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The Morphology and Sculpture of Ossicles in the Cottidae (Teleostei) of the Baltic Sea

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The Morphology and Sculpture of Ossicles in the Cottidae (Teleostei) of the Baltic Sea Estonian Journal of Earth Sciences, 2010, 59, 3, 216–237 doi: 10.3176/earth.2010.3.04 The morphology and sculpture of ossicles in the Cottidae (Teleostei) of the Baltic Sea Tiiu Märssa, Janek Leesb, Mark V. H. Wilsonc, Toomas Saatb and Heli Špilevb a Institute of Geology at Tallinn University of Technology, Ehitajate tee 5, 19086 Tallinn, Estonia; [email protected] b Estonian Marine Institute, University of Tartu, Mäealuse Street 10a, 12618 Tallinn, Estonia; [email protected], [email protected], [email protected] c Department of Biological Sciences and Laboratory for Vertebrate Paleontology, University of Alberta, Edmonton, Alberta T6G 2E9 Canada; [email protected] Received 31 August 2009, accepted 27 May 2010 Abstract. Small to very small ossified structures (ossicles) such as cephalic horns, dermal tubercles, fin-ray rods, lateral-line canal segments, branchial tooth plates and gill-raker tubercles of three species of the teleostean fish family Cottidae inhabiting the Baltic Sea are described and for the first time illustrated with SEM images, with emphasis on their microscopic external features. The morphology and sculpture of these small ossicles notably differ in the three genera, but differences can also occur between males and females, as documented here in Myoxocephalus scorpius (Linnaeus). The detailed features of such ossicles are potentially useful for taxonomic identification in sea-floor sediments or gut contents, in a manner similar to their use in many modern and fossil fishes, and may also prove useful as phylogenetically important characters at generic and familial levels. Key words: dermal ossicles, Cottidae, Cottoidei, Cottiformes, Actinopterygii, Baltic Sea, Myoxocephalus, Triglopsis, Taurulus. INTRODUCTION Beznosov 2001, 2005) and extant chondrichthyans (Reif 1985), and has been explained as imprints of epidermal Exoskeletal elements such as scales, tesserae and bony cells (Gross 1973) or as a reflection of the composition plates of Palaeozoic jawless vertebrates (agnathans) and of the underlying tissue (e.g. Schultze 1977; Gayet & fishes with jaws (gnathostomes) have been shown to Meunier 1986; Meunier & Gayet 1992). According to exhibit characteristic morphology, sculpture and micro- Afanassieva (2004), what she called microsculpture structure, all of which have been useful for taxonomic (= ultrasculpture of this paper) is a result of the mode identification, systematics and phylogenetic analyses of of ossification or may have provided better attachment these fossils (Gross 1967; Karatajūtė-Talimaa 1978; of the epidermis on the scale’s surface. Märss (2006) Märss & Ritchie 1998; Wilson & Caldwell 1998; Wilson distinguished seven main ultrasculpture types, including & Märss 2004, 2009; Märss et al. 2006, 2007). The microtubercles and united micronodules. She showed shape and the sculpture pattern of the elements also that on the one hand similar main types of ultrasculpture vary, depending on their position on the body, the occur on the scales of representatives of more than one developmental age of the individual and the habitat of the higher taxonomic group (i.e. the same general type is species or group. The sizes of these microscopic sculpture found in two or more subclasses), but on the other hand, elements differ greatly. For example, the width of ridges the finer details of the ultrasculpture are more specific and grooves on agnathan thelodont scales is 20–200 µm, and allow the subdivision, for example, of the subclass while the length may coincide with the length of the scale Thelodonti, into four orders. She also supposed that (up to 3.7 mm). Smaller sculptural elements, measured in the different ultrasculpture types within Thelodonti tens of micrometres only and referred to as ultrasculpture may indicate the polyphyletic nature of this subclass. by Märss & Gagnier (2001) and Märss (2006), have Nevertheless, the origin of most ultrasculpture types has been illustrated during the last few decades, ever since remained unclear. the scanning electron microscope was available for use. The work of Cockerell (1913) convinced ichthyo- Such a very fine pattern has been observed on agnathans logists of the great value of modern fish scales for (Gross 1967; Blieck 1982; Märss 1986; Long et al. identification and classification purposes. His investi- 1988; Afanassieva 2004), early gnathostomes (e.g. Gross gations of the scales of Teleostomi were illustrated by 1973; Schultze 1977; Derycke & Chancogne-Weber 1995; many drawings of structural details and photographs. 216 T. Märss et al.: Ossicles of Baltic cottids Similar studies on the morphology of teleostean scales change in different fish habitats and populations, whereas were continued by Taylor (1916), Haque (1955), Batts such effects can be observed in recent fishes (e.g. Poulet (1964) and many others. et al. 2004). The introduction of scanning electron microscopy The results of this investigation might serve for (SEM) initiated a new approach in the study of fish several purposes in ichthyology: for analysis of feeding scale morphology and surface ornamentation (DeLamater preferences of fishes (while studying their stomach & Courtenay 1973, 1974; Hughes 1981; Bonwick et al. content), for determination of fish migrations (e.g. Daniels 1989; Sire & Meunier 1993) and ontogenic develop- & Peteet 1998), for taxonomic and systematic studies ment of the ornamentation on scales (Sire 1986). The and for phylogenetic analyses. The microscopic ossicles comprehensive work by Roberts (1993) gave descriptions, studied here can provide much of the same kind of SEM investigations and comparison of spined scales of taxonomic information as available under ideal circum- Teleostei, including representatives of the Scorpaeni- stances from the study of fish otoliths (e.g. Assis 2003), formes in Percomorpha sensu lato. The sculpture char- but ossicles and otoliths can be recovered in different acteristics of scales have occasionally been used as preservational situations. Microremains obtained from taxonomic criteria for cypriniform (Cyprinidae and drill cores of relatives of modern fishes can help elucidate Cobitidae) fish species (e.g. Kaur & Dua 2004; Jawad the formation of the fish fauna of the Baltic Sea during 2005; Johal et al. 2006; Jawad & Al-Jufaili 2007), but different phases of its development. Such data can also the fine details of the external surface of scales and show the content and abundance of fishes in the fresh tubercles have more often been illustrated in publications water of the Baltic Ice Lake through the Preboreal on histology (e.g. Sire et al. 1997; Meunier & Brito brackish phase of the Yoldia Sea and freshwater Ancylus 2004). Usually, however, when a new fish was dis- Lake stages, and the brackish-water Litorina Sea Stage. covered, not much attention was paid to its smaller The results obtained can be utilized also in archaeology. bony ossicles. The ultrasculpture of dermal tubercles Different fish and seal bones have been used in archaeo- becomes visible only at very high magnifications and is logical studies (Lõugas 1997, 1999; Paaver & Lõugas therefore even less investigated than the sculpture. An 2003), but these are never as numerous as microscopic exception was the groundbreaking monograph of Reif external elements, denticles, tubercles, scales, etc. of (1985), who gave examples of ultrasculpture, including fishes. peculiar polygons on the scale surfaces of sharks. This paper deals with the dermal tubercles and other To some extent the material under study in the ossicles of representatives of three genera of the family present work (tiny ossified structures (ossicles), which Cottidae found in the Baltic Sea. The results of the study have survived from chemical preparation of the fresh were first reported during the 13th European Congress or preserved fish specimen) bears comparison with the of Ichthyology in Klaipeda in 2009. Short descriptions material dealt with in palaeontology (fossilized micro- of the units (within Scorpaeniformes) are given in Märss remains, which have escaped decay, sedimentation and et al. (2009). the processes of fossilization and chemical preparation Cottidae have at times been classified within the of the rocks). The main aim of the present study is to order Scorpaeniformes in the suborder Cottoidei, super- examine and describe the small ossicles of Cottiformes family Cottoidea (e.g. Yabe 1985), but inclusion in from the Baltic Sea, with particular emphasis on their Scorpaeniformes is controversial (Nelson 2006). Recent fine surface structures, sculpture and ultrasculpture, at a molecular studies suggest that cottoids are not close to level of detail that has proven useful in the study of Scorpaeniformes (e.g. Shinohara & Imamura 2007), fossil vertebrate microremains. This comparison can be and Wiley & Johnson (2010) most recently classified informative for palaeontological work. the former Cottoidei in a separate order Cottiformes. One potential benefit for fossil studies is that under- The families Cottidae (possibly not monophyletic; Smith standing the morphology and degree of variability of & Wheeler 2004), Cyclopteridae, Liparidae, Psychrolutidae ossicles in living groups of fishes may contribute to our and Agonidae (also possibly not monophyletic), among understanding of the possible sources of variation in others, are included within the suborder Cottoidei. Palaeozoic taxa. For example, in palaeoichthyology no Descriptions of comparable ossicles of selected species differences have yet been recognized
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