Estonian Journal of Earth Sciences, 2017, 66, 2, 59–76 https://doi.org/10.3176/earth.2017.05
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A new psammosteid (Agnatha, Heterostraci) from the Amata Regional
Stage of the Main Devonian Field and morpho-histological types of
discrete micromeric elements in the family Psammosteidae
In memory of an outstanding palaeoichthyologist Elga Mark-Kurik
Vadim N. Glinskiy and Sergey P. Nilov
Saint Petersburg State University, 7/9 Universitetskaya nab., St. Petersburg, 199034 Russia; vadim.glinskiy@gmail.com
Received 28 October 2016, accepted 8 February 2017
Abstract. The range of diversity of psammosteids from the family Psammosteidae is still poorly known. Here a new species,
Psammosteus ramosus sp. nov. Glinskiy, from the Amata Regional Stage of the Main Devonian Field is described. Its
morphology, ornamentation, histology of exoskeletal plates, and micromeric elements are compared with those of other
representatives of the family Psammosteidae. The comparison shows a close relationship of the new species with Psammosteus
falcatus Obruchev, P. kiaeri Halstead Tarlo and P. pectinatus Obruchev, a group of species that is significantly different from other
representatives of the genus Psammosteus and constitutes a separate evolutionary lineage. On the basis of morphological and
histological features we here differentiate in the fields of tesserae of Psammosteidae the discrete micromeric elements of the
‘basic type’, known in Psammosteus bergi (Obruchev), P. levis Obruchev, P. livonicus Obruchev, P. maeandrinus Agassiz,
P. megalopteryx (Trautschold), P. praecursor Obruchev and Karelosteus weberi Obruchev, and micromeric elements of the
‘progressive type’, known in Psammosteus falcatus, P. cf. kiaeri and P. ramosus sp. nov. Glinskiy.
Key words: Agnatha, Heterostraci, Psammosteida, Late Devonian, Main Devonian Field, new species.
INTRODUCTION
Psammosteids are a group of jawless vertebrates from
the order Pteraspidiformes (suborder Psammosteida sensu
Tarlo 1962). They are known from the Early–Late
Devonian (from Pragian to Frasnian stages), primarily
of Laurussia (Halstead Tarlo 1967a; Halstead 1987)
and, to a lesser extent, of the peri-Gondwanan shelf,
Armorica, Barentsian, Kara-Tajmyr and Siberian palaeo-
continents (Bystrow 1959; Blieck et al. 2002; Delsate et
al. 2004; Vaškaninová & Kraft 2016). The exoskeleton
of psammosteids is composed of macromeric and
mesomeric elements (plates of the cephalothorax, scales
in the tail region) and micromeric elements (mostly
tesserae). Discrete micromeric elements, situated between
the main plates of the cephalothorax, form so-called
‘fields of tesserae’. Praepineal and paired lateral fields
on the dorsal surface, and postoral and ventrolateral
fields on the ventral surface of the cephalothorax can
be distinguished (Obruchev & Mark-Kurik 1965, p. 40).
In numerous articulated specimens of Drepanaspis
gemuendenensis Schlüter, 1887 and in a single
articulated cephalothorax of Psammosteus megalopteryx
(Trautschold, 1880), fields of tesserae may be found
in their original articulation (Gross 1963; Halstead
Tarlo 1965, pl. XVI, figs 1, 2), but in most cases their
micromeric elements are found isolated (e.g. Mark-
Kurik 1999; Blom et al. 2006). As a rule, these elements
can be determined to the species level (Halstead Tarlo
1965; Obruchev & Mark-Kurik 1965; Glinskiy & Mark-
Kurik 2016) on the basis of species-specific tubercles.
Discrete micromeric elements in the family
Psammosteidae sensu Novitskaya (2004) are the most
diverse morphologically. They possess a smaller base
than other psammosteids. Rooted tesserae are also weakly
connected with the cancellous aspidin of the plates
(Gross 1933, p. 15, fig. 4; Obruchev 1947a, p. 197;
Mark-Kurik 1999, p. 7). The small base size of discrete
micromeric elements in representatives of Psammosteidae
results in their higher concentration in the fields of
tesserae and on the plates of the cephalothorax. Our
study of micromeric elements of psammosteids from
Frasnian deposits of the Main Devonian Field has
shown that discrete elements of the representatives of
the family Psammosteidae can generally be classified
into two morpho-histological types: the ‘basal’ and
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