Palaeontologia Electronica a Possible Fossil Paralarva (Cephalopoda
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Palaeontologia Electronica palaeo-electronica.org A possible fossil paralarva (Cephalopoda: Coleoidea) from the Solnhofen Lithographic Limestones (Upper Jurassic, southern Germany) Joachim T. Haug, Isabelle Kruta, and Carolin Haug ABSTRACT We describe a possible fragmentary gladius of a paralarval cephalopod from the Solnhofen Lithographic Limestones, southern Germany (Upper Jurassic, Lower Titho- nian). The specimen is 3 mm long and leaf-shaped, representing mainly the median field of the gladius. Faint remains indicate the former positions of the lateral fields. The median line is marked by a sharp keel. On the posterior end there is a pear-shaped structure. Anteriorly a kite-shaped area is ornamented with small pits. The principle morphology resembles, for example, certain trachyteuthidids, based on shape, pres- ence of a keel and an ornamented anterior kite-shaped region. Also resemblances of certain muensterellid characters are pointed out. The special morphology of the here described specimen is interpreted as ontogenetically caused. This together with its small size makes a paralarval status of the specimen possible, although we cannot provide the necessary ecological evidence for this assignment. A clear ascription to any systematic group is hindered by the fact that there is no comparable material of similarly small size available. Joachim T. Haug, University of Greifswald, Zoological Institute and Museum, Department of Cytology and Evolutionary Biology, Soldmannstr. 23, 17487 Greifswald, Germany [email protected] Isabelle Kruta, Yale University, Department of Geology and Geophysics, PO Box 208109, New Haven Connecticut 06520-8109, USA and American Museum of Natural History, Division of Paleontology (Invertebrates), New York, New York 10024, USA [email protected] Carolin Haug, University of Greifswald, Zoological Institute and Museum, Department of Cytology and Evolutionary Biology, Soldmannstr. 23, 17487 Greifswald, Germany [email protected] Keywords: palaeo-evo-devo; cephalopods; gladius; fossilized ontogenies; larva; fluorescence microscopy PE Article Number: 15.3.28A Copyright: Palaeontological Association November 2012 Submission: 20 March 2012. Acceptance: 15 October 2012 Haug, Joachim T., Kruta, Isabelle, and Haug, Carolin. 2012. A possible fossil paralarva (Cephalopoda: Coleoidea) from the Solnhofen Lithographic Limestones (Upper Jurassic, southern Germany) . Palaeontologia Electronica Vol. 15, Issue 3;28A,8p; palaeo-electronica.org/content/2012-issue-3-articles/313-possible-fossil-paralarva HAUG, KRUTA, & HAUG: POSSIBLE FOSSIL PARALARVA INTRODUCTION The hatching stage of a metazoan that differs significantly in morphology, behavior or other aspects from the adult is termed larva in many dif- ferent taxa. The term is treated differently in the lit- erature, varying in the exact criteria that need to be fulfilled to apply it (see, for example, discussions in Hickman, 1999; Young, 2002; Minelli, 2003). For cephalopod molluscs this dispute has partly been avoided by introducing the term paralarva for the early developmental phase that follows hatching and resembles the adult in gross morphology, but differs in its ecological niche (Young and Harman, 1988); paralarvae are planktonic. This specialised early life stage is found in some, but not all modern cephalopods. As for many metazoan taxa, the ontogeny of cephalopods is an important field of research (e.g., Tsuchiya and Okutani, 1991; Boletzky, 2003; O'Shea et al., 2007), including studies of paralarvae (Tsuchiya et al., 1991; Vec- chione et al., 1992, 2001; Haimovici et al., 2002). Investigating the ontogeny of fossil represen- tatives of Cephalopoda is, as for any other taxon, more complex and restricted in many aspects as compared to investigating the ontogeny of extant cephalopods. For fossil cephalopods the ontogeny of a certain species is usually reconstructed by investigating the shell, as this is what remains of the animal, the soft body parts being only pre- served in exceptional cases (e.g., Weitschat, 1986; Fuchs et al., 2009; Kruta et al., 2011). Coleoid cephalopods possess an interior shell or remains of it that gained different names in dif- FIGURE 1. Comparison of gladius shapes between ferent subtaxa. For coleoids with a so-called glad- older (1.1, 1.3) and younger stages (1.2, 1.4) of the same species of Coleoidea. 1.1-2. Gladii of Onykia ius, growth lines on the latter are used for robusta (Verrill, 1876). Gladius of younger stage much reconstructing the life histories in extant species stouter than that of older stage. Redrawn after (Perez and O'Dor, 2000), but these growth lines Tsuchiya and Okutani (1991, figures 18, 19). 1.3-4. also yield significant information in fossil coleoids Gladii of Pholidoteuthis massyae (Pfeffer, 1912), (Fuchs et al., 2009). Thus, aspects of both extant exhibiting shape differences between the different and fossil ontogenies of Cephalopoda can be stages. Redrawn after O’Shea et al. (2007, figures 36, reconstructed based on few or even a single 37). mature specimen. With such an approach many aspects of the ferences demands for the availability of several ontogeny of a fossil cephalopod can be accessed. specimens representing consecutive developmen- Yet, differences in the total shape of the gladius tal stages (= semaphoronts). The approach to can be difficult to infer exclusively from the growth reconstruct the ontogeny of coleoids based on a marks of a single mature specimen. Although onto- series of individuals has been applied for details of genetic changes of the coleoid gladius are gener- the soft parts of extant species (O’Shea et al., ally considered to be minor (Boletzky, 2003), in the 2007). For such approaches small-sized speci- few examples in which actual gladii of different mens are of importance, especially for accessing growth stages of the same species have been information on the earliest post-hatching phase, depicted in detail, their shapes appear to differ the paralarva. Early post-hatching specimens of quite significantly (Tsuchiya and Okutani, 1991; fossil cephalopods are relatively rare (e.g., Mapes O’Shea et al., 2007; Figure 1). Identifying such dif- et al., 2007; Mapes and Nützel, 2009). No speci- 2 PALAEO-ELECTRONICA.ORG men of a fossil coleoid being small enough to pos- (2007). Drawings from the literature were redrawn sibly represent a paralarva has been described to using Adobe Illustrator CS3 and Inkscape. date. We report here a single specimen from the RESULTS lithographic limestones of Solnhofen (Upper Juras- Description of the Specimen (Figure 2.1-3) sic, southern Germany) that is supposed to repre- sent an early developmental stage of a coleoid, We use the terminology as applied by Fuchs possibly a paralarva. To emphasize the importance et al. (2009) for fossil gladii. The specimen is rela- of this find we discuss the different approaches of tively flat, but slightly more elevated in the median reconstructing developmental sequences, either region (Figure 2.1, 2.3). Based on this elevation the from growth marks of a single mature specimen or specimen is most likely seen in dorsal view. The based on a series of semaphoronts. outline is more or less oval to leaf-shaped. Its max- imal length is 3 mm. On the widest part it measures MATERIAL AND METHODS 1.6 mm. The area preserved in this specimen is interpreted as representing mainly the median field Material of a gladius. Remains of the hyperbolar zone/lat- The single specimen described here was eral fields can still be detected under fluorescence found by Michael Fecke, Langenberg, in the hobby settings (Figure 2.2), but are too weakly preserved quarry Blumenberg near Eichstätt, which belongs to infer the outline. Thus, the original maximal to the area of the Solnhofen Lithographic Lime- width of the specimen is not accessible. Whether stones (Upper Jurassic, Lower Tithonian, lower the weak preservation of the lateral fields reflects Hybonotum zone, Riedense subzone; Schweigert, original structural conditions or is a preservational 2007). The specimen is preserved as part only, a artifact cannot be determined. The midline of the counterpart is not available. It is tiny (ca. 3 mm), median field is marked by a prominent keel. The but is preserved with substance differing from the keel is very narrow (ca. 60 µm) along the anterior surrounding matrix as its autofluorescence capabil- part of ca. 2 mm length. Then the keel widens to a ities suggest, possibly consisting of calcium phos- pear-shaped structure towards the apex with a phate (Haug et al., 2009). The fossil displays a maximum width of 0.45 mm. Closer to the apex the recognizable relief. It is now part of the collection of keel becomes more slender again, measuring the Staatliches Museum für Naturkunde Stuttgart about 0.13 mm in width. A kite-shaped area in the under SMNS 67904b. Also preserved on the same anterior 40% (median) to 30% (lateral) of the entire slab is a small specimen of a mantis shrimp, tenta- gladius length bears a surface ornamentation. tively assigned to ?Sculda pusilla (SMNS 67904a; Under fluorescence this ornament appears like a cf. Haug et al., 2010). reticulate pattern (Figure 2.2). In combination with the stereo image it becomes clear that this pattern Methods is in fact a cluster of small pits (Figure 2.3). The Due to the small size of the specimen it was diameter of the pits ranges from 20-70 µm. necessary to apply higher magnification imaging. A simple macrograph was recorded under standard DISCUSSION North-West lighting. Under the same setting a ste- Cephalopod Affinities reo image was recorded. Additionally, the speci- men was documented under a Zeiss Axioskop 2 The here described specimen is interpreted fluorescence microscope under green light (546 as a small fragmentarily preserved gladius of a nm). Images were processed following the protocol coleoid cephalopod (Figure 3.1). Its shape is rela- of composite fluorescence imaging (Haug et al., tively stout, yet early developmental stages of 2008; Haug et al., 2009; Kerp and Bomfleur, 2011) cephalopods are usually less elongate than they using CombineZM/ZP, Microsoft Image Composite are as adults.