Jennifer M. Bannister

New Zealand Eocene, Oligocene and Miocene Macrofossil and Pollen Records and Modern Plant Distributions in the Southern Hemisphere

The modern New Zealand flora has a relatively low number of families and genera in relation to land area, but well-preserved macrofossils and pollen from three sites in southern New Zealand suggest that the floras in Eocene, Oligocene and Miocene times were much more diverse at the generic level. At Pikopiko, Southland, a late Eocene in situ forest with fern understory was dominated by conifers, Casuarinaceae, Lauraceae, Nothofagus, Proteaceae, and mesothermal angiosperms including palms (aff. Calamus), Sapindaceae: Cupaniae and Picrodendraceae. At Newvale Mine, Southland, a leaf bed within a thick lignite seam represents leaf fossils preserved in a late Oligocene oligotrophic bog. This site demonstrates that Agathis, Dacrycarpus, Dacrydium, Halocarpus, Microcachrys, Podocarpus and Phyllocladus coexisted with diverse angiosperms including Nothofagus, Gymnostoma, Cunoniaceae, Ericaceae, Sapindaceae and several Proteaceae. Pollen data add Meliaceae, Myrtaceae, Onagraceae and Rubiaceae to the flora. At Foulden Maar, Otago, mummified leaves and flowers, including several with in situ pollen, demonstrate the existence of a diverse flora surrounding an Early Miocene lake. This site contains numerous monocot macrofossils including Astelia, Cordyline, Ripogonum and Typha, as well as the oldest fossils known for Orchidaceae and Luzuriagaceae. This flora was dominated by Lauraceae with affinities to Cryptocarya and Litsea, but other families include Araliaceae, Cunoniaceae, Elaeocarpaceae, Euphorbiaceae sensu lato, Menispermaceae, Myrsinaceae, Myrtaceae, Onagraceae, Proteaceae and Sterculiaceae. Many ferns, conifers, and Nothofagus are from lineages with Gondwanan ancestors, whereas other taxa show links to Australia (e.g., Gyrostemonaceae), New Caledonia (e.g., Beauprea) and South America (e.g., Luzuriaga, Fuchsia). Many of these taxa are now extinct in New Zealand, and therefore indicate much wider biogeographic ranges for many families and genera in the past.

Late Oligocene‐Early Miocene leaf macrofossils confirm a long history of Agathis in New Zealand

Abstract The antiquity of lineages in the extant indigenous flora of New Zealand is controversial, and plant macrofossils provide important evidence for testing hypotheses for in situ survival of ancient lineages or their geologically recent arrival by longdistance dispersal. Cuticle analysis of organically preserved leaf fossils confirms the presence of Agathis in New Zealand since at least the Late Oligocene to Early Miocene. Well‐preserved Agathis foliage occurs in a leaf litter bed within a thick seam of resiniferous lignite in the middle Gore Lignite Measures, Newvale Mine, Waimumu, Southland. The Agathis leaf fossils have some affinities with extant Agathis australis.

Morphology and palaeoenvironmental context of Fouldenia staminosa, a fossil flower with associated pollen from the Early Miocene of Otago, New Zealand

Abstract A new genus and species, Fouldenia staminosa, are proposed for an angiosperm flower with associated pollen bearing anthers found in Early Miocene finely laminated lacustrine diatomite at Foulden Hills, Otago. This is the first New Zealand record of a Miocene flower with in situ pollen and among few such records worldwide. The pollen is closely similar to Rubipollis oblatus, of unknown affinity, known from Late Oligocene to Early Miocene strata elsewhere in New Zealand and Australia, but differs in being spheroidal to subprolate. In spite of the relatively good preservation of both the flower and its pollen it has not been possible to assign this species to an extant genus or even family. The highly diverse angiosperm pollen flora from the same bedding plane in the diatomite indicates that Fouldenia grew in a dry, warm temperate to subtropical climate.

Fossil Ericaceae from New Zealand: deconstructing the use of fossil evidence in historical biogeography

The Australasian Ericaceae epitomize many problems in understanding the biogeography of the southern hemisphere, especially the relative contributions of Gondwanan vicariance and dispersal. Late Cretaceous fossil pollen of the family suggests extreme antiquity of the group in Australasia, but recent phylogenetic evidence suggests much younger histories for most of the groups in that region. This paper documents two new species of latest Oligocene-Early Miocene macrofossils of Ericaceae from New Zealand. Cyathodophyllum novae-zelandiae G.J.Jord. & Bannister gen. and sp. nov. is the oldest record of the tribe Styphelieae, but is of a clade now extinct in New Zealand, possibly related to the Tasmanian genus Cyathodes. Richeaphyllum waimumuensis G.J.Jord. & Bannister sp. nov. is a member of Richeeae, but it is ambiguous as to whether it is a member of the impressive modern New Zealand radiation in Dracophyllum. These fossils emphasize the fact that at least some of the fossil pollen of Ericaceae may have been derived from extinct lineages and therefore should not be used as evidence for the antiquity of any modern New Zealand clade of Ericaceae. New fossils and/or detailed analysis of fossil and extant pollen may help resolve such uncertainty.

Fossil scale insects (Hemiptera, Coccoidea, Diaspididae) in life position on an angiosperm leaf from an early Miocene lake deposit, Otago, New Zealand

Abstract The first fossil scale insects to be described from New Zealand are 14 well‐preserved female scale covers, including nine old, mature, adult females, attached to an angiosperm leaf found in Lower Miocene finely laminated lacustrine diatomite at Foulden Hills, Otago. These insects are interpreted as belonging to Family Diaspididae, Subfamily Aspidiotinae, a group that is well‐represented in the modern Australasian fauna. Their attachment in situ on a fossil leaf (possibly from the family Elaeocarpaceae) provides an extremely rare example of a 20‐million‐year‐old plant‐animal association, and demonstrates that before the Early Miocene Aspidiotinae had a neotenic, wingless female, and its modern form of scale cover, with a fully developed, fibrous, waxy scale cover.

High conifer diversity in Oligo-Miocene New Zealand

Eight species of Podocarpaceae foliage are recognised from the late Oligocene or early Miocene Newvale site, South Island, New Zealand, and the following five new species are described: two of Dacrydium Lamb. and one each of Dacrycarpus (Endl.) de Laub., Phyllocladus Rich. ex Mirb. and Halocarpus Quinn. The latter is the first macrofossil record of this New Zealand endemic genus. All these conifers, plus Agathis Salisb., Microcachrys Hook.f. and Podocarpus Pers., co-occurred in the local vegetation at Newvale. In conjunction with prior records of macrofossils and pollen, these fossils indicate that the late Paleogene to early Neogene conifer flora of New Zealand was very diverse, with all but one of the extant genera present, together with several regionally or globally extinct genera, and multiple species of many of these genera. This fossil diversity is similar to the Paleogene fossil conifer diversity observed in south-eastern Australia (and particularly Tasmania) and in parts of North America.

Subtropical rainforest vegetation from Cosy Dell, Southland: plant fossil evidence for Late Oligocene terrestrial ecosystems

A fossil flora from the basal Chatton Formation at Cosy Dell farm near Waimumu, Southland, New Zealand comprises wood, seeds, pollen and spores. A Late Oligocene age (25.4–24.4 Ma) determined from macro- and nannofossils constrains the beginning of marine transgression in this area. The palynoflora comprises more than 100 taxa, in addition to a few spores recycled from underlying Jurassic Murihiku basement. At least 16 ferns are present. Conifers include Araucariaceae and Podocarpaceae (Dacrydium, Dacrycarpus, Lagarostrobos, Microcachrys, Phyllocladus and Podocarpus). The angiosperm palynoflora comprises monocots including Arecaceae, Asparagaceae, Asteliaceae, Pandanaceae and Typhaceae and numerous dicots including Casuarinaceae, Cunoniaceae, Euphorbiaceae, Loranthaceae, Malvaceae, Myrtaceae, Nothofagaceae, Proteaceae and Strasburgeriaceae. The drift flora includes wood and seeds including the tropical legume Entada. Elongate limpets and mangrove snails provide indirect evidence for seagrasses and mangroves. The predominance of rainforest tree pollen and fern spores suggests a regional subtropical coastal forest and high rainfall.

Leaf fossils of Luzuriaga and a monocot flower with in situ pollen of Liliacidites contortus Mildenh. & Bannister sp. nov. (Alstroemeriaceae) from the Early Miocene

PREMISE OF THE STUDY The Foulden Maar lake sediments in Otago, South Island, New Zealand, date to the earliest Miocene and provide an important picture of the diversity of the Australasian biota, paleoecology, and climate at a time when New Zealand had a smaller land area than today. The diverse rainforest contains many taxa now restricted to Australia, New Caledonia, or South America. The presence of Luzuriaga-like fossils in these deposits is important for understanding Alstroemeriaceae evolution and the biogeography of genera shared between New Zealand and South America. METHODS Leaves and a flower with in situ pollen that resemble extant Luzuriaga are described and placed phylogenetically. Geographic range information and a molecular clock model for the Alstroemeriaceae were used to investigate possible biogeographic scenarios and the influence of the new fossil on inferred divergence times. KEY RESULTS Luzuriaga peterbannisteri Conran, Bannister, Mildenh., & D.E.Lee sp. nov. represents the first macrofossil record for Alstroemeriaceae. An associated Luzuriaga-like flower with in situ fossil pollen of Liliacidites contortus Mildenh. & Bannister sp. nov. is also described. The biogeographic analysis suggests that there have been several dispersal events across the Southern Ocean for the genus, with the fossil representing a now-extinct New Zealand lineage. CONCLUSIONS Luzuriaga was present in Early Miocene New Zealand, indicating a long paleogeographic history for the genus, and L. peterbannisteri strengthens biogeographic connections between South America and Australasia during the Oligocene and earliest Miocene.

Leaf fossils of Proteaceae tribe Persoonieae from the Late Oligocene–Early Miocene of New Zealand

Fossils from the Newvale lignite mine, Southland, are the first substantiated foliar records of Proteaceae subfamily Persoonioideae. The fossils possess very large stomata, a probable synapomorphy for Persoonioideae, and within Proteaceae the combination of this feature and more or less parallel-aligned, brachyparacytic stomatal complexes and undulate anticlinal epidermal cell walls is uniquely found in this subfamily. The new genus Persoonieaephyllum is described to recognise affinity of the fossil leaves and cuticles with tribe Persoonieae of Persoonioideae and their distinction from its only other extant representative, Placospermum. Two new species are described. P. ornatum is represented by linear leaves less than 20mm wide and possessing more or less parallel-aligned major veins. These leaves closely match those of extant hypostomatic, broad-leaved species of tribe Persoonieae and are distinct from Placospermum in venation and several cuticular details. P. villosum has so far been recovered only as cuticular material in disaggregated lignite. It is distinct from P. ornatum in having abundant trichome bases, an absence of surface tubercules, and even larger stomata (guard cells often >70 mm long). The fossils extend the known record of Persoonioideae in the New Zealand–New Caledonia region by ~20 million years.

Proteaceae leaf fossils from the Oligo–Miocene of New Zealand: new species and evidence of biome and trait conservatism

Abstract. At least seven foliar taxa of Proteaceae occur in Oligo–Miocene lignite from the Newvale site. These taxa include two new species of the fossil genus Euproteaciphyllum, and previously described species of tribe Persoonieae and Banksia. Other specimens from Newvale are not assigned to new species, but some conform to leaves of the New Caledonian genus Beauprea, which is also represented in the lignite by common pollen. Two other Euproteaciphyllum species are described from the early Miocene Foulden Maar diatomite site. One of these species may belong to Alloxylon (tribe Embothrieae) and the other to tribe Macadamieae, subtribe Gevuininae. Ecologically, the species from Newvale represented important components of wet, oligotrophic, open vegetation containing scleromorphic angiosperms and very diverse conifers. In contrast, Proteaceae were large-leaved and rare in Lauraceae-dominated rainforest at the volcanic Foulden Maar site. Overall, the Oligo–Miocene fossils confirm that Proteaceae was formerly much more diverse and dominant in the New Zealand vegetation, and provide fossil evidence for biome conservatism in both leaf traits and lineage representation.