion, however, remain skeletons is poor. Several pigments, including spino- enigmatic (COMFORT 1950). Continuous secretion of chromes, do not only occur in the hardparts of echinids, colour causes radial or spiral bands, or, in extreme, an but also in soft tissue (ANDERSON et al. 1969, PANTAZIS entirely coloured surface. Interruptive secretion results 2006). As a result, it is likely that these tissues are capable in spots, blotches, or dashed lines. Oblique patterns are of colour deposition also within the hard parts, because the result of continuous displacement of the loci for pigments also occur in membranes that are responsible secretion. Combination of these elements produces nearly for test- and spine-repair (CUÉNOT 1848). However, the all of the complex patterns found in mollusc shells. In details of the secretion process have not yet been docu- general, colour patterns are genetically coded (HOARE & mented.

Text-fig. 8. A-C. Isognomon rugosus (MÜNSTER, 1835), Upper Kimmeridgian, Santa Cruz. A. GML 4237. B, C. BSPG 2005 V 1. Scale bar =

10 mm. D. Isognomon bicolor (C.B. ADAMS, 1845), Recent, San Andres, Colombia, BSPG 2007 I 6. Scale bar = 5 mm. Color pattern preservation in Fürsichella n. gen., bivalves, and echinid spines (Upper Jurassic, Portugal) 153 154 SIMON SCHNEIDER & WINFRIED WERNER

Text-fig. 9. Coelastarte discus (SHARPE, 1850), Alcobaça Formation, Lower Kimmeridgian, Consolação. A. View on right valve; GML 4232. B. View on left valve; BSPG 1984 XXI 545. Scale bar = 10 mm.

Possible models for colour preservation

Apart from the Neritimorpha, colour preservation in during mechanical or chemical erosion (TICHY 1980). In fossils is a rare phenomenon (COX 1960, HOLLINGWORTH fossil echinoderms, the cleavage plains show that colour

& BARKER 1991). Pigments may be destroyed or removed is incorporated in the outer layer of the skeleton (BEAVER prior to preservation by various circumstances. The ultra- & FABIAN 1998, own observations) and was preserved in violet rays of sunlight may cause bleaching and decay, the spines regardless of recrystallization. However, colour especially of carotenoids (TICHY 1980, HOLLINGWORTH & preservation also depends on the original mineralogy of

BARKER 1991), and indigoids are subject to biological the organisms. The calcitic skeletons of echinoderms, as

(bacterial) decay that occurs during putrefaction (TICHY well as the outer calcitic layer of neritimorphs, are much 1980). Moreover, oxygenation, along with temperature more likely to preserve colour than the relatively instable and chemical composition of circulating pore fluids, plays aragonitic shells of many other molluscs. a major role in the destruction of pigments. Colours may In general, features that are indicative of many conser- be altered or substituted, or become pseudomorphic vation type Lagerstätten, including rapid burial, oxygen

(TICHY 1980) or inverted by incorporated oxides (STRAUCH isolation, and early cementation, also favour colour pre-

1985). servation (HOLLINGWORTH & BARKER 1991). Such condi- On the other hand, melanins are highly resistant to tions sometimes occur throughout an entire sediment layer organic and acidic solvents (COMFORT 1951) and also or package, which is the case in both localities of coloured porphyrins are remarkably durable (HOLLINGWORTH & Isognomon, for the banded Coelastarte shells, and for

BARKER 1991). If solution or recrystallization of shells part of the blotchy Pseudocidaris spines (e.g. locality occurs conservatively, certain original patterns maintain Salgados) from Portugal. In other instances, clayey sedi- in the sparitic matrix, or even in silicified gastropod shells ment or concretions are only locally distributed, and so

(HAAS 1953, YOCHELSON & KRÍZ 1974, KRÍZ & LUKES are invertebrates showing colour preservation. In most

1974) or on steinkerns (MAPES & DAVIS 1996, own Portuguese Late Jurassic localities yielding coloured observations). However, colour in molluscs is typically echinid spines, only a few coloured specimens occur restricted to the outer shell layer(s) (HOLLINGWORTH & among hundreds of colourless spines.

BARKER 1991), and thus is easily removed by abrasion

Text-fig. 10. A, E. Pseudocidaris cf. spinosa. B, D, F. Pseudocidaris sp.. C, G, I, J. Pseudocidaris lusitanicus LORIOL, 1890. H. Pseudocidaris spinosa LORIOL, 1890. Scale bar = 5 mm A. Lower Kimmeridgian, Alcobaça Fm, Salgados; BSPG 2005 V 3. Circular rows of small spots at the base of the shaft; vertical rows or irregular pattern in the distal portion of the spine. B. Lower Kimmeridgian, Alcobaça Fm, São Martinho do Porto; BSPG 1984 XXII 351. C. Lower Kimmeridgian, Alcobaça Fm, Vestiaria; BSPG 1984 XXII 352. D. Portlandian, Cabo Espichel; BSPG 1978 XIX 8. E. Lower Kimmeridgian, Alcobaça Fm, Salgados; BSPG 2005 V 4. Extremely irregular shape and dispersion of blotches. F. Lower Kimmeridgian, Sobral da Lagoa, limestone quarry; BSPG 1984 XXII 353. G. Lower Kimmeridgian, Alcobaça Fm, Vestiaria; GML 4115. Spine with central constriction; irregular distribution of large rounded spots. H. Upper Kimmeridgian, Santa Cruz; GML 4238. I. Lower Kimmeridgian, Alcobaça Fm, Vestiaria; BSPG 1984 XXII 354. Club-shaped spine with fine colour pattern composed of small single spots and few spot fusions. J. Lower Kimmeridgian, Alcobaça Fm, Vestiaria; GML 4239. Elongated spots are arranged in rows running from the base to near the distal end of the shaft. Color pattern preservation in Fürsichella n. gen., bivalves, and echinid spines (Upper Jurassic, Portugal) 155 156 SIMON SCHNEIDER & WINFRIED WERNER

Possible function of colour patterns A relatively complete survey of the circumstances that established. We suppose that C. discus was an epibenthic may have resulted in the development of colour patterns recliner based on the fact that the large and extremely and their functions in marine invertebrates has been pre- flat shell appears completely unsuitable for burrowing. sented by KOBLUK & MAPES (1989). These authors pro- Attachment by byssus can be excluded because the shell pose that the necessary disposal of metabolic by-products lacks a byssal gap. Moreover, byssal threads have not was the initial step towards colour secretion; this been recorded for any astartids. Nevertheless, it has been hypothesis is based on biochemical analyses (COMFORT reported from „normally-shaped“, rapid-burrowing extant 1950, 1951). Moreover, a relationship of colour and diet astartids that some spend part of their life on the bottom has been documented for certain extant gastropods (TICHY surface (STANLEY 1970).

1980, HEDEGAARD et al. 2006). Nevertheless, HEDEGAARD Finally, the evolution of the visual sense in animals et al. (2006) question the hypothesis that mollusc colours may have resulted in a variety of interactive functions of are the result of waste disposal based on the fact that colour (KOBLUK & MAPES 1989). In bivalves and echinids, molluscan excretory organs are quite effective. colour patterns are hardly sensible for inner-specific In a second step, pigments may have been utilized in communication, since these animals are not socially or- stabilization of the shells or reduction of abrasion ganized, and mating just occurs in the form of releasing

(KOBLUK & MAPES 1989). This may be true of melanins sexual products into the seawater. Moreover, the visual or porphyrines, which are intimately associated with the abilities of these animals are rather low. The latter is also shell (COMFORT 1950, KOBLUK & MAPES 1989), but certain- true of most gastropods, regardless of whether they ly does not apply to soluble pigments such as indigoids engage in mating sensu stricto. On the other hand, it is or carotenoids. In a single fragment of Coelastarte, we unlikely that any of the creatures discussed herein was observed radial depressions (Text-fig. 5), resulting from poisonous and therefore coloured for deterrence. Since the erosion of coloured shell material. This feature seems all of the organisms treated herein are grazers or feed on to be indicative of a structural function of the pigments, suspension, colour patterns may have been effective because the colour could only be removed together with against detection by predators. It is well documented that the shell. Whether these shell segments were originally blotches, spots or vivid stripes render well-defined shapes strengthened by the pigments cannot be determined. and result in an optical break-up of outlines (COTT 1957,

Colour in exoskeletons may also protect soft tissue from WICKSTEN 1983). Of course, camouflage is most sensible light or overheating (ENDLER 1978) or serve as a filter for for solitary organisms (WICKSTEN 1983). In Isognomon protection against certain wavelengths, especially harmful rugosus, this utility is lost where the animal occurs as a

UV-radiation (COMFORT 1951, KOBLUK & MAPES 1989). semi-infaunal mud-sticker in dense colonies, e.g. in vari- However, the latter is rather unlikely with regard to Fuer- ous Jurassic localities in Portugal. If Coelastarte discus sichella since these large neritimorphs were relatively was an epibenthic recliner, this organism may have pro- thick-shelled and therefore probably opaque. All the fited by a rather effective camouflage produced by the more, the heavy valves of Isognomon rugosus have cer- banded shell, especially in light-flooded shallow waters tainly been sufficiently effective in protecting the animal or between seaweeds. On the other hand, all other banded from excessive radiation. In living echinids, the spines astartids mentioned above were predominantly infaunal, are covered by a thin layer of soft tissue that is more or and thus colour was developed without a „visible“ sense. less translucent. Moreover, the spines do not contain any In neritimorph gastropods, colour patterns are general organs or structures that require protection. Accordingly, equipment (COX 1960), and therefore their effectiveness Coelastarte discus may be the only organism considered can be studied based on extant representatives. To the in this study that may have benefited from the light-protec- human eye, they generally appear to be relatively success- tion effect of colour since it is relatively thin-shelled. fully hidden in their natural habitat while grazing on the However, the life position of C. discus has not yet been sea bottom, so this may also be true of the Jurassic Fuer- Color pattern preservation in Fürsichella n. gen., bivalves, and echinid spines (Upper Jurassic, Portugal) 157 sichella. The broad convex callus of Fuersichella seems Although these sea urchins were well-armoured with their to be rather unsuitable for living on hard substrates. On big club-shaped spines, blotchy colouration acting against the other hand, the extended auriculate outer lip may have detection might have provided additional protection. On protected the animal from sinking too far into the soft shelly sand or mud, or among seaweeds, the echinids may sediment. have blended in with the background, and thus became The colour pattern developed by Pseudocidaris is best nearly invisible for pycnodont fishes, which probably compared to as military camouflage, and may have had were their most vicious predators based on the frequent exactly the same function in the natural environment. documentation of pycnodont teeth in the same localities. Conclusions

The preserved colour patterns reported in this study occur coincide. While Isognomon preserves colour in the outer in five quite different organisms, which, however, all calcitic shell layer that is subject to fading, colour is thrived in similar marginal marine habitats. The neriti- intimately associated with radial shell parts in Coel- morph gastropod Fuersichella bicornis was probably astarte. In the regular echinids Pseudocidaris lusitanica adapted to soft bottoms based on shell shape, and exhibits and P. spinosa, coloured blotches occur all over the club- a considerable intra-specific variability with regard to shaped spines. In these organisms, colouration may have the narrowly striped colour pattern. Both the semi- been effective as camouflage. In Isognoman, this is less infaunal pteriacean bivalve Isognomon rugosus and epi- probable because the organism often occurs in dense benthic reclining astartid Coelastarte discus are vividly colonies. banded, and the patterns on the opposite valves do not Acknowledgements

We thank H. Hess (Basel), B. Hostettler (Glovelier), A. of Fuersichella. The photographs were taken by G. Kroh (Vienna), and K. Wolkenstein (Innsbruck) for useful Janssen (Munich) and S. Tracey (London). We are also advice regarding coloured echinids. J. Koppka (Greifs- grateful to M. Krings (Munich) for improving the English. wald) and A. Nützel (Munich) are acknowledged for M. Heinze and B. Niebuhr (Würzburg) are thanked for helpful comments on astartids and neritimorphs. J. Todd improvement of the manuscript. The project was partially (London) kindly provided images and data of the holotype finanzed by the DFG (Fu 131/31-2, and We 1152/2-2).

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