Atavograptus primitivus (Li) from the earliest Silurian of Arctic Canada: implications for monograptid evolution

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Abstract
It has long been considered that all monograptids have theca 1 originating from a primary porus in the sicula (i.e., sinus and lacuna stages; Bulman, 1970). The major basis for this assumption has been studies of Late Silurian monograptid species such as Saetograptus chimaera (Barrande), Pristiograptus bohemicus (Barrande), and Pristiograptus dubius (Suess) (e.g., Eisenack, 1942; Walker, 1953; Urbanek, 1958). That all monograptids possessed this type of development was not questioned due to the lack of well-preserved, early monograptid specimens clearly showing the sicula–theca 1 relationship. Hutt et al. (1970) described Monograptus gregarius Lapworth, from the gregarius Zone, as having theca 1 originating through a primary porus (Hutt et al., 1970; Rickards and Hutt, 1970, referred to this as a “primary notch” although, according to Bulman, 1970, p. V11, the notch corresponds only to the sinus stage). Their figures of these specimens, however, do not unequivocally confirm its presence (Hutt et al., 1970, p. 14, Pl. 3, figs. 65, 67). Rickards and Hutt (1970) alluded to the possible presence of a primary porus in Atavograptus ceryx (Rickards and Hutt) from the persculptus Zone, but could not determine its existence with certainty. Mitchell (1987) was the first to question the presence of this mode of development of theca 1 in relation to the origin of the monograptids. He proposed, however, that the development of the fully uniserial rhabdosome was coincident with a change in both the sicular structure (the sinus and lacuna stages) and the early ontogeny of theca 1. He suggested that the sinus and lacuna stages represented a homologue of the hooded foramen seen in the th 11 protheca of some Pattern H species, from which th 12 emerges and that, in the uniserial monograptids, this feature is accelerated in the astogenetic sequence, appearing in the metasicular ontogeny rather than that of the first theca. He pointed out that this hypothesis could only be tested once suitable, isolated specimens of earliest monograptids could be studied to see if the sinus and lacuna stages were, indeed, coincident with the achievement of a fully uniserial rhabdosome. New evidence from isolated, early monograptid material allows us to test Mitchell's theory on monograptid origin.

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