Zemes un vides zinātnes Earth and Environment Sciences - Latvijas ...

Zemes un vides zinātnes Earth and Environment Sciences - Latvijas ... Zemes un vides zinātnes Earth and Environment Sciences - Latvijas ...

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ACTA UNIVERSITATIS LATVIENSIS, 2004, Vol. 679, pp. 14-21 Microrelief on the exoskeleton of some early osteostracans (Agnatha): preliminary analysis of its significance OLGA B. AFANASSIEVA Olga B. Afanassieva, Paleontological Institute of Russian Academy of Sciences, 123, Profsoyuznaya St., Moscow 117997, Russia; oafan@paleo.ru The surface of the osteostracan exoskeleton has been studied using the SEM on isolated microremains, and small fragments taken from complete cephalothoracic shields. The material comes from the Silurian and Lower Devonian deposits of Severnaya Zemlya Archipelago, Russia, and Saaremaa Island, Estonia. Imprints of epidermal cells on the exoskeleton surface are described for the first time in osteostracans. It is concluded that the sculpture on the osteostracan exoskeleton, both macrosculpture and microsculpture, reflects processes of the probable mode of ossification of the osteostracan hard cover. On the other hand, various types of microsculpture (microtubercles, fine ribs or stripes, microapertures) in general are related to the functional peculiarities responsible for animals’ adaptation to the ambient environment, and were necessary for the implementation of metabolic processes in different covering tissues of early vertebrates. Key words: Palaeozoic agnathans, osteostracans, exoskeleton, surface sculpture. Introduction In the last few decades considerable progress has been made in the study of the dermal skeleton of Palaeozoic vertebrates. Use of the scanning electron microscope (SEM) produced new interesting data on the exoskeleton microstructure of different groups, including the fine sculpture of the exoskeleton surface (Smith 1977; Schultze 1977; Deryck and Chancogne-Weber 1995; Märss 2002; Beznosov 2003; also see references in Märss 2002). For instance, fine sculptural elements, about ten microns in diameter, were found on the exoskeleton surface in different groups (in chondrichthyans, acanthodians, and dipnoans) and were explained as imprints of the epidermal cells of integument. However, for osteostracans little is known about the exoskeleton microrelief, and there are no special papers on the subject. The present paper attempts to analyze some of the relevant data for the osteostracan exoskeleton. Material and methods Isolated microremains of exoskeleton, extracted by V.N. Karatajute-Talimaa from rock samples by dissolving with formic acid, were determined as Oeselaspis pustulata (Patten), Tremataspis obruchevi Afanassieva et Karatajute-Talimaa, Tremataspis cf. schmidti, T. cf. milleri, and Tremataspis sp. (Afanassieva and Märss 1999; Afanassieva 2000). The material comes from the Ust’-Spokojnaya Formation, Ludlow, Upper Silurian of October Revolution Island, Severnaya Zemlya Archipelago, Russia.

Olga Afanassieva. Microrelief on the exoskeleton of early osteostracans 15 Ungulaspis arctoa, described by Afanassieva and Karatajute-Talimaa (1998), occurs in the upper part of the Severnaya Zemlya Formation deposits of the Lower Devonian of October Revolution Island. Exoskeleton microfragments belonging to the holotype of Ungulaspis arctoa were taken from the anterolateral and marginal parts of the dorsal shield. All fragments were studied by the present author using the SEM technique. Complete and fragmented shields of osteostracans from the collections of the Paleontological Institute of the Russian Academy of Sciences in Moscow (mainly, Tremataspidoidei from Silurian of Saaremaa Island) were used for comparison. Material described in this paper is housed in the Paleontological Institute of the RAS (PIN) and in the Institute of Geology and Geography of Lithuania in Vilnius (LIGG). Description and discussion The surface of the cephalothoracic shields of Tremataspis species is smooth and shiny (Robertson 1938; Denison 1951a; Afanassieva and Karatajute-Talimaa 1998). There are only a few, low tubercles on the dorsal side of the interzonal part of the shield. The tubercles have smooth, non-perforated walls and tops. The sensory-line system opens on the surface through the relatively wide pores. The diameter of the pores varies from 10-15 microns (T. milleri Patten) up to 40 microns (T. obruchevi Afanassieva et Karatajute- Talimaa). The surface of trunk scales of Tremataspis is usually shiny and smooth (Fig. 1 A). The scales are thick, with all three layers of exoskeleton well developed. The basal layer has a typical cross-laminated structure and forms a significant part of the exoskeleton. No fragments and scales with tubercles were found. Some scales demonstrate an infrequent type of fine sculpture. On the surface of the scale (specimen PIN 4765/20) irregular polygons reminding a honeycomb pattern are clearly visible (Fig. 1 B, C). Their diameter is about 10 microns. Similar ultrasculpture has been found on the surface of cephalothoracic shields, and of several scales of Tremataspis species from the Silurian deposits of Saaremaa Island, Estonia (Fig. 1 D). The surface of the cephalic shield of Ungulaspis arctoa in the anteromedial parts of the dorsal side are covered with small rounded and elongated tubercles (Afanassieva and Karatajute-Talimaa 1998; Afanassieva 1999, pl. 1, fig. 1). The tubercle sides are composed of relatively compact tissue without any foramina. The surface of the tubercles is covered with fine ribs which form a peculiar microrelief (Fig. 1 E, F). The distance between the ribs is about 5 microns. The ribs usually meet at the top of the rounded tubercle and at the apical part (ridge) of the elongated one. Similar ribbing is found on the side surfaces of the large (about 1 mm in length) elongated tubercles situated along the anterior and lateral edges of the cephalic shield of Ungulaspis arctoa (Afanassieva 1999, pl. 1, fig. 4). To clarify the significance of the thin ribbing of the osteostracan exoskeleton we should focus our attention on the external skeleton of Thyestes verrucosus Eichwald. The surface of the Thyestes shield is covered with numerous tubercles (Stensiö 1932; Denison 1951b) that can be divided into three types: large tubercles with curved tips,

Olga Afanassieva. Microrelief on the exoskeleton of early osteostracans<br />

15<br />

Ungulaspis arctoa, described by Afanassieva <strong>and</strong> Karatajute-Talimaa (1998), occurs<br />

in the upper part of the Severnaya Zemlya Formation deposits of the Lower Devonian<br />

of October Revolution Isl<strong>and</strong>. Exoskeleton microfragments belonging to the holotype<br />

of Ungulaspis arctoa were taken from the anterolateral <strong>and</strong> marginal parts of the dorsal<br />

shield.<br />

All fragments were studied by the present author using the SEM technique. Complete<br />

<strong>and</strong> fragmented shields of osteostracans from the collections of the Paleontological<br />

Institute of the Russian Academy of <strong>Sciences</strong> in Moscow (mainly, Tremataspidoidei<br />

from Silurian of Saaremaa Isl<strong>and</strong>) were used for comparison.<br />

Material described in this paper is housed in the Paleontological Institute of the<br />

RAS (PIN) <strong>and</strong> in the Institute of Geology <strong>and</strong> Geography of Lithuania in Vilnius<br />

(LIGG).<br />

Description <strong>and</strong> discussion<br />

The surface of the cephalothoracic shields of Tremataspis species is smooth <strong>and</strong> shiny<br />

(Robertson 1938; Denison 1951a; Afanassieva <strong>and</strong> Karatajute-Talimaa 1998). There are<br />

only a few, low tubercles on the dorsal side of the interzonal part of the shield. The<br />

tubercles have smooth, non-perforated walls <strong>and</strong> tops. The sensory-line system opens<br />

on the surface through the relatively wide pores. The diameter of the pores varies from<br />

10-15 microns (T. milleri Patten) up to 40 microns (T. obruchevi Afanassieva et Karatajute-<br />

Talimaa).<br />

The surface of tr<strong>un</strong>k scales of Tremataspis is usually shiny <strong>and</strong> smooth (Fig. 1 A).<br />

The scales are thick, with all three layers of exoskeleton well developed. The basal layer<br />

has a typical cross-laminated structure <strong>and</strong> forms a significant part of the exoskeleton.<br />

No fragments <strong>and</strong> scales with tubercles were fo<strong>un</strong>d. Some scales demonstrate an<br />

infrequent type of fine sculpture. On the surface of the scale (specimen PIN 4765/20)<br />

irregular polygons reminding a honeycomb pattern are clearly visible (Fig. 1 B, C). Their<br />

diameter is about 10 microns. Similar ultrasculpture has been fo<strong>un</strong>d on the surface of<br />

cephalothoracic shields, <strong>and</strong> of several scales of Tremataspis species from the Silurian<br />

deposits of Saaremaa Isl<strong>and</strong>, Estonia (Fig. 1 D).<br />

The surface of the cephalic shield of Ungulaspis arctoa in the anteromedial parts of<br />

the dorsal side are covered with small ro<strong>un</strong>ded <strong>and</strong> elongated tubercles (Afanassieva<br />

<strong>and</strong> Karatajute-Talimaa 1998; Afanassieva 1999, pl. 1, fig. 1). The tubercle sides are<br />

composed of relatively compact tissue without any foramina. The surface of the tubercles<br />

is covered with fine ribs which form a peculiar microrelief (Fig. 1 E, F). The distance<br />

between the ribs is about 5 microns. The ribs usually meet at the top of the ro<strong>un</strong>ded<br />

tubercle <strong>and</strong> at the apical part (ridge) of the elongated one. Similar ribbing is fo<strong>un</strong>d on<br />

the side surfaces of the large (about 1 mm in length) elongated tubercles situated along<br />

the anterior <strong>and</strong> lateral edges of the cephalic shield of Ungulaspis arctoa (Afanassieva<br />

1999, pl. 1, fig. 4).<br />

To clarify the significance of the thin ribbing of the osteostracan exoskeleton we<br />

should focus our attention on the external skeleton of Thyestes verrucosus Eichwald.<br />

The surface of the Thyestes shield is covered with numerous tubercles (Stensiö 1932;<br />

Denison 1951b) that can be divided into three types: large tubercles with curved tips,

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