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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.
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.
Three isolated and well−preserved retiolitid taxa, two of them new, and one taxon of unknown affinity, are described from the upper Llandovery of Arctic Canada. All taxa display unusual characteristics The three retiolitids display extraordinary morphological features: Pseudoplegmatograptus cf. obesus preserves very delicate and lacey or cobweb−like list structures external to the main skeletal lists, structures previously seen only rarely on flattened material; Pileograptus pileatus gen. et sp. nov. possesses a thecal framework typical of retiolitines, particularly Stomatograptus, but its thecal orifices possess broad, meshwork genicular hoods similar to those in the plectograptines; and Giganteograptus giganteus, formerly attributed to Pseudoplegmatograptus, with its very coarse and relatively simple skeletal framework, well−developed paired thecal apertural spines, and a pustulose micro−ornamentation on the lists. The latter feature suggests an assignment to the subfamily Plectograptinae, rather than to the Retiolitinae as has been previously presumed. The presence of pustulose lists in Giganteograptus adds a further complication to understanding of retiolitid evolution. Mirorgraptus arcticus gen. et sp. nov., the fourth taxon and of unknown systematic affinity, lacks the proximal end, but preserves lists with seams and some fusellar material suggestive of some type of new retiolitid. Unlike known retiolitids or other graptolites, however, the skeletal list development is apparently confined to a narrow region along one side of the nema.
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Convergent evolution of two Silurian graptolites

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The two graptolite speciesCochlograptus veles(Telychian, Upper Llandovery) and Testograptus testis(lower Homerian, upper Wenlock) are remarkably and uniquely similar in being strongly ventrally and planispirally coiled and in demonstrating an abrupt deflection in the immediate post−sicular regions of their rhabdosomes. The two species, however, are separated by a relatively large biostratigraphic gap and a global mass extinction, and they differ morphologically in the proportion of thecal overlap, different angles of inclination of the interthecal septa, relative proportions of the widths occupied by the free metathecae, the position of the sharp dorsal flexure relative to the tip of the sicula, and the presence of a distinctive, keel−like structure on T. testis. It is suggested therefore, that in spite of the strong proximal morphological parallelism between the two species, their origin is best explained as a remarkable example of convergent evolution. It is suggested that C. veles perhaps evolved from some modified monograptid such as Stimulograptus, whereas the small Testograptus group may have derived from some monograptid such as Monograptus flemingii. Cladistic analysis fully supports the independent derivation the two species. If correct, this hypothesis supports the validity of separate generic names for the two species, despite the close and unique rhabdosomal similarities, including proximal metathecal form.
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