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Volume 81, 1953
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1. The Generic Content of the Tertiary Echinoderm Faunas of Australasia

(a) Early Tertiary.

Our knowledge of the Australasian echinoid faunas of the Cretaceous and Eocene is very fragmentary. In New Zealand occurred the Cretaceous Micraster,

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and the Senonian starfish Ophryaster is represented by a species which closely recalls those of the Danish and English Senonian. From Western Australia a Cretaceous cidarid is known, but its generic position is uncertain.

During the Eocene the holasterid echinoid Cardiaster was present in both Australia and New Zealand, though it had by then already become extinct at the close of the Cretaceous in all other areas, so far as we know. Nucleopygus (syn. Nucleolites), another Mesozoic genus, was established in Australasia; we know it with certainty from Australia, and it may well yet turn up in the New Zealand Eocene. Elsewhere it was in process of extinction. Its lineage was destined to follow a long history in Australia and-New Zealand, with Recent representatives in both areas (Apatopygus). In Australian Eocene sediments are also found Salenidia and Prenaster. In New Zealand are found Schizaster, Galeraster, Loveniidae of undetermined generic position (probably representing an undescribed genus), and some Echinometrid, possibly Heliocidaris. Brochopleurus, listed as an Oligocene immigrant below, has also been found in the Upper Eocene (Kaiatan) since this paper was set in type.

(b) Mid Tertiary.

Mesozoic holasterids continued to survive in New Zealand at the opening of the Oligocene, with Cardiaster and also Echinocorys, the latter genus not yet known earlier in New Zealand sediments. The two genera, however, do not seem to have survived into the mid-Oligocene. Another, and later, development of the holasterid stock is seen in Duncaniaster, which appears in the mid-Oligocene, and enters upon a prolonged history in the Australasian region.

Genera which first appear in the lower or middle Oligocene of New Zealand include a number of forms of large size. The assemblage includes the following, together with a few others not yet determined:-Histocidaris, Phyllacanthus, Stereocidaris, Goniocidaris, Eucidaris, Prionocidaris, Grammechinus, Brochopleurus (see Eocene), Echinocorys, Duncaniaster, Fibularia, Studeria, Echinolampas, Planilampas, Maretia, Lovenia, Pericosmus, Linthia, Brissopsis, Cyclaster, Eupatagus, Brissus. Also till such time as Nucleopygus is known with certainty from the New Zealand Eocene, it must be included here.

Contemporary Australian lower and middle Oligocene beds of marine origin are believed to be unrepresented stratigraphically. They may well have carried a similar faunal assemblage to those of New Zealand, since we find much the same kind of fauna in the Janjukian, which Finlay (1947) attributes to upper Oligocene plus lower Miocene. The Janjukian yields the following genera:-Phyllacanthus, Stereocidaris, Goniocidaris, Eucidaris (to which the unnamed club-shaped spines of Chapman and Cudmore (1934) must certainly be ascribed), Prionocidaris, Paradoxechinus, Brochopleurus, Duncaniaster, Fibularia, Studeria, Echinolampas, Plesiolampas, Australanthus, Cassidulus, Echinoneus, Maretia, Lovenia, Pericosmus, Linthia, Schizaster, Brissopsis, Cyclaster, Eupatagus, Scutellina, Sismondia, Conoclypus, Monostichia, Coelopleurus, Gualtieria and Hemiaster; as well as Nucleopygus which was earlier present.

New Zealand Miocene marine sediments are mainly of a non-calcareous and sandy facies, yielding little direct information on the echinoid faunas. We do know that Schizaster and Pericosmus persisted, the latter at least into the lower Miocene, the former till near the close of the period. It would seem probable that, where the environment was suitable, the Oligocene genera would have per-

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sisted—in other words, the fauna would have resembled that known from the Australian Janjukian.

It will be seen that the two portions of the mid-Tertiary represented in Australia and New Zealand by echinoderm-bearing beds contain a similar assemblage of genera. The similarities, indeed, extend to the specific level, as in the case of Brochopleurus, which has similar or identical species on either side of the Tasman (Fell, 1949a), and Duncaniaster (unpublished evidence).

(c) Late Tertiary.

Profound changes in the Australasian echinoid faunas towards the close of the Miocene parallel those observed in the mollusca and other groups. These latter have been attributed to a cooling climate (Fleming, 1949), and the same conclusion might be drawn from the echinoid evidence. Thus, the disappearance of Schizaster, a supposedly warm-water genus, takes place in New Zealand at the close of the Tongaporutuan stage (Upper Miocene). The warm-water genus Phyllacanthus last occurs in New Zealand in the Kapitean stage (Uppermost Miocene).

Genera which disappeared from south-eastern Australia include Duncaniaster, Lovenia, Linthia, Hemiaster, Gualtieria, Coelopleurus, Brochopleurus, Paradoxechinus, Monostichia, Sismondia, Scutellina, Nucleopygus, Cassidulus, Australanthus, Studeria, Plesiolampas and Echinolampas.

The New Zealand pre-Pliocene extinctions removed the following genera from the fauna:-Schizaster, Duncaniaster, Lovenia, Linthia, Brochopleurus, Studeria, Planilampas, Echinolampas, Grammechinus, Fibularia, Maretia, Pericosmus, Eupatagus, and all the cidarids, except apparently Stereocidaris, and perhaps Goniocidaris.

The following Miocene genera of Australia are still represented in the Recent East Australian Fauna, though not known from the Pliocene; they may have survived in warmer waters of the northern or north-eastern coasts:-Prionocidaris, Eucidaris, Fibularia, Echinoneus, Maretia, Lovenia, Pericosmus, Schizaster, Brissopsis, Eupatagus; and also the Nucleopygus-Apatopygus lineage, known from the Recent of Western Australia.

These losses were partly balanced by an accretion of new genera of modern aspect. Clypeaster, Arachnoides, Peronella and probably also Pseudechinus appeared in the Australian Lower Pliocene, whilst in the corresponding sediments of New Zealand (Waitotaran and Nukumaruan) we first meet with Arachnoides, Pseudechinus, Echinocardium and Evechinus.

The Recent echinoderm faunas of the two regions need not be summarized here; see, for Australia, Clark (1946), and for New Zealand, Fell (1949b).