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The new retiolitid species, Spinograptus tubothecalis, is described from the Colonograptus praedeubeli and C. deubeli biozones from two localities in Poland: a borehole on the East European Platform and the Holy Cross Mountains. This was a recovery phase after the severe Silurian Cyrtograptus lundgreni Event. The new species has a unique, previously undescribed form of finite rhabdosome. Unlike the species Spinograptus reticulolawsoni and S. lawsoni, in which the fi− nite rhabdosomes taper distally, its rhabdosome is parallel−sided with the two distal thecae developed as isolated tubes without genicular processes, with a small appendix between them. The new species also has preserved membranes of the sicula, thecae and ancora sleeve, similar to a few species of Spinograptus from the lower Homerian. Spinograptus tubothecalis, like Spinograptus clathrospinosus and S. spinosus, has paired reticulofusellar genicular processes on the pre−thecal ventral orifices, similar to but shorter than thecal processes. Transverse rods, a rare character in post−Cyrto− graptus lundgreni Event retiolitids occur in the new species in rudimentary form.
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Evolution of retiolitid graptolites - a synopsis

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Twenty million years of retiolitid evolution reflect environmental changes, the most severe being the Silurian Cyrtograptus lundgreni Event. Five biostratigraphically and morphologically constrained retiolitid faunas are distinguished and characterized according to their rhabdosomal modifications: (1) the oldest and long−ranging Llandovery group of mostly large and morphologically complex rhabdosomes, (2) the less diverse Telychian−Sheinwoodian group, (3) the Cyrtograptus lundgreni Biozone varied group of intermediate size, and two short−ranged (4) late Homerian, and (5) early Ludlow groups with small rhabdosomes. Although the evolutionary history of retiolitids was complex and not linear, a common tendency toward reduction of rhabdosome size in most lineages is observed. The greatest reduction in both number and volume of thecae, and in skeletal elements is demonstrated in the Gothograptus and Plectograptusfaunas. Contrary to the thecal decrease, a distinctive increase of sicula size is observed in retiolitids. Two types of colonies are distinguished: L−colonies with a small sicula and numerous large thecae of similar size, and S−colonies with a long sicula and a few, small thecae. These changes imply modification of the soft body: an increase in siculozooid length and a decrease in the size of the zooids. Thus, the siculozooid probably produced great amounts of morphogen inhibiting zooid growth. In consequence the phenomenon of colony reduction occurred. The most extreme stages of rhabdosome reduction in Ludlow retiolitids can be seen in Plectodinemagraptus gracilis of the Plectograptuslineage and in the new species Holoretiolites helenaewitoldi, possibly representing the last stage of skeletal reduction in the Gothograptus lineage; the next hypothetical stage would be its total loss. The sicula length of Holoretiolites, about 2 mm, is reported herein for the first time.
Agastograptus robustus Obut and Zaslavskaya, 1983, the type species of the genus Agastograptusis herein recognized as a species of Plectograptus. Large proximal lateral orifices are one of the significant characters of Plectograptus. P. robustus differs from the type species, Plectograptus macilentus Törnquist, 1887, in possessing paired apertural processes. Other species of Agastograptus have been assigned to three different genera: Spinograptus, S. clathrospinosus (Eisenack, 1951), S. munchi (Eisenack, 1951), Neogothograptus, N. balticus (Eisenack, 1951), and Cometograptus, C. nevadensis (Berry and Murphy, 1975). The main diagnostic feature used for Agastograptus, the spinoreticular paired apertural processes, is recognized as a species feature, characteristic mostly for Spinograptus, whereas the generic features are the arrangement of the proximal end, ventral walls, and ancora sleeve of the rhabdosome. Therefore the genus Agastograptus is a synonym of Plectograptus.
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Silurian retiolitids of the East European Platform

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Late Llandovery to late Ludlow isolated retiolitid assemblages from deep boreholes drilled in the Polish part of the East European Platform are used to restore phylogeny of the group. The process of reduction of the network skeleton (reticulum and clathria), as well as reduction of the transverse rods and change of virgula position started in the Wenlock. All Ludlow retiolitids have a central virgula and reduced rhabdosomes. Colonies of the Gothograptus lineage with finite growth, related to reduction in the number of thecae, first appeared in the Late Wenlock and continued in the Ludlow. Their evolution leads either to development of finite rhabdosomes with few thecae (Holoretiolites) or to almost complete reduction of clathria and reticulum (Plectodinemagraptus gen. n. of the Plectograptus lineage). The latter homeomorph of the Ordovician abrograptid Dinemagraptus is the last known retiolitid of late Ludlow age (Cucullograptus hemiaversus /C. aversus Zone). The family Retiolitidae consists of five separate lineages. Sokolovograptus polonicus sp. n., Neogothograptus gen. n., Holoretiolites atrabecularis sp. n., Semiplectograptus urbaneki gen. et sp. n., Plectodinemagraptus gracilis gen. et sp. n. are proposed. S. polonicus from the Cyrtograptus murchisoni Zone has a strongly reduced rhabdosome.
The Ludlow genus Plectograptus, with the type species Retiolites macilentus Törnquist, 1887, collected from Thuringia (Germany), has been a widely−identified, monospecific, but poorly understood taxon for almost one hundred years. This was due to poor and incomplete preservation of the type material, and misidentification by subsequent authors up to 1995. The original, and only, type specimen of P. macilentus collected by Törnquist being lost, a neotype is herein selected from a small collection of Thuringian material. The genus has now been redefined and based on this, and SEM studies of isolated material, the defining characteristics of the genus are (i) the possession of a simple ancora umbrella with five radial lists with an incompletely developed rim; (ii) an ancora umbrella separated from lateral ancora sleeve walls by exceptionally large lateral orifices; (iii) the possession of mid−ventral lists; (iv) simple, orderly zigzag lateral wall ancora sleeve lists. Recently, two additional species, P. robustus and P. wimani, previously placed in different genera, were assigned to Plectograptus. This study recognizes three new species: P. mobergi, P. toernquisti, and P. trijunctus, bringing the total number of species to six. Species are distinguished by the presence or absence of genicular processes, inclination of the thecal ventral walls and mid−ventral lists, presence or absence of reticular lists, and three−way or four−way sleeve/lateral rod/apertural lip junctions.
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The new retiolitid graptolite Kirkigraptus inexpectans gen. et sp. nov., from the Neodiversograptus nilssoni Biozone of the Bartoszyce borehole, Poland is described. It is unique among the retiolitids not having a preserved virgella or ancora. Instead the most proximal structures are two round proxi−lateral lists, joining the two genicular lists of the first thecae, connecting the two sides of the rhabdosome. The lists are interpreted as a possible homologue of the distal edge of the ancora umbrella in typical retiolitids. The size of rhabdosome with large proximal lateral orifices, and the ventral panels of thecae with mid−ventral lists, are similar to those of Plectograptus, whereas the two ancora sleeve panels consisting of spaced horizontal lists only, resemble those of Valentinagraptus. It is possible that the new retiolitid may represent a new pattern of development of the proximal end of the rhabdosome, different from that in all other retiolitids.
Neogothograptus reticulatus sp. nov. from the upper Homerian Colonograptus praedeubeli Biozone, and N. thorsteinssoni and N. alatiformis from the Lobograptus progenitor Biozone, are described for the first time from three localities: borehole, Baltic erratic boulder of East European Platform and Holy Cross Mountains of Poland. N. reticulatus, presently the oldest known species of Neogothograptus, is also tentatively identified from upper Homerian strata of southeastern Australia. The two other species are previously known only from Arctic Canada, and possibly China. The morphology of the Neogothograptus reticulatus rhabdosome, its appendix, thecal profile, densely reticulated rhabdosome and genicular hoods suggest a close relationship to N. eximinassa from Colonograptus ludensis Biozone. N. reticulatus and N. eximinassa are most similar to Gothograptus nassa, the earliest−known retiolitid to appear immediately following the Cyrtograptus lundgreni extinction event. The biostratigraphic position of N. reticulatus suggests it might be considered as a potential ancestor to all younger (Ludlow) species of Neogothograptus. Cladistic analysis, however, provides no direct support for such an interpretation and, instead, suggests that Baculograptus batesi may be the ancestor. The occurrences of Neogothograptus, as well as G. nassa, from a number of Silurian terranes mostly from low paleolatitude regions, but also from high paleolatitudes, demonstrate their tolerance to a broad range of paleoenvironments.
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