A taxonomic revision of the tertiary echinoid genus monostychia

Abstract

For over 100 years the genus Monostychia (Echinoidea: Clypeasteroida) and its type species M. australis Laube, 1869 have been a taxonomic home for a wide range of genera and species with the commonality of a rounded to pentagonal, discoidal test and a submarginal periproct. The specimens comprising this group are all extinct and from the Tertiary strata of southern Australia. While there have been a few minor species identified beyond M. australis, notably M. etheridgei Woods, 1877 and P. loveni (Duncan, 1877), it has been clear to many researchers that the variability remaining in M. australis was representative of numerous other taxa awaiting discovery. Recent taxonomic works on the Clypeasteroida suggested that the number of interambulacral plates on the oral surface of the test of some species was a useful diagnostic character. Of interest were the plates that first come into contact with the periproct. However, there appeared little evidence in the literature that it had been established that the number of such plates remained constant with test length and age, or that the variability in each taxon, of those plate numbers, has been determined. Without understanding those two issues the utility of plate numbers was questionable. This study set out to resolve some of those issues for Monostychia and its relatives. It was found that the number of interambulacral and ambulacral plates on the oral surface was fixed and did not change with increasing test length and therefore there was potential utility for plate numbers as a taxonomic tool. However, there was substantial variability in the numbers. As a result, the use of plate numbers in the paired interambulacra, paired ambulacra, and ambulacrum III on the oral surface appears to have limited utility at genus level. At the species level, however, such numbers can be quite useful diagnostically, particularly when paired interambulacral, paired ambulacral and ambulacrum III plate numbers are used in combination. The plates that first come into contact with the periproct was shown to have little value taxonomically at the genus or species level within the monostychioids, largely because most species had the same plate number dominating, but also because of the variability. At subfamily level the taxonomic value of this feature is yet to be established. A previously unreported structural feature was identified in many of the specimens. This was a thin circumferential wall of stereom present on the right-hand side of the test, lying half way between the marginal and central buttressing. It was a form of intestinal buttressing referred to hereafter as the intermurum. Its presence enabled the establishment of a new subfamily, Monostychinae in the family Arachnoididae. Four genera have been placed in the Monostychinae; Monostychia Laube, 1869, Quinquestychia gen. nov., Rotundastychia gen. nov. and Deltoidstychia gen. nov. A key to these genera is provided. Several species have been established and others redescribed in this study. In Monostychia there are seven species; M. australis Laube, 1869, M. macnamarai Sadler et al. 2017, M. alanrixi Sadler et al. 2017, M. merrimanensis Sadler et al. 2019, M. etheridgei Woods, 1877, M. glenelgensis Sadler et al. 2019 and M. robheathi sp. nov. A new genus, Quinquestychia gen.nov., has also been established. It contains four species: Q. mannumensis sp. nov., Q. gigas sp. nov., Q. berylmorrisae sp. nov. and Q. robertirwini (Sadler et al. 2017). The last of these species was published as a Monostychia earlier in this study but reassigned later on the basis of further data. A second new genus, Rotundastychia gen. nov., has also been established. It contains three species: R. pledgei sp. nov., R. aquilaensis sp. nov. and R. eyriei sp. nov. A third new genus, Deltoidstychia gen. nov., has also been established. It currently contains a single species, D. erioaster sp. nov. In addition to the above, two other new genera were established but they do not belong in the subfamily Monostychinae. Instead they are tentatively placed in the subfamily Ammotrophinae. The first of these is Obscurostychia gen. nov. with two new species: O. spirographica sp. nov. and O. curtus sp. nov. Keys to all the genera discussed above that contain more than one species have been provided.