New insights into the phylogeny of Rhuddanian (Lower Llandovery) graptolites  

Michael J. Melchin1, Aleksandra Naczk–Cameron2 and Tatiana N. Koren3  

1 Department of Earth Sciences, St. Francis Xavier University, Antigonish, Nova Scotia, B2G 2W5, Canada.  E–mail:

2 Canadian Natural Resources Limited, 2500, 855 – 2 Street S.W., Calgary, AB T2P 4J8.

3 All-Russia Geological Research Institute (VSEGEI), Sredny Pr. 74, 199026, St. Petersburg, Russia.  E–mail:


Keywords: Graptolites. Astogeny. Phylogeny. Llandovery. Silurian.

Study of the thecal form, proximal development, and internal structure of some late Hirnantian and Rhuddanian biserial graptolites has led to some new insights into the phylogenetic relationships among the major groups of Silurian diplograptids. Cladistic analyses suggest that Glyptograptus s.s. with a Pattern I (tamariscus) proximal development form a sister group with Pattern I species of Sudburigraptus. These taxa are probably ancestral to the later Llandovery petalolithids. The earliest known Pattern I taxa appear to belong to a group of species related to Paraclimacograptus? kiliani, which first appear at the beginning of the Rhuddanian. Preliminary evidence suggest that these latter forms were ancestral to all subsequent petalolithid (including Glyptograptus) and retiolitid taxa.

Species of Rivagraptus and possibly Agetograptus appear to form a distinct clade among Silurian diplograptids, characterized by a very short sicula, no descending portion of theca 11 and distinctive ridges marking the margins of the fuselli and cortical bandages. These taxa appear to be derived from a mid–Rhuddanian "normalograptid" with similar fuselli and bandages and a short sicula, but also a very short descending th11.

Preliminary evidence also indicates that the akidograptids, dimorphograptids, and early monograptids appear to share a common ancestry, from a nomalograptid ancestor like Normalograptus avitus at the beginning of the Rhuddanian. This group appears to form a clade distinct from other Rhuddanian taxa.

At the present time, the relationships among the remaining groups of Rhuddanian normalograptines remain to be studied.

It is recognized that biserial species with a full median septum clearly show two separately growing series of thecae (two stipes) and that those that are aseptate normally have a single series of thecae (one stipe) alternately arranged in two "series" along the rhabdosome. However, many taxa that possess a partial median septum throughout the length of the rhabdosome also appear to have only a single series of alternately arranged thecae. In addition, at least one species, Normalograptus indivisus, appears to possess two stipes but no preserved median septum. Thus, the terms "septate" and "aseptate" do not adequately characterize the range of variation of biserial taxa with one vs. two stipes. In addition, the terms "biserial" and "uniserial" are firmly entrenched as referring to rhabodosomes with theca with two sides and one side of the rhabdosome, respectively. We, therefore, proposed that the term "bistipular" be used to refer to any taxa with two separately growing stipes (i.e., those with a dicalycal theca in the proximal end) and "unistipular" to taxa with a single stipe (i.e., no dicalyc). Thus, monograptids are both uniserial and unistipular, septate diplograptids are biserial and bistipular, but most aseptate diplograptids and some taxa with a partial median septum are biserial and unistipular.



Received: February 15, 2003

Accepted: June 15, 200