Palaeontologia Electronica http://palaeo-electronica.org

DISPERSED LEAF CUTICLE FROM THE EARLY MIOCENE OF SOUTHERN

Mike Pole

ABSTRACT

This paper describes 115 parataxa of dispersed leaf fossil cuticle from 120 sam- ples from the Early Miocene of Central Otago (the fluvial-lacustrine Manuherikia Group) and Southland (the coastal deltaic East Southland Group), New Zealand. The modern affinities include Argophyllaceae (Argophyllum), Atherospermataceae, Casuarinaceae (Gymnostoma), Cunoniaceae-Elaeocarpaceae, Ericaceae, Gneta- ceae, Grisseliniaceae (Grisellinia), Meliaceae, Menispermaceae, (Hedy- carya), Myrsinaceae, Proteaceae (incl. Lomatia and Placospermum), Santalaceae (Notothixos), Sapindaceae, Strasburgeriaceae (Strasburgeria), and Winteraceae. The records of Argophyllaceae, Menispermaceae, Placospermum and Notothixos are the first of these families and genera for New Zealand. For the Argophyllaceae and Notothixos at least, these are the first known fossil records. With the exception of Cunoniaceae-Elaeocarpaceae, Ericaceae, Grisseliniaceae, Myrsinaceae, and Winter- aceae, which occur in the south of New Zealand today, the fossils indicate a more southerly range extension in the Early Miocene than today. This evidence of extended range along with a previously published high diversity of Lauraceae and conifers is probably the result of warmer conditions despite the fossil localities lying at about 50ºS in the Early Miocene – about 5 degrees further south than today. Argophyllum and Strasburgeria are evidence of a biogeographical link with , where they are now restricted. The were components of rainforest vegetation growing in microthermal to mesothermal temperatures.

Mike Pole. Queensland Herbarium, Brisbane Botanic Gardens Mt Coot-tha, Mt Coot-tha Rd, Toowong QLD 4066, [email protected]

KEY WORDS: Early Miocene, cuticle, stomata, epidermis, biogeography, biodiversity

INTRODUCTION perature which rapidly followed (Miller et al. 1991; Zachos et al. 2001). During the optimum rainforest The Miocene covers a period of extraordinary was widespread and thermophilic species global climate change, extending though for exam- grew poleward of their current extent. The suc- ple, the early Middle Miocene Climate Optimum ceeding temperature drop saw rainforest retreat (Flower and Kennett 1994; Shevenell et al. 2004; and its replacement with deciduous forest while although Böhme et al. 2007 identify a German opti- mum slightly earlier) and the profound drop in tem-

PE Article Number: 11.3.15A Copyright: Palaeontological Association October 2008 Submission: 12 January 2008. Acceptance: 10 September 2008

Pole, Mike S., 2008. Dispersed Leaf Cuticle from the Early Miocene of Southern New Zealand. Palaeontologia Electronica Vol. 11, Issue 3; 15A: 117p; http://palaeo-electronica.org/2008_3/153/index.html POLE: MIOCENE LEAF CUTICLE

NEW ZEALAND

Grey Lake St Bathans + Blue Lake 45S˚ Mata Ck 45S

Bannockburn 170E Nevis

50 km Harliwichs Pit

46S˚ 170E˚ DUNEDIN

Chatton d1324 46S˚ d1027 d1115 d1051 d1026 d1057 GORE d1108 d1024 Waimumu d1052 d1109 d1110 Hedgehope d1243 d1121 d1124 MATAURA d1077 d1053 d1246 d1078 d1049 Makarewa d1107 Morton d1106 Mains INVERCARGILL d1105 d1297 Kapuka d1104 d1298 Waimatua d1102 d1299 d1141 d1101 d1142 d1143 d1144 d1296 d1294 10 km d1295 169E˚

Figure 1. Location Maps. Upper map shows the extent of the Manuherikia Group schematically as a grey circle. Blue Lake is the location of sample numbers prefixed with “BL-“, Grey Lake is the location of sample numbers prefixed with “GL-“, Mata Creek is the location of samples prefixed with “Mata-“, Harliwichs Pit is the loca- tion of sample numbers prefixed with “Harliwich-“. Lower map shows the position of all drill-cores in South- land (the location of sample numbers prefixed with “Sthd-“) from which fossils were obtained. For precise locations of all samples, see Appendix 1 and 2. some deciduous forest became scrubland or The fluvial-lacustrine Manuherikia Group of Central grassland (Flower and Kennett 1994). Otago (Douglas 1986) lay to the north of the Southern New Zealand has proven to be an coastal fluvial –deltaic East Southland Group important source of Early Miocene plant fossils. (Isaac and Lindqvist 1990). Extensive palynologi-

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cal studies have been carried out in both areas Groups. To this end it has no taxonomic focus, and (e.g. Pocknall and Mildenhall 1984; Mildenhall and includes a large proportion of taxonomic unknowns Pocknall 1989). Leaf and fruit impressions found in but is a necessary step prior to a synthesis of the the Manuherikia Group include Eucalyptus (Pole vegetation. 1993a), Allocasuarina (Campbell and Holden 1984) and a legume (Pole 1992). They demon- METHODS strate the variety of vegetation types which existed Fossil-cuticle bearing samples of carbon- in the Early Miocene and from which significant cli- aceous mud were made in 120 locations in the mate change can be inferred (Pole 2003). Palyno- Manuherikia and East Southland Groups (Fig. 1). logical studies also indicate the importance of Sample numbers prefixed by “BL-“ and “GL-“ come burning during part of the deposition of the Manu- from the St Bathans Paleovalley, those prefixed herikia Group (Mildenhall 1989; Pole and Douglas with “Mata-“ come from Mata Creek and prefixed 1998). Along with the vegetation there was a fauna with “Harliwich-“ come from Harliwich’s Coal Pit, which is becoming increasingly well-known (Feld- those prefixed with “Sthd-“ come from the East man and Pole 1994; McDowall and Pole 1997; Southland Group. Most Manuherikia Group sam- Pole et al. 2003; Worthy et al 2006, 2007). Earlier ples were about 300g of sediment, with about half work on plant macrofossils focussed on impres- of that being processed. Seven samples were sions (see Pole 1993b and references therein) more intensively sampled by repeat visits as they although the pioneering investigations of Holden have more abundant and intact angiosperm fossils. (1987) documented the presence of leaf cuticle in These samples are BL-30, GL-01, GL-02, GL-32, the East Southland Group. More recently dis- H41/f038, BL-32, Mata-01 where up to one or two persed cuticle has been central to research in the kilos were processed. Samples from the East field (Pole 1997, 1998a, 2007a, 2007b) and has Southland Group come from drillcore and were lim- resulted in the discovery of two cycads (Hill and ited by the diameter of the drill core (c. 50 mm) and Pole 1994, Pole 2007b), 16 conifers (Pole 2007b), the desire to preserve the integrity of the core as and 11 Proteaceae (Pole 1998a) and 25 Lau- much as possible. Sample size was about 50 g and raceae (Pole 2007a) and 17 monocots or possible the amount of sediment prepared was about half of monocots (Pole 2007c). The identification of these that. taxa may help quantify aspects of the warmer glo- For the St Bathans Paleovalley samples a bal climate at a time when New Zealand was fur- simple stratigraphic column is inappropriate as the ther south than it is today. For instance the sampled units are lensoid and widely-spread. How- southern limit of the Lauraceae family lies within ever, a grid-reference and notes will allow reloca- New Zealand today, where only four species of tion of the sample site. These details can be found broad-leaved Lauraceae are present (Allan 1982; in Appendix 1 and 2. Wright 1984). However, in the Early Miocene 25 The samples were disaggregated by covering species are known (Pole 2007a). In the debate with hot water and adding 40 % hydrogen perox- over the importance of long distance dispersal in ide, fines were then removed by sieving through a the origin of New Zealand’s flora (Knapp et al. 1 mm mesh. What remained was typically a “hash” 2005; McGlone 2005; Pole 1994; Winkworth et al. of cuticle fragments, typically around 1 - 10 mm 2002) genetic data provided by extant plants across. Cuticle could then be purified with further seems to be strongly favouring the dispersal hydrogen peroxide treatment (which removes option, but good plant fossil evidence is still opaque cellular material) and hydrofluoric acid to needed to date nodes on phylogenetic trees and to remove silicates. It was then stained using either demonstrate where extinction has entirely elimi- safranin or crystal violet. This concentrate was nated taxa from major biogeographic regions. then scanned in a Petri dish under binocular micro- Basic description of cuticle morphologies will also scope and individual cuticle fragments were iso- prove useful for stratigraphic purposes. While pre- lated and mounted on microscope slides using vious papers have documented the more diverse thymol glycerine jelly for Transmitted Light Micros- taxonomic groups (see previous discussion) and a copy (TLM). When sufficient additional material further paper documents the presence of families was present, also mounted on stubs for scanning currently endemic to New Caledonia, the aim of electron microscopy (SEM) using double-sided this paper is to complete the documentation of the tape, and coated with platinum. Catalogue num- dispersed fossil leaf cuticle morphologies currently bers for material mounted on microscope slides known from the Manuherikia and East Southland

3 POLE: MIOCENE LEAF CUTICLE are prefixed with “SB” or “SL” and SEM stubs are to the stomatal pore and the pair of guard cells, prefixed with “S-“. and ‘stomatal complex’ for the stoma plus subsid- The task of distinguishing taxa was an itera- iary cells. The inadvisability of constructing a termi- tive process that has taken over 15 years. This nology with a mixture of purely morphological work mainly used TLM and enlarged TLM photo- terms as well as terms based on developmental graphs were employed which could be laid side by processes have been discussed by these authors. side to facilitate comparison. SEM observation was However, there are occasions where some evi- useful to interpret the three-dimensional structure dence of developmental process is obvious in and view very fine details. However, for pragmatic cuticular fragments (for instance where one cell purposes this study focuses on distinguishing char- has been divided in two by a new wall) and need to acters which can be viewed under TLM. be indicated. For these instances I have used As a general observation, it is relatively simple terms (heliocyctic and tangenticytic) from Timonin to study one sample and separate out distinct taxa. (1995). More recently Carpenter (2005) has intro- However, as more and more samples are added, duced several terms which incorporate develop- perhaps with slightly differing degrees of preserva- mental considerations. He did not discuss these tion, staining, environmental differences, etc, the terms in the context of several previous authors issue of taxon distinction becomes critical. As more (for instance Timonin 1995) but some of them are samples are studied, typically many ‘provisional’ also clearly applicable to the fossils described here taxa are merged with one another. The issue then and so they are listed alongside Timonin’s. The becomes one of finding similarities rather than dif- reader should be aware that what appear to be fun- ferences. The benefits of a large sample-base damentally different terms describing stomatal being that a more realistic concept of taxa complex structure may result from the subjectivity emerges. However, it is certain that a few taxa dis- involved in deciding what is and isn’t a subsidiary tinguished here are likely to cover more than one cell. “real” species, and thus the final number will be an To clarify some further terms; OSL = outer sto- underestimate. Where there may be some issues matal ledge. By “outer stomatal ledge thickness” of distinction; these are addressed under the sub- here is meant the combined thickness of ledges heading “Distinguishing features”. and any underlying cuticle as seen under TLM In this work, cuticle morphologies are view. By “giant stomatal complexes” is meant any described as parataxa. Each parataxon is prefixed distinctly different population of complexes, either by “CUT-“, then a letter. This is a purely pragmatic in terms of size, or by obviously increased develop- subdivision of the cuticle taxa into large groups. ment of subsidiary cells around them. “Networking” For instance, “L” includes Lauraceae, “M” the Myrt- is used to describe the situation where contact or aceae, and “P” for Proteaceae. Most parataxa subsidiary cells are shared between stomatal com- described in this report were given a “Z” as they plexes (Pole 1998b). When stomata are known to are the large group “left over” once the more imme- be on one leaf surface only the distribution is diately identifiable groups had been dealt with. assumed to be hypostomatic. In all other cases the Finally, each parataxon gets a unique string of distribution with respect to leaf surfaces is three letters. This is meant as a flexible system to unknown. Stomatal size classes follow Wilkinson deal with the disparate and poorly hierarchical mor- (1979). “Texture” refers to a pattern on the outer or phologies of cuticle fragments. The intention is that inner surfaces of cuticle, such as “granular” which parataxa are equivalent to species. For each is of much smaller dimensions than normal epider- parataxon a reference specimen and sample is mal cells. Ornamentation refers to a pattern on the nominated, which are equivalent to the holotype outer cuticular surface, such as ridging, which is and type locality of a Linnean species. A single comparable to the size of normal epidermal cells. It specimen is also nominated from each other sam- does not include papillae or scales. ple that the taxon is recorded from. For examples Taxonomic identification into the Linnean sys- of this methodology see Pole (2007a). tem is based on morphological characters which Standard epidermal terminology is used have been detailed in the published literature as to describe the cuticle taxa and is based on well as a large cuticle reference collection devel- authors such as Baranova (1987, 1992), Stace oped by the author. To date this collection includes (1965); Dilcher (1974), Hewson (1988), Payne around 4000 species in about 1500 genera and (1978), and Wilkinson (1979). Carpenter (2005) is 285 families of mostly rainforest taxa from around followed in the use of ‘stoma’ (stomata pl.) to refer the world. In this paper cuticle preparations of

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Table 1. Taxonomic summary of families and genera identified in this study and their present distribution (Family distri- butions from Watson and Dallwitz, 1992 onwards, genera from the Global Biodiversity Information Facility, http://us.mir- ror.gbif.org).

Family and current distribution Argophyllaceae Argophyllum. Australia, New Caledonia. Atherospermataceae South America to Australasia. Restricted to northern parts of New Zealand. Casuarinaceae Gymnostoma. Througout Malesia, New Guinea, New Caledonia and highly restricted in Australia to northeast Queensland, absent in New Zealand. Cunoniaceae-Elaeocarpaceae Widespread in New Zealand and from Southeast Asia, to Australia, Central and South America and (Cunoniaceae) South Africa. Ericaceae Cosmopolitan Gnetaceae Closest to extant Gnetum although the fossil is regarded as an extinct genus. Gnetum occurs from Amazonia, tropical west Africa, and tropical Asia from Bombay to Fiji (absent in New Caledonia and New Zealand). Grisseliniaceae Grisellinia. South America and widespread in New Zealand. Meliaceae Pantropical to subtropical and warm. To latitude 41° 30' S in New Zealand. Menispermaceae Pantropical and warm. The family is absent from New Zealand today although it does occur in Australia to about 37 °S. The fossil localities are further south than any living occurrence today. Monimiaceae Hedycarya. Australia, New Guinea, New Caledonia, widespread in New Zealand. Myrsinaceae Comparable with Ardisia although other genera are not yet ruled out. Ardisia is pantropical to subtropical, but is absent from New Zealand. Proteaceae Incl. Lomatia and Placospermum The family is present in the northern half of New Zealand. Placospermum is an Australian endemic, Lomatia is found in Australia and South America. Santalaceae Notothixos. Ceylon, Myanmar, Philippines, Malesia, Australia. Sapindaceae Temperate to tropical. Not extending to the far south of New Zealand. Strasburgeriaceae Strasburgeria. New Caledonian endemic. Winteraceae Distinct from extant New Zealand Pseudowintera. Malaysia, Pacific, Australia, New Zealand, Central and South America. Pseudowintera is widespread in New Zealand.

extant herbarium material cite the original herbar- Southland Groups suggest at least 100 tree spe- ium sheet number (“AQ” refers to catalogue num- cies grew with them, and probably at least 120 bers of specimens in the Queensland Herbarium, (Pole 2007a). Brisbane; “CANB” of specimens in the Australian Some of the families described as fossils in National Herbarium, Canberra; “OTA” of speci- this study occur in the extant flora of New Zealand, mens in the Botany Department Herbarium, Uni- but others represent the first known occurrence of versity of Otago, Dunedin) and material in the the family in New Zealand, and as such, add to author’s own reference herbarium is prefixed with another dimension of biodiversity. Among the uni- “OPH”. All material is stored in the Queensland dentified taxa there are also likely to be totally Herbarium, Toowong. extinct genera represented, but these will need more information, for instance evaluation of the RESULTS AND DISCUSSION corresponding whole leaf architecture. A brief discussion of the significance of the One hundred and fifteen morphologies of cuti- identified taxa (Summarised in Table 1) follows and cle parataxa were distinguished in the 120 samples the detailed descriptions are relegated to Appendix (Appendix 3). Fifty eight taxa have been placed 3. A fuller synthesis will await publication of some into extant families, and eight into extant genera. further taxa and will be integrated with more The remainder have not been identified, or have detailed stratigraphy and sedimentology. less certain, and only suggested affinities. The spe- Evidence for the presence of Gnetales in New cies-level biodiversity represented here makes a Zealand is not new – palynological works have substantial contribution to that predicted from the reported the pollen Ephedripites notensis Cookson diversity of one family in the fossil record, the Lau- (1957) for many years (e.g. Couper 1960). This raceae. Based on the current relationship between pollen type has been said to have affinities with the number of Lauraceae species and total tree extant Ephedra, an arid-land plant of North Amer- diversity for rainforest sites in Australia, the 25 Lau- ica. Its presence in an obviously wet-environment raceae taxa known from the Manuherikia and East

5 POLE: MIOCENE LEAF CUTICLE

is something of an anomaly, and the likelihood is is a feature of the wet, mesothermal rainforests of that the family identification is correct, but that it northern Australia (Webb and Tracey 1981). represents some other, extinct genus. The pres- The Menispermaceae represent a new addi- ence of cuticle with similarities (but distinct from) tion to the New Zealand record. They are a family Gnetum reinforces the view. The cuticle morphol- dominated by climbers or creepers, mostly tropical, ogy is also present in the Late Eocene of the reaching as far south as Victoria in Australia. Waikato Coal Measures (Pole unpublished) and a Added to the Rhipogonum already known (Pole similar parataxon is in the Early Eocene of Tasma- 1993c) this identification allows the inference of a nia (Pole 2007d). It may represent a plant lineage second climber in the Manuherikia Group based on which continued through the putative Oligocene macrofossil evidence. drowning of New Zealand (Cooper and Cooper The genus Notothixos is a new addition to the 1995), or it may represent long-distance dispersal. New Zealand record and is the first fossil record of In this context it is of interest that DNA evidence the genus, and perhaps of the Viscaceae. has suggested that Gnetum has dispersed globally Notothixos is a mistletoe and occurs today from within the Neogene (Won and Renner 2006). Ceylon to Malesia and down the east coast of Aus- The Winteraceae are represented in New tralia to Victoria (Barlow 1983). Zealand today by three species in the genus Strasburgeria (Strasburgeriaceae) is currently Pseudowintera (Allan 1982). The cuticle record endemic to New Caledonia, but has been identified suggests that other genera of the family were with the pollen Bluffopollis scabratus (Couper) present in the Miocene (there is no Miocene cuticle Pocknall & Mildenhall (1984), which is present record of Pseudowintera). The inference of higher throughout the Tertiary of New Zealand (Jarzen generic diversity at this time supports the conclu- and Pocknall 1993) and is a common and wide- sions of the palynological record (Pocknall and spread component in many of the samples studied Mildenhall 1984; Raine et al. 2006) where Harrisi- here (pers. obs.). This is the first fossil record of the pollenites annulatus Mildenhall & Crosbie (1979) is cuticle. regarded as representing Zygogynum, a New Griselinia is the only genus in the Griselini- Caledonian genus, and Pseudowinterapollis cou- aceae, which is found today in both New Zealand peri Krutzsch (1970) is consistent with Drimys, and South America. This is the first fossil record of Pseudowintera and Tasmannia and P. cranwelliae the genus. (Stover and Partridge 1973) represents Tasman- The Meliaceae and Sapindaceae both have a nia. Today the family is a prominent component of single rainforest species in New Zealand today; rainforests in the cool southern Hemisphere as well Dysoxylon and Alectryon respectively. In the Sap- as New Caledonia. indaceae there is also Dodonaea, a shrub of more The Atherospermataceae and Monimiaceae open vegetation. The cuticle evidence confirms the are also important families in cooler Southern palynological evidence for these families in the Hemisphere rainforests and both are in the current Neogene, but like the pollen, is not helpful to New Zealand flora (Hedycarya in the Monimiaceae generic identification. Although Couper (1960) and Laurelia in the Atherospermataceae). Little is identified pollen of Dysoxylum in the Cenozoic, nei- known of their prehistory; the long-ranging pollen ther the genus nor the Meliaceae is included in type Liliacidites variegatus Couper (1953) may rep- Raine et al. (2006). The fossil pollen may not be resent Atherospermataceae, though it may also be distinguishable from that of many other genera of produced by the Lilliaceae (Raine et al. 2006), but Meliaceae (e.g. Mildenhall 1980). fossil wood has been reported from Antarctica, but The Cunoniaceae-Elaeocarpaceae are promi- cannot be related to any single extant genus nent components of the extant New Zealand flora (Poole and Francis 1999). as the genera Elaeocarpus and Weinamnnia. The Proteaceae has two genera in the extant Miocene Elaeocarpaceae leaf impressions were flora of New Zealand, Knightia, and Toronia, both described by Pole (1993; which on the basis of members of the warmer rainforests. The cuticle their basic shape, domatia, and spinose attach- evidence supports the palynological evidence that ments on the teeth, I am now certain are Elaeocar- the Proteaceae were much more diverse in New pus). The Myrsinaceae is present in New Zealand Zealand. The identification of Placospermum in this today as the widespread genus Myrsine and also study adds another genus to the New Zealand the highly restricted Elingamita (Allan 1982). A record and those known from the Miocene (Pole compression fossil of the Mysinaceae was 1998a). Today a diverse component of Proteaceae described by Pole (1996) from the Miocene Foul-

6 PALAEO-ELECTRONICA.ORG den Hills Diatomite (and based on the vein archi- REFERENCES tecture, I am now certain that this is an Ardisia). Allan, H.H. 1982. Flora of New Zealand. Volume 1. Indig- The Manuherikia Group would have lain at enous Tracheophyta - Psilopsida, Lycopsida, Filicop- about 50ºS in the Early Miocene (Lawver and sida, Gymnospermae, Dicotyledons. Government Gahagan 2003), about 5º further south than its Printer, Wellington. present position. The latitude of the East Southland Angiosperm Phylogeny Group. 2003. An update of the Group, like today, would have been only a degree Angiosperm Phylogeny Group classification for the or two higher than the Manuherikia Group. Both of orders and families of flowering plants: APG II. these areas lie southwards of the southern limit of Botanical Journal of the Linnean Society, 141: 399- several plant families represented by the cuticle 436. (Table 1), and therefore indicate a more southerly Bailey, F.M. 1885. Contributions to the Queensland flora. range extension than found today. This likely Part III. Proceedings of the Royal Society of Queen- sland 1: 140-150. refects warmer conditions, consistent with other Baker, R.T. 1899. Contributions to a knowledge of the fossil (Hornibrook 1992) and isotopic evidence . No. II. Proceedings of the Linnean (Shackleton and Kennett 1975). Judged on the Society of New South Wales, Series 2, 24: 437-447. present distribution of these taxa, temperatures Baranova, M. 1987. Historical development of the were in the microthermal to mesothermal range present classification of morphological types of sto- (sensu Nix 1982) and the vegetation was rainforest mas. The Botanical Review, 53: 53-79. (sensu Bowman 2000). Baranova, M. 1992. Principles of comparative stomato- graphic studies of flowering plants. The Botanical CONCLUSION Review, 58: 49-99. Barlow, B.A. 1983. A revision of the genus Notothixos A diverse range of leaf cuticle is present in a (Viscaceae). Brunonia, 6: 1-24. wide range of localities in the Early Miocene of Barnes, R.W., Hill, R.S. and Bradford, J.C. 2001. The southern New Zealand, ranging from the inland to history of Cunoniaceae in Australia from macrofossil coastal sites. The taxa described include families evidence. Australian Journal of Botany, 49: 301-320. still present in New Zealand today, as well as some Barrera, E.M. and Meza, I.P. 1992. Arquitectura foliar de which are now locally extinct, and some which are arboles chilenos I. Subclase Magnoliidae. Boletin now restricted to New Caledonia. A large propor- Museo Nacional de Historia Natural Chile, 43: 41-54. Bentham, G. 1870. Flora Australiensis, vol. 5. L. Reeve tion cannot yet be placed into the Linnean hierar- and Co., London. chy with any confidence, but will be useful in Böhme, M., Bruch, A.A. and Selmeier, A. 2007. The stratigraphic and fine-scale differentiation of vege- reconstruction of Early and Middle Miocene climate tation and habitat types, as well as providing useful and vegetation in Southern Germany as determined information on biodiversity and climate. In this way from the fossil wood flora. Palaeogeography, Palaeo- cuticle parataxa will compliment the palynological climatology, Palaeoecology, 253: 91-114. record as additional taxa to help refine relative Bongers, J.M. 1973. Epidermal leaf characteristics of the ages and distinguish assemblages. Winteraceae. Blumea, 21: 381-411. The fossils also have implications for biogeog- Bowman, D.M.J.S. 2000. Australian rainforests: Islands raphy. The new family and generic records provide of green in the land of fire. Cambridge University interesting range-extensions to be explained. In Press, Cambridge. Brown, R. 1814. Prodromous Florae Novae Hollandiae particular they add to the growing evidence of a et Insulae Van-Diemen. Taylor, London. link between New Zealand’s past vegetation and Candolle, de A.P. 1816. Essai sur les propriétés médi- the extant vegetation of New Caledonia. They also cales des plantes. Crochard, Paris. add to the established knowledge of a warmer than Campbell, J.D. and Holden, A.M. 1984. Miocene casuar- present Early Miocene, and will help elucidate the inacean fossils from Southland and Central Otago, specific details of this climate. New Zealand. New Zealand Journal of Botany, 22: 159-67. ACKNOWLEDGEMENTS Carpenter, K.J. 2005. Stomatal architecture and evolu- tion in basal angiosperms. American Journal of Bot- I am most grateful to the Queensland Herbar- any, 92: 1595-1615. ium, the Australian National Herbarium, and the Carpenter, R.J. 1994. Cuticular morphology and aspects Botany Department of the Otago University for of the ecology and fossil history of North Queensland access to their collections. Many thanks to two rainforest Proteaceae. Botanical Journal of Linnean anonymous referees for their help in improving this Society, 116: 249-303. paper.

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Carpenter, R.J., Hill, R.S., Greenwood, D.R., Partridge, Forster, J. G. 1786. Florulae Insularum Australium Pro- A.D. and Banks, M.A. 2004. No snow in the moun- dromus. Joann. Christian. Dieterich, Göttingen, tains: Early Eocene plant fossils from Hotham Gray, A. 1866. Monimiaceae. Journal of Botany, British Heights, Victoria, Australia. Australian Journal of Bot- and Foreign, 4: 80-85. any, 52: 685-718. Guillaumin, A. 1942. Contribution a la flore de la Nou- Cookson, I.C. 1957. On some Australian Tertiary spores velle-Calédonie 80. Bulletin de Museum National and pollen grains that extend the geological and geo- Histoire Naturelle (sér. 2) 14: 451-. 456. graphical distribution of living genera. Proceedings of Hewson, H.J. 1988. Plant Indumentum - A Handbook of the Royal Society of Victoria, 69: 41-53. Terminology. Australian Flora and Fauna Series 9. Cooper, A. and Cooper, R.A. 1995. The Oligocene bot- Canberra, Australian Government Publishing Service tleneck and New Zealand biota: genetic record of a Hill, R.S. 1989. Early Tertiary leaves of the Menisper- past environmental crisis. Proceedings of the Royal maceae from Nerriga, New South Wales. Alcheringa, Society of London B, 261: 293-302. 13: 37-44. Couper, R.A. 1953. Upper Mesozoic and Cainozoic Hill, R.S. and Pole, M.S. 1994. Two new species of spores and pollen grains from New Zealand. New Pterostoma R.S. Hill from Cenozoic sediments in Zealand Geological Survey Palaeontological Bulletin, Australasia. Review of Palaeobotany and Palynol- 22: 1-77. ogy, 80: 123-130. Couper, R.A. 1960. New Zealand Mesozoic and Caino- Holden, A.M. 1987. Vegetation-lithotype correlations in zoic plant microfossils. New Zealand Geological Sur- New Zealand lignites. New Zealand Energy vey Paleontological Bulletin, 32: 1-88. Research and Development Committee Publication Cronquist, A. 1981. An Integrated System of Classifica- R150. tion of Flowering Plants. Columbia University Press, Hooker, J. D. 1864. Handbook of the New Zealand Flora: New York. a systematic description of the native plants of New Cunningham, A. 1838. Florae Insularum Novae Zealand and the Chatham, Kermadec's, Lord Auck- Zelandiae Precursor; or a Specimen of the land's, Campbell's and Macquarie's Islands. Part I. Reeve,.London, Botany of the Islands of New Zealand. Hornibrook, N.de.B. 1992. New Zealand Cenozoic Annals of Natural History, 1: 376-381. marine paleoclimates: a review based on the distribu- Cunningham, A. 1839. Specimen of the botany of New tion of some shallow water and terrestrial biota. p. Zealand. Annals of Natural History, 3: 260-261. 83-106. In: Tsuchi, R., and Ingle J.C. Jr. (eds), Pacific Dilcher, D.L. 1974. Approaches to the identification of Neogene. Environment, Evolution, and Events. Uni- angiosperm leaf remains. The Botanical Review, 40: versity of Tokyo Press, Tokyo. 1-157. Isaac, M.J. and Lindqvist, J.K. 1990. Geology and lignite Domin, K. 1930. Beitrage zur Flora und Pflanzengeogra- resources of the East Southland Group, New phie Australiens. E. Schweizerbart, Stuttgart. Zealand. New Zealand Geological Survey Bulletin Douglas, B.J. 1986. Lignite resources of Central Otago. n.s., 101: 1-202. New Zealand Energy Research and Development Jarzen, D. M. and D. T. Pocknall. 1993. Tertiary Bluf- Committee, Publication P104, 1-388. fopollis scabratus (Couper) Pocknall & Mildenhall, Endlicher, S.L 1841. Iconographia Generum Plantarum. 1984 and modern Strasburgeria pollen: a botanical Vindobonae, Vienna. comparison. New Zealand Journal of Botany, 31: Endlicher, S.L. 1843. Genera Plantarum Secundum 185-192. Ordines Naturales Disposita. Botanische Zeitung, 3: Johnson, L.A.S. 1980 Notes on Casuarinaceae. Telopea 288-290 2: 83-84. Feldmann, R. and Pole, M.S. 1994. A new species of Jordan, G.J. 1997. Evidence of Pleistocene plant extinc- Paranephrops White, 1842: a fossil freshwater cray- tion and diversity from Regatta Point, western Tas- fish (Decapoda: Parastaciadae) from the Manu- mania, Australia. Botanical Journal of Linnean herikia Group (Miocene), Central Otago, New Society, 123: 45-71. Zealand. New Zealand Journal of Geology and Geo- Jordan, G.J. and Hill, R.S. 1995. Oligocene leaves of physics, 37: 163-167. Epacridaceae from Little Rapid River, Tasmania, and Flower, B.P. and Kennett, J.P. 1994. The middle Miocene the identification of fossil Epacridaceae leaves. Aus- climatic transition: East Antarctic ice sheet develop- tralian Systematic Botany, 8: 71-83. ment, deep ocean circulation and global carbon Jordan, G.J. and Hill, R.S. 1996. The Fossil Record of cycling. Palaeogeography, Palaeoclimatology, Palae- the Epacridaceae. Annals of Botany, 77: 341-346. oecology, 108: 537-555. Jussieu, A.L. 1789. Genera Plantarum. Herissant, Paris. Forster, J. R., and Forster, G. 1776. Characteres Jussieu, A.L. 1809. Memoire sur les Monimiees, nouvel Generum Plantarum. White, Cadwell, and Elmsly, ordre de plantes. Annales Muséum National D'His- London. toire Naturelle, Paris 14: 116-135.

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Knapp, M., Stöckler, K. Havell, D., Delsuc, F., Sebastiani, 47-76.In: Evolution of the Flora and Fauna of Arid Aus- F. and Lockhart, P.J. 2005. Relaxed Molecular Clock tralia. (Eds) Barker W.R. and. Greenslade, P.J.M. Provides Evidence for Long-Distance Dispersal of Peacock Publications, Adelaide. Nothofagus (Southern Beech). PLoS Biology, 3 (1) Oliver, D. 1863. Notes on the Loranthaceae, with a syn- e14. opsis of the genera. Journal of the Proceedings of Koorders, S. H. 1898. Mededeelingen uit 's Lands Plant- the Linnean Society, Botany 7: 90-105. entuin. Kolff, Batavia. Payne, W.W. 1978. A glossary of plant hair terminology. Krutzsch, W. 1970. Zur Kenntnis fossiler disperser Tet- Brittonia, 30: 239-255. radenpollen. Palaeontologische Abhandlungen Pocknall, D.T. and Mildenhall, D.C. 1984. Late Oligocene Abteilung B., Palaeobotanik, 3: 399-430. -Early Miocene spores and pollen from Southland, Lange, R.T. 1978. Some Eocene leaf fragments compa- New Zealand. New Zealand Geological Survey Pale- rable to Proteaceae. Journal of the Royal Society of ontological Bulletin, 51: 1-66. Western Australia, 60: 107-114. Pole, M.S. 1992. Fossils of Leguminosae from the Lawver, L.A. and Gahagan, L.M. 2003. Evolution of Cen- Miocene Manuherikia Group of New Zealand. p. 251- ozoic seaways in the circum-Antarctic region. 258. In Herendeen, P.S. and Dilcher, D.L. (eds), Palaeogeography, Palaeoclimatology, Palaeoecol- Advances in Legume Systematics: Part 4. The Fossil ogy, 198: 11-37. Record. The Royal Botanic Gardens, Kew. Lindley, J. 1830. An Introduction to the Natural System of Pole, M.S. 1993a. Early Miocene floras of the Manu- Botany. Longman, London. herikia Group, New Zealand. 7. Myrtaceae, including Lindley, J. 1834. Botanical Register 20. James Ridgway Eucalyptus. Journal of the Royal Society of New and Sons, Piccadilly. Zealand, 23: 313-328. McDowall, R.M. and Pole, M.S. 1997. A large galaxiid Pole, M.S. 1993b. Early Miocene flora of the Manu- fossil (Teleostei) from the Miocene of Central Otago, herikia Group, New Zealand. 10. Paleoecology and New Zealand. Journal of the Royal Society of New stratigraphy. Journal of the Royal Society of New Zealand, 27: 193-198. Zealand, 23: 393-426. McGlone, M.S. 2005. Goodbye Gondwana. Journal of Pole, M.S. 1993c. Early Miocene flora of the Manuherikia Biogeography, 32: 739–740. Group, New Zealand. 5. Smilacaceae, Polygo- Metcalfe, C. R. 1987. Anatomy of the Dicotyledons. III. naceae, Elaeocarpaceae. Journal of the Royal Soci- Magnoliales, , and Illiciales. Oxford, Claren- ety of New Zealand, 23: 289-302. don Press. Pole, M.S. 1994. The New Zealand flora: entirely long- Miers, J. 1867. A few remarks on the Menispermaceae. distance dispersal? Journal of Biogeography, 21: Annals and Magazine of Natural History (series 3), 625–635. 19: 85-95. Pole, M.S. 1996. Plant macrofossils from the Foulden Mildenhall, D.C. 1980. New Zealand Late Cretaceous Hills Diatomite (Miocene), Central Otago, New and Cenozoic plant biogeography: a contribution. Zealand. Journal of the Royal Society of New Palaeogeography, Palaeoclimatology, Palaeoecol- Zealand, 26: 1-39. ogy, 31: 197-233. Pole, M.S. 1997. Miocene conifers from the Manuherikia Mildenhall, D.C. 1989. Summary of the age and paleo- Group, New Zealand. Journal of the Royal Society of ecology of the Miocene Manuherikia Group, Central New Zealand, 27: 355-370. Otago, New Zealand. Journal of the Royal Society of Pole, M.S. 1998a. The Proteaceae record in New New Zealand, 19: 19-29. Zealand. Australian Systematic Botany, 11: 343-372. Mildenhall, D.C., and Crosbie, Y.M. 1979. Some porate Pole, M.S. 1998b. Paleocene gymnosperms from Mount pollen from the upper Tertiary of New Zealand. New Somers, New Zealand. Journal of the Royal Society Zealand Journal of Geology and Geophysics, 22: of New Zealand, 28: 375-403. 499-508. Pole, M.S. 2003. New Zealand climate in the Neogene Mildenhall, D.C. and Pocknall, D.T. 1989. Miocene-Pleis- and implications for global atmospheric circulation. tocene spores and pollen from Central Otago, South Palaeogeography, Palaeoclimatology, Palaeoecol- Island, New Zealand. New Zealand Geological Sur- ogy, 193: 269-284. vey Palaeontological Bulletin, 59: 1-128. Pole, M. 2007a. Lauraceae Macrofossils and Dispersed Miller, K.G., Wright, J.D., and Fairbanks, R.G. 1991. Cuticle from the Miocene of Southern New Zealand. Unlocking the Ice House: Oligocene-Miocene oxygen Palaeontologia Electronica 10; 3A:38p, 65MB; http:// isotopes, eustasy, and margin erosion. Journal of palaeo-electronica.org/paleo/2007_1/zealand/ Geophysical Research, 96 (B4): 6829–6848. index.html Mueller, F.J.H. von, 1861. Fragmenta Phytographiae Pole, M.S. 2007b. Conifer and Cycad Distribution in the Australiae 2. Auctorite Gubern, Melbourne. Miocene of Southern New Zealand. Australian Jour- Mueller, F.J.H. von, 1868. 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Pole, M. 2007d. Early Eocene Dispersed Cuticles and Stace, C.A. 1965. Cuticular studies as an aid to plant tax- Mangrove to Rainforest Vegetation at Strahan- onomy. Bulletin of the British Museum (Natural His- Regatta Point, Tasmania. Palaeontologia Electronica tory) Botany, 4: 1-78. Vol. 10, Issue 3; 15A:66p. Stover, L.E. and Partridge, A.D. 1973. Tertiary and Late Pole, M. and Douglas, B.J. 1998. A quantitative palynos- Cretaceous spores and pollen from the Gippsland tratigraphy of the Miocene Manuherikia Group, New Basin, southeastern Australia. Proceedings of the Zealand. Journal of the Royal Society of New Royal Society of Victoria, 85: 237-286. Zealand, 28: 405-420. Takhtajan, A.L. 1987. System of Magnoliophyta. Acad- Pole, M.S., Douglas, B.J. and Mason, G. 2003. The ter- emy of Sciences, Leningrad. restrial Miocene biota of southern New Zealand. Timonin, A.C. 1995. 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Wright, A.E. 1984. Beilschmiedia Nees (Lauraceae) in Zachos, J., Pagani, M., Sloan, L., Thomas, E., and Bil- New Zealand. New Zealand Journal of Botany, 22: lups, K. 2001. Trends, Rhythms, and Aberrations in 109-125. Global Climate 65 Ma to Present. Science, 292, 686– 693.

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APPENDIX 1. DETAILS FOR ST BATHANS PALEOVALLEY, MATA CREEK AND HARLIWICH’S PIT SAMPLES (ALL MANUHERIKIA GROUP

Sample Grid reference Stratigraphic notes

AFW-18 H41/6006 8493 Uncertain precise relationship

AFW-23 H41/6003 8490 Uncertain precise relationship BL-01 H41/5799 8824 c. 6 m below BL-02

BL-02 H41/5799 8824 c. 6 m above BL-01

BL-03 H41/5799 8824 c. 6 m below BL-01 BL-04 H41/5796 8827 c. 7 m above BL-02

BL-05 H41/5796 8827 c. 6 m above BL-04

BL-06 H41/5796 8829 c. 10 m above BL-05

BL-07 H41/5796 8842 possibly equivalent to BL-30 BL-08 2257934 5588425 c. 15 m above BL-07

BL-09 H41/5791 8843 c. 20 m above BL-08

BL-10 H41/5776 8866 c. 23 m above BL-30 BL-11 H41/5777 8865 c. 2.5 m above BL-10

BL-12 H41/5778 8864 c. 7 m above BL-10

BL-13 H41/5778 8862 c. 9 m above BL-10 BL-14 H41/5787 8861 c. 6 m below BL-15

BL-15 H41/5782 8865 c. 6 m below BL-33

BL-16 H41/5779 8872 c. 6 m below BL-14, possibly equivalent to BL-30 BL-17 H41/5778 8878 c. 15-20 m below Bl-16

BL-18 H41/5780 8877 c. 2 m above BL-17

BL-19 2257596 5588825 10 cm above BL-21 BL-20 2257596 5588825 c. 2 m below BL-26

BL-21 2257596 5588825 c. 2 m below BL-26

BL-22 2257550 5588771 c. 3 m above BL-26

BL-23 2257550 5588771 c. 7 m above BL-22

BL-24 2257562 5588748 c. 3 m above BL-23

BL-25 2257603 5588745 possibly equivalent to BL-24

BL-26 H41/5764 8881 c. 10 m below BL-27

BL-27 H41/5781 8876 c. 10 m below BL-28

BL-28 H41/5781 8870 probably 1-2 m lower than BL-25

BL-29 H41/5764 8881 c. 10 m below BL-27

BL-30 H41/5795 8831 (H41/f045) c. 10 m above BL-06

BL-31 H41/5800 8823 (H41/f048) possibly equivalent to BL-04 BL-32 H41/5800 8823 (H41/f072) possibly equivalent to BL-05

BL-33 2257911 5588447 (H41/f073) Uncertain precise relationship

12 PALAEO-ELECTRONICA.ORG

Sample Grid reference Stratigraphic notes GL-01 2259089 5590115 estimated 40 m above basement

GL-02 2259079 5590201 approximately equivalent to GL-01

GL-03 H41/5905 9040 c. 10 m above GL-01

GL-04 H41/5898 9040 c. 5 m above GL-01 GL-05 H41/5942 8955 probably equivalent to GL-12

GL-07 2259326 5589510 broadly equivalent to GL-05, est. 60 m above basement

GL-08 2259326 5589510 50 cm above GL-07 GL-09 2259326 5589510 c. 10 m above GL-08

GL-10 2259326 5589510 c. 13 m above GL-09

GL-11 H41/5933 8953 c. 4 m above GL-10 GL-12 2259420 5589504 10-14 m above GL-11

GL-13 2259453 5589724 10-14 m above GL-12

GL-14 2259258 5589617 40 cm above GL-14 GL-15 2259258 5589617 c. 1 m above GL-16

GL-16 2259258 5589617 c. 9 m above GL-19

GL-17 H41/5917 8968 c. 1 m above GL-18, approx equivalent to GL-19 GL-18 H41/5917 8968 broadly equivalent to GL-20

GL-19 H41/5917 8968 approximately equivalent to GL-17

GL-20 H41/5914 8970 Estimated c. 40 m above basement GL-21 H41/5890 9001 c. 3 m above basement

GL-22 2258994 5589951 approximately equivalent to GL-30

GL-23 2258994 5589951 c. 1 m above GL-22 GL-24 2259039 5589957 c. 5 m above GL-23

GL-25 2259082 5589965 c. 10 m above GL-24

GL-26 2259082 5590032 c. 2 m below GL-1 GL-27 H41/5909 8999 c. 4 m below GL-26

GL-28 H41/5917 8990 probably equivalent to GL-27

GL-29 2259266 5589801 broadly equivalent to GL-05 GL-30 2258940 5590041 c. 10 m above basement

GL-31 2258940 5590041 directly overlying GL-30

GL-32 2258940 5590041 directly overlying GL-31

Harliwich-3 G43 211193 Mud within about 2 m above the top of the McPherson coal seam above the high wall of Harliwich’s Coal Mine, Roxburgh

Mata-01 2259921 5585353 (H41/f053) Uncertain precise relationship

Mata-03 2260009 5585216 (H41/f074) Uncertain precise relationship

Mata-06 H41/6000 8514 (H41/f077) Uncertain precise relationship

Mata-18 H41/6006 8493 One of four closely-spaced carbonaceous layers in the left bank of Mata Ck. Mata-18 is the second one downstream.

Mata-23 H41/6003 8490 Carbonaceous mud on right bank of Mata Ck.

13 POLE: MIOCENE LEAF CUTICLE

APPENDIX 2. DETAILS FOR GORE LIGNITE MEASURES, EAST SOUTHLAND GROUP SAMPLES

Sample Drill Hole Depth Grid reference

Sthd-002 d.1024 118.45 m 2186734 5439866

Sthd-004 d.1026 27.62 m 2179606 5441890

Sthd-010 d.1027 168.02 m 2181460 5441941 Sthd-011 d.1027 174.42 m 2181460 5441941

Sthd-012 d.1027 207.25 m 2181460 5441941

Sthd-016 d.1051 4.96 m 2182300 5439100 Sthd-017 d.1051 84.36 m 2182300 5439100

Sthd-018 d.1051 86.00 m 2182300 5439100

Sthd-019 d.1051 86.42 m 2182300 5439100 Sthd-020 d.1051 92.62 m 2182300 5439100

Sthd-022 d.1051 94.41 m 2182300 5439100

Sthd-024 d.1051 105.09 m 2182300 5439100 Sthd-026 d.1051 151.95 m 2182300 5439100

Sthd-027 d.1052 4.80 m 2184396 5438625

Sthd-029 d.1052 9.78m 2184396 5438625 Sthd-030 d.1052 20.21 m 2184396 5438625

Sthd-032 d.1052 49.52 m 2184396 5438625

Sthd-033 d.1052 70.75 m 2184396 5438625 Sthd-034 d.1057 92.84 m 2179645 5439947

Sthd-040 d.1102 35.15 m 2177520 5427926

Sthd-041 d.1102 55.80 m 2177520 5427926 Sthd-043 d.1102 58.00 m 2177520 5427926

Sthd-044 d.1102 58.15 m 2177520 5427926

Sthd-045 d.1102 58.40 m 2177520 5427926

Sthd-046 d.1102 59.80 m 2177520 5427926

Sthd-047 d.1102 60.00 m 2177520 5427926

Sthd-051 d.1105 88.95 m 2183637 5427618

Sthd-054 d.1106 74.35 m 2185817 5427315

Sthd-055 d.1107 3.75 m 2187857 5427096

Sthd-056 d.1108 120.75 m 2173129 5428234

Sthd-058 d.1109 48.50 m 2170584 5428226

Sthd-059 d.1109 200.85 m 2170584 5428226

Sthd-060 d.1109 201.00 m 2170584 5428226

Sthd-067 d.1115 102.29 m 2192600 5450900 Sthd-068 d.1121 102.30 m 2170600 5422800

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Sample Drill Hole Depth Grid reference Sthd-069 d.1121 150.70 m 2170600 5422800

Sthd-072 d.1124 115.45 m 2176557 5419776

Sthd-073 d.1124 157.8 or 160.3 m 2176557 5419776

Sthd-074 d.1124 c. 158.30 m 2176557 5419776 Sthd-076 d.1141 15.10 m 2163781 5408891

Sthd-078 d.1143 56.33 m 2166000 5305600

Sthd-086 d.1246 38.35 m 2188861 5435907 Sthd-087 d.1246 95.13 m 2188861 5435907

Sthd-088 d.1294 80.35 m 2163219 5411606

Sthd-089 d.1295 37.50 m 2164509 5410216 Sthd-090 d.1295 41.50 m 2164509 5410216

Sthd-091 d.1296 30.10 m 2165620 5408988

Sthd-094 d.1298 21.52 m 2162214 5407900 Sthd-095 d.1299 14.35 m 2158369 5407117

Sthd-097 d.1299 51.62 m 2158369 5407117

Sthd-098 d.1299 54.30 m 2158369 5407117 Sthd-099 d.1299 73.10 m 2158369 5407117

Sthd-100 d.1324 22.30 m 2186533 5451035

Sthd-102 d.1324 28.22 m 2186533 5451035 Sthd-106 d.1324 86.75 m 2186533 5451035

Sthd-107 d.1324 88.18 m 2186533 5451035

Sthd-108 d.1324 89.44 m 2186533 5451035 Sthd-109 d.1324 115.79 m 2186533 5451035

Sthd-110 d.1324 124.75 m 2186533 5451035

Sthd-111 d.1324 143.25 m 2186533 5451035

15 POLE: MIOCENE LEAF CUTICLE

APPENDIX 3.

In this publication identified fossil cuticle taxa 4. Stomatal complexes with clearly visible guard are presented in order following the Angiosperm cells, which are distinctly elongate, almost Phylogeny Group (APG 2003): rectangular guard cells, outer stomatal Gnetales ledges and separating walls between guard Gnetaceae cells absent. Gnetales Canellales 4. Stomatal complexes with guard cells often par- Winteraceae tially obscured by an outer stomatal ledge, Laurales separating walls between guard cells Atherospermataceae present, guard cells irregular or rounded. 5. Monimiaceae Proteales 5. Epidermis distinctly granular. Winteraceae Proteaceae 5. Epidermis not distinctly granular. 6. Ranunculales 6. Cuticle glabrous. Monimiaceae Menispermaceae Santalales 6. Trichomes or trichome attachment sites present Santalaceae 7. Fagales 7. Multicelluar trichome attachment sites present Casuarinaceae (trichomes deciduous). Proteaceae Oxalidales 7. Persistent branching trichomes present. Santal- Cunoniaceae-Elaeocarpaceae aceae Sapindales Meliaceae 8. Stomatal complexes anisocytic, ornamented by Sapindaceae prominent ridging. Myrsinaceae Ericales 8. Stomatal complexes not anisocytic. 9. Myrsinaceae 9. Stomatal complexes with clear subsidiary cells. Apiales 10. Griseliniaceae Asterales 9. Stomatal complexes with no clear subsidiary Argophyllaceae cells (anomocytic, although there may be The following key is a general guide for how occasional divisions of contact cells). Sto- the cuticle in this work has been allocated to fami- matal complexes with prominent outer sto- lies. matal ledges. Strasburgeriacae 10. Stomatal complexes with a distinctive outer sto- Key to families matal rim, which is typically ovate, of equal 1. Plant remains found as three-dimensional, four- breadth around the stoma, and under TLM sided articles. Stomatal complexes oriented appears thicker than any other area of cuti- at right angles to long axis (The cuticle is cle. Atherospermataceae very delicate and difficult to prepare.) Gym- 10. No distinct outer stomatal rim as above. 11. 11. nostoma Distinctive form of massively-thickened tri- 1. Remains found as dispersed cuticle and typically chome attachment scars (probably poral) as sheets.2. present. Menispermaceae 2. Raised, thick, peg-like attachment scars of 11. No massively-thickened trichome attachment deciduous trichomes common and ridges of scars as above. 12. cuticle commonly partially projecting over 12. Stomatal complexes typically aligned. Eri- OSL. Argophyllaceae caceae 2. Peg-like attachment scars absent, or OSL not 12. Stomatal complexes randomly oriented 13. obscured by ridges of cuticle. 3. 13. Stomata size large (and with prominent outer 3. Stomatal complexes brachyparacytic 4. stomatal ledges). Grisselinaeaceae 3. Stomatal complexes not brachyparacytic 8. 13. Stomata size medium 14.

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14. Subsidiary cell flanges end abruptly at or over- Group 14. Stomatal complexes prominently cyclo- lap the outline of the guard cells. Sapin- cytic (developmentally tangeticytic) with daceae. prominent outer stomatal ledges 14. Subsidiary cell flanges do not end abruptly or Group 15. Stomatal complexes anomocytic overlap the outline of the stomata 14. Group 16. Cuticle either non-stomatiferous or sto- 15. Outer stomatal ledges so thin and smooth that matal complexes very inconspicuous. they leave an unobstructed view of classic ‘paired kidney’ stomata. giant stomata DESCRIPTIONS (hydathodes), simple trichome attachment scars, and cork-warts may be present. Elae- Gnetalaceae Lindley 1834 ocarpaceae – Cunoniaceae. CUT-Z-ADE 15. Margin of the guard cell pair is indistinct, while Figure 2.1-2.4 the outer stomatal ledge is well-defined but Reference Specimen and locality: SL0093, BL-08. narrow, covering just the inner part of the Referred specimens and occurrence: SL1309, BL- guard cells. The contact zone between guard 32; SB0841, GL-02; SL1740, Sthd-017; SL2434, cells and subsidiary cells is often covered by Sthd-033; SL2076, Sthd-113. very thin cuticle which leads to the appear- ance under TLM of the outer stomatal ledge Stomatal complexes. Stomatal complexes evenly seeming to ‘float’. The pattern of subsidiary spread, isolated, randomly oriented, tetracytic cells is often complex, the result of tangential (sometimes brachyparacytic), developmentally tan- cell division. Meliaceae genticytic or stephanocytic bicylic, with distinct lat- This leaves a large group of taxa which could eral subsidiary cells parallel to the stomatal axis not be identified at all, or identified with reasonable and polar cells at right angles to the axis and which confidence. To deal with this remainder they were may 'enclose' the lateral cells. Sometimes subsid- grouped using morphological characters, but with- iary cells are modified by a tangential division, giv- out any implication that they were taxonomically ing two lateral and two polar cells, size range related. The morphological groups are segregated unimodal. Subsidiary cells (4–6) typically elongate, in this order (i.e. it is to be read sequentially as a or wedge-like, periclinal walls thicker than over nor- dichotomising key): mal epidermal cells, smooth, unornamented. Guard cell pair outline distinctly elongate, often Group 1. Papillae present with flattened poles, like a rounded rectangle, out- Group 2. Stomatal complexes generally aligned lined by a well-defined anticlinal wall, length 27–37 Group 3. Epidermal cells highly sinuous m (medium), at same level as subsidiary cells (exposed on surface), little polar development Group 4. Stomatal complexes surrounded by a between guard cells (guard cells appear as contin- prominent, broad peristomatal ring uous ring). Outer stomatal ledge possibly absent, Group 5. Persistent trichomes present with the visible cuticle lying directly over the guard Group 6. Stomatal complexes paracytic cells, thinner than normal epidermal cells (often broken away), and with a narrowly elliptic pore. Group 7. Stomatal complexes anisocytic Epidermal Cells. Epidermal cell flanges clearly visi- Group 8. Multi cellular trichome attachment scars ble using TLM, normal cells highly variable from present isodiametric to elongate (cells over major veins Group 9. Fine surface ridging or striae around sto- more isodiametric shape than normal epidermal matal complexes cells), distinctly larger than the stomata, anticlinal Group 10. Stomatal complexes in islands walls straight to curved, unbuttressed, smoothly textured, unornamented. Group 11. Outer stomatal ledges prominent or unusually shaped Indumentum. Glabrous. Group 12. Epidermal cells sinuous and stomatal Distinguishing features. Distinguished by the elon- complexes highly networked gate guard cell pair with no division at the polar ends, lack of outer stomatal ledges and subsidiary Group 13. Stomatal complexes typically having two cells which include both typically paracytic forms rings of narrow subsidiary cells as well as polar forms which have formed by tan- gential cut-off.

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Figure 2. 1-8. Fossil and extant Gnetales. 1. CUT-Z-ADE, TLM view showing stomatal complexes (SL0093, scale-bar = 50 µm); 2. CUT-Z-ADE, TLM detail of single stomatal complex (SL0093, scale-bar = 20 µm). Note the distinctive elongate shape of the guard cell pair, with no sign of separating walls, and no sign of an outer stomatal ledge; 3. CUT- Z-ADE, SEM view of outer cuticular surface showing a single stomatal complex (S-1125, scale-bar = 20 µm); 4. CUT- Z-ADE, SEM view of inner cuticular surface showing a single stomatal complex (S-1125, scale-bar = 10 µm); 5. Extant Gnetum microcarpum, TLM view showing stomatal complexes (AQ142225, scale-bar = 50 µm); 6. Extant G. gnemon, TLM detail of single stomatal complex (AQ142124, scale-bar = 20 µm); 7. Extant G. tenuifolium, TLM detail of single stomatal complex (AQ142229, scale-bar = 20 µm); 8. Extant G. latifolium, TLM detail of single stomatal complex (AQ142207, scale-bar = 20 µm).

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Identification. On the basis of its highly distinctive subsidiary cells. Outer stomatal ledge elliptical, stomatal complexes, this taxon is regarded as an thicker than normal epidermal cells, extending over extinct taxon in the Gnetaceae. Extant Gnetum centre of stoma, with a broad, elliptical pore, species (for comparison G. microcarpum (Earl) plugged with cuticular material. Blume, G. gnemon L., G. tenuifolium Ridley, and G. Epidermal Cells. Epidermal cell flanges clearly visi- latifolium Blume are illustrated; Figs 2.5-2.8), have ble using TLM, normal cells highly variable from uniquely-shaped stomata which are almost isodiametric to elongate (cells over veins not distin- rounded rectangles. There is no clear division guished by shape), approximately the same size between the two guard cells and an outer stomatal as the stomata, anticlinal walls curved to wavy, ledge appears to be absent (the cuticle that is unbuttressed, with moderately granular texture, present lies directly over the guard cells). CUT-Z- unornamented. ADE is distinct from Gnetum because it has distinct polar subsidiary cells, which are essentially absent Indumentum. Glabrous. in Gnetum. For further discussion see Pole (2007d) Identification. Winteraceae based on large para- where CUT-Z-GDB from the Early Eocene of Tas- cytic stomata and moderately coarse cuticle. Com- mania is also regarded as Gnetalean. The clearly pare with illustrations of extant Winteraceae related cuticle of the Tasmanian specimens differs (Belliolum haplopus (B. L. Burtt) A.C. Sm., B. burtti- in the walls of the epidermal cells being distinctly anum A. C. Smith (1985), Exospermum stipitatum wavy. (Baillon) van Tieghem (1900) ex Morot, Zygogy- Comments. Gnetum has no fossil record, but Won num balansae Tiegh.; Figs 4.1-4.4). and Renner (2006) have used genetics to infer that the major divergences amongst the extant clades CUT-Z-FJF of the genus date from the Late Oligocene. Figure 3.3-3.6 Reference Specimen and locality: SL2538, GL-02. Magnoliopsida Cronquist 1981 Stomatal complexes. Stomatal complexes evenly Winteraceae Lindley 1830 spread, isolated, randomly oriented, brachypara- Family identification: All extant Winteraceae gen- cytic, size range unimodal. Subsidiary cells with era are characterized by large paracytic stomata periclinal walls of same thickness as normal epi- and a highly granular or ornamented cuticle dermal cells, smooth, ornamented with thick ridges (Bongers 1973). generally parallel to the stomatal axis on either side of the stoma, and at right angles to it at either end. Key to Winteraceae Guard cell pair outline difficult or impossible to see 1. Cuticle ornamented with closely-spaced, thick in TLM view (obscured by surface ornamentation), (5-10 m) discontinuous ridges. CUT-Z-FJF at same level as subsidiary cells, length 38–40 m 1. Cuticle unornamented. 2. (large). Outer stomatal ledge elliptical, thinner than normal epidermal cells, extending over whole 2. Cuticle moderately granular (outline of outer sto- stoma, with a narrowly elliptic pore, plugged with matal ledge clearly visible under TLM) CUT- cuticular matter. Z-FFD Epidermal Cells. Epidermal cell flanges not clearly 2. Cuticle strongly granular (plugging and obscur- visible under TLM because of surface thickenings, ing stomatal outline under TLM). CUT-Z-FJH normal cells elongate (cells over veins not distin- guished by shape), approximately the same size CUT-Z-FFD as the stomata, anticlinal walls curved to wavy, Figure 3.1-3.2 unbuttressed, smoothly textured ornamented with Reference Specimen and locality: SL3140, GL-01. closely-spaced, thick (5-10 m) discontinuous Stomatal complexes. Stomatal complexes evenly ridges. spread, isolated, randomly oriented, brachypara- Indumentum. Glabrous. cytic, size range unimodal. Subsidiary cells elon- Distinguishing features. Distinguished from CUT-Z- gate, periclinal walls of same thickness as normal FJH by the ornamentation of massive cuticular epidermal cells, unornamented. Guard cell pair ridges. outline difficult or impossible to see in TLM view (obscured by outer stomatal ledge), but obviously Identification. Winteraceae based on large para- elliptical, length 45–53 m (large), at same level as cytic stomata and highly ornamented cuticle. Com-

19 POLE: MIOCENE LEAF CUTICLE

Figure 3. 1-8. Fossil Winteraceae: CUT-Z-FFD, CUT-Z-FJF, and CUT-Z-FJH. 1. CUT-Z-FFD. TLM view showing sto- matal complexes (SL3140, scale-bar = 50 µm); 2. CUT-Z-FFD. TLM detail of single stomatal complex (SL3140, scale- bar = 20 µm). The outer stomatal ledge is complex and may involve overlapping ledges; 3. CUT-Z-FJF. TLM view showing stomatal complexes (SL2538, scale-bar = 50 µm). Note the ornamentation of massive ridges; 4. CUT-Z-FJF. TLM detail of single stomatal complex (SL2538, scale-bar = 20 µm); 5. CUT-Z-FJF. SEM view of outer cuticular sur- face showing massive ridging obscuring stomatal complexes (S-1197, scale-bar = 20 µm); 6. CUT-Z-FJF. SEM view of outer cuticular surface showing a single stomatal complex (S-1197, scale-bar = 20 µm); 7. CUT-Z-FJH. TLM view showing stomatal complexes (SL2185, scale-bar = 50 µm). Note the dark material plugging the stomatal apertures; 8. CUT-Z-FJH. TLM detail of single stomatal complex (SL2185, scale-bar = 20 µm). Note the highly granular texture.

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Figure 4. 1-4. Extant Winteraceae. Note the highly granular texture in each example. 1. Belliolum haplopus, TLM detail of single stomatal complex (AQ117672, scale-bar = 20 µm); 2. B. burttianum, TLM detail of single stomatal complex (AQ463392, scale-bar = 20 µm); 3. Exospermum stipitatum, TLM detail of single stomatal complex (AQ391245, scale-bar = 20 µm); 4. Zygogynum balansae, TLM detail of single stomatal complex (AQ391241, scale- bar = 20 µm). pare with illustrations of extant Winteraceae (Figs ric to elongate (cells over veins not distinguished 4.1-4.4). by shape), approximately the same size as the sto- mata, anticlinal walls curved to wavy, unbuttressed, CUT-Z-FJH texture strongly granular, unornamented. Figure 3.7-3.8 Indumentum. Glabrous. Reference Specimen and locality: SL2185, GL-10. Distinguishing features. Distinguished from CUT-Z- Referred specimens and occurrence: SL3163, BL- FJF and CUT-Z-FFD the strongly granular pericli- 33. nal epidermal cell walls. Stomatal complexes. Stomatal stomatal complexes Identification. Winteraceae based on large brachy- evenly spread, isolated, randomly oriented, brac- paracytic stomata and highly granular cuticle. hyparacytic, size range unimodal. Subsidiary cells Compare with illustrations of extant Winteraceae with periclinal walls of same thickness as normal (Figs 4.1-4.4). epidermal cells, texture granular, unornamented. Guard cell pair outline ovate, outer margin Atherospermataceae Brown 1814 obscured under TLM by surface ornamentation, at Family identification: Based on the reference col- same level as subsidiary cells, length 35–45 m lection the Atherospermataceae have a distinctive (large). Outer stomatal ledge elliptical, thicker than outer stomatal rim, which is typically ovate, of normal epidermal cells, extending over whole equal breadth around the stoma, and under TLM stoma with a narrowly elliptic pore, plugged with appears thicker than any other area of cuticle cuticular material. (Doryphora sassafras Endlicher (1841), D. sassa- Epidermal Cells. Epidermal cell flanges somewhat fras, Laurelia novae-zealandiae Cunningham diffuse, normal cells highly variable from isodiamet- (1838), L. novae-zealandiae, D. aromatica (

21 POLE: MIOCENE LEAF CUTICLE

Figure 5. 1-6. Extant Atherospermataceae. 1. Doryphora sassafras, TLM view showing stomatal complexes and a massively-thickened trichome attachment scar (AQ217598, scale-bar = 50 µm); 2. D. sassafras, TLM view showing two stomatal complexes (AQ217598, scale-bar = 20 µm). Note the distinctive outer stomatal rim; 3. Laurelia novae- zealandiae, TLM view showing stomatal complexes and a massively-thickened trichome attachment scar (OPH7027, scale-bar = 50 µm); 4. L. novae-zealandiae, TLM detail of single stomatal complex (OPH7027, scale-bar = 20 µm). Note the distinctive outer stomatal rim; 5. D. aromatica, TLM view showing stomatal complexes (AQ607275, scale-bar = 50 µm); 6. D. aromatica, TLM detail of single stomatal complex (AQ607275, scale-bar = 20 µm). Note the distinctive outer stomatal rim.

F.M.Bailey) Smith (1958), are illustrated as extant nent periclinal rims have been shown on examples; Figs 5.1-5.6 and see also Barrera and Doryphora (Metcalfe (1987). Meza (1992) for the Chilean Laurelia and Laureli- Key to Atherospermataceae opsis). This feature immediately distinguishes the family from the Monimiaceae, within which it has 1. Cuticle glabrous. CUT-Z-CEF sometimes been included. Furthermore, there is 1. Trichome attachment scars present. 2. often a ring of subsidiary cells, and often with mas- sively-thickened trichome attachment scars. Promi- 2. Ornamented by cuticular ridges. CUT-Z-CFA 2. Unornamented. 3.

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Figure 6. 1-4. Fossil Atherospermataceae: CUT-Z-CEF. 1. TLM view showing two stomatal complexes (SL0100, scale-bar = 50 µm); 2. TLM detail of single stomatal complex (SL0100, scale-bar = 20 µm). Note the distinctive outer stomatal rim; 3. SEM view of inner cuticular surface showing a single stomatal complex (S-1112, scale-bar = 20 µm); 4. SEM view of outer cuticular surface showing a single stomatal complex with prominent outer stomatal ledges (S- 1112, scale-bar = 20 µm).

3. Massively thickened trichome attachment sites complexes have more subsidiary cell development, present. CUT-Z-CFC but they are no larger than normal stomatal com- 3. Massively thickened trichome attachment sites plexes). Subsidiary cells (4–5) irregularly shaped, absent. 4. but often elongate radially to the stoma, periclinal walls thicker than over normal epidermal cells, 4. Epidermal cells distinctly smaller than the sto- smooth, unornamented. Guard cell pair outline dis- mata. CUT-Z-ECG tinctly circular, length c. 34 m (medium), at same 4. Epidermal cells of roughly similar size to the sto- level as subsidiary cells (exposed on surface), with mata 5. prominent T-piece thickenings at polar ends. Outer 5. Epidermal cell anticlinal walls markedly sinuous, stomatal ledge subcircular, thicker than normal epi- slightly buttressed, CUT-Z- CGB dermal cells, extending over centre to inner edge of stoma, with a broad, sub-circular pore. 5. Epidermal cell anticlinal walls not sinuous or but- tressed, CUT-Z-JIF Epidermal Cells. Epidermal cell flanges clearly visi- ble using TLM, normal cells elongate (cells over CUT-Z-CEF veins not distinguished by shape), approximately Figure 6 the same size as the stomata, anticlinal walls curved to wavy, unbuttressed, smoothly textured, Reference Specimen and locality: SL0100, BL-08. unornamented. Referred specimens and occurrence: SL2409, BL- Indumentum. Glabrous. 32; SB1392, BL-33; SL3136, GL-02. Distinguishing features. Distinguished from CUT-Z- Stomatal complexes. Stomatal complexes evenly CFC by being glabrous and from CUT-Z-EHI by the spread, isolated, randomly oriented, actinocytic, or subsidiary cells being level with the other epider- staurocytic, size range unimodal, (some stomatal mal cells.

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Figure 7. 1-8. Fossil Atherospermataceae: CUT-Z-CFA. 1. TLM view showing stomatal complexes and a massively- thickened trichome attachment scar (SB1373, scale-bar = 50 µm); 2. TLM detail of single stomatal complex (SB1373, scale-bar = 20 µm). Note the distinctive outer stomatal rim; 3. TLM view showing stomatal complexes (SL2960, scale- bar = 50 µm); 4. TLM detail of single stomatal complex (SL2960, scale-bar = 20 µm). Note the distinctive outer sto- matal rim; 5. TLM view showing stomatal complexes (SL0304, scale-bar = 50 µm); 6. TLM detail of single stomatal complex (SL0304, scale-bar = 20 µm). Note the distinctive outer stomatal rim; 7. SEM view of outer cuticular surface showing a single stomatal complex with prominent outer stomatal ledges surrounded by a discontinuous rim (S-1096, scale-bar = 20 µm); 8. SEM view of inner cuticular surface showing a single stomatal complex. Note granular texture of subsidiary cell periclinal walls compared with epidermal cells (S-1096, scale-bar = 20 µm).

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Figure 8. 1-8. Fossil Atherospermataceae: CUT-Z-CFC, and CUT-Z-CGB. 1. CUT-Z-CFC, TLM view showing sto- matal complexes and a massively-thickened trichome attachment scar (SL0066, scale-bar = 50 µm); 2. CUT-Z-CFC, TLM detail of single stomatal complex (SL0066, scale-bar = 20 µm); 3. CUT-Z-CFC, SEM view of outer cuticular sur- face showing stomatal complexes with prominent outer stomatal ledges (S-1108, scale-bar = 20 µm); 4. CUT-Z-CFC, SEM view of inner cuticular surface showing a single stomatal complex (S-1108, scale-bar = 20 µm); 5. CUT-Z-CGB, TLM view showing stomatal complexes and a massively-thickened trichome attachment scar (SL0088, scale-bar = 50 µm); 6. CUT-Z-CGB, TLM detail of single stomatal complex (SL0088, scale-bar = 20 µm). Note the distinctive outer stomatal rim; 7. CUT-Z-CGB, SEM view of inner cuticular surface showing a single stomatal complex (S-1111, scale- bar = 20 µm); 8. CUT-Z-CGB, SEM view of outer cuticular surface showing a single stomatal complex (S-1111, scale- bar = 20 µm).

25 POLE: MIOCENE LEAF CUTICLE

CUT-Z-CFA opmentally tangenticytic or incomplete Figure 7 stephanocytic bicyclic), development size range Reference Specimen and locality: SB1373, BL-32. unimodal. Subsidiary cells (5–6) irregularly- shaped, periclinal walls of same thickness as nor- Referred specimens and occurrence: SL0289, BL- mal epidermal cells, smooth, unornamented. 01; SL3064, BL-26; SL3151, GL-01; SL2543, GL- Guard cell pair outline circular, outer margin 02; SL2682, GL-04; SL2130, GL-08; SL2157, GL- obscured under TLM by surface ornamentation, at 09; SL2919, GL-22; SL4799, GL-23; SL2960, GL- same level as subsidiary cells, length 23–25 m 25. (medium). Outer stomatal ledge sub circular, cir- Stomatal complexes. Stomatal complexes evenly cumscribed by a ring of very thin cuticle, the ledge spread, isolated, randomly oriented, cyclocytic, itself thicker than normal epidermal cells, extending with all some, or none of the subsidiary cells hav- over centre of stoma, with a broad, sub-circular ing been modified by a tangential division, size pore. range unimodal. Subsidiary cells (4–6) typically Epidermal Cells. Epidermal cell flanges clearly visi- elongate, or wedge-like, often including distinct ble using TLM, normal cells isodiametric (cells over polar cells (at right angles to the stomatal axis) veins not distinguished by shape), distinctly smaller which may 'enclose' the lateral cells, periclinal than the stomata, anticlinal walls straight, unbut- walls of same thickness as normal epidermal cells, tressed, smoothly textured, unornamented. smooth, ornamented with one or two peristomatal ridges and there may be varying degrees of ridges Indumentum. Attachment scars of deciduous tri- which tend to flow around the stoma. Guard cell chomes common, diameter similar or larger in size pair outline ovate, outlined by a well-defined anticli- than a normal epidermal cell. Epidermal cells nal wall, length 27–45 m (large to medium), at around trichome attachment scar (4–11) modified same level as subsidiary cells (exposed on sur- with massive thickening to form a rim and radiating face), with prominent T-piece thickenings at polar walls. ends. Outer stomatal ledge sub circular, thicker Distinguishing features. Distinguished from CUT-Z- than normal epidermal cells, extending over whole CEF by having massively thickened trichome stoma, with a broad, sub-circular pore. attachment sites and from CUT-Z-EHI by the sub- Epidermal Cells. Epidermal cell flanges clearly visi- sidiary cells being level with the other epidermal ble using TLM, normal cells isodiametric (cells over cells. major veins more elongate), distinctly smaller than the stomata, anticlinal walls curved to wavy, unbut- CUT-Z-CGB tressed, smoothly textured, ornamented through- Figure 8.5-8.8 out with short ridges or just in the area of the Reference Specimen and locality: SL0088, BL-08. stomatal complexes. Referred specimens and occurrence: SL0239, BL- Indumentum. Scars of deciduous trichomes 04; SL3281, BL-05; SL3193, BL-32; SL1258, BL- sparse, scattered over venal and non-venal 33; SL1173, GL-01; SL2882, GL-20; SL3214, Sthd- regions, inserted between (8–13, hard to count) 158. epidermal cells modified by tangential divisions to Stomatal complexes. Stomatal complexes evenly form a sub-circular zone of foot cells with massive spread, isolated, randomly oriented, cyclocytic, wall thickening, scar diameter much smaller than size range unimodal. Subsidiary cells (6–7) irregu- normal epidermal cell. larly-shaped, periclinal walls of same thickness as normal epidermal cells, smooth, unornamented. CUT-Z-CFC Guard cell pair outline ovate, outlined by a well- Figure 8.1-8.4 defined anticlinal wall, at same level as subsidiary Reference Specimen and locality: SL0066, Mata- cells, length 30–45 m (medium). Outer stomatal 18. ledge sub circular, thicker than normal epidermal Referred specimens and occurrence: SL0370, cells, extending over whole stoma, with a broad, Mata-03. sub-circular pore. Stomatal complexes. Stomatal complexes evenly Epidermal Cells. Epidermal cell flanges clearly visi- spread, isolated, randomly oriented, cyclocytic, ble using TLM, normal cells highly variable from with all some, or none of the subsidiary cells hav- isodiametric to elongate (cells over major veins ing been modified by a tangential division (devel- more elongate), approximately the same size as

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Figure 9. 1-8. Fossil Atherospermataceae: CUT-Z-JIF and CUT-Z-ECG. 1. CUT-Z-JIF, TLM view showing stomatal complexes and a massively-thickened trichome attachment scar (SB0851, scale-bar = 50 µm); 2. CUT-Z-JIF, TLM detail of single stomatal complex (SB0851, scale-bar = 20 µm); 3. CUT-Z-JIF, SEM view of outer cuticular surface showing stomatal complexes and (just left of centre) a trichome attachment scar (S-319, scale-bar = 0.1 mm); 4. CUT-Z-JIF, SEM view of inner cuticular surface showing a single stomatal complex (S-319, scale-bar = 10 µm); 5. CUT-Z-ECG, TLM view showing stomatal complexes (SL1954, scale-bar = 50 µm); 6. CUT-Z-ECG, TLM detail of sin- gle stomatal complex (SL1954, scale-bar = 20 µm); 7. CUT-Z-ECG, SEM view of inner cuticular surface showing a single stomatal complex (S-1073, scale-bar = 20 µm); 8. CUT-Z-ECG, SEM view of outer cuticular surface showing a single stomatal complex (S-1073, scale-bar = 10 µm).

27 POLE: MIOCENE LEAF CUTICLE the stomata, anticlinal walls markedly sinuous, CUT-Z-ECG slightly buttressed, smoothly textured, unorna- Figure 9.5-9.8 mented. Reference Specimen and locality: SL1954, Sthd- Indumentum. Attachment scars of deciduous tri- 051. chomes common, inserted between (5–7) epider- Referred specimens and occurrence: SL3042, BL- mal cells modified into radially elongate foot cells, 22; SL1563, Sthd-004; SL1730, Sthd-017; SL1716, scar poral diameter much smaller than a normal Sthd-018; SL1706, Sthd-019; SL1776, Sthd-022; epidermal cell. SL1779, Sthd-030; SL1937, Sthd-059; SL1894, Distinguishing features. Distinguished from CUT-Z- Sthd-073; SL1906, Sthd-074; SL1833, Sthd-095. CFC by the sinuous and slightly buttressed walls. Stomatal complexes. Stomatal complexes evenly spread, isolated, randomly oriented, laterocyclic, CUT-Z-JIF all some, or none of the subsidiary cells may have Figure 9.1-9.4 been modified by a tangential division (develop- Reference Specimen and locality: SB0851, Mata- mentally incomplete stephanocytic bicyclic), size 03. range unimodal (but with wide range of stomatal Referred specimens and occurrence: SL1354, BL- size). Subsidiary cells (7–8) irregularly-shaped, 05; SL3166, BL-33; SL2210, GL-12; SL1060, periclinal walls thicker than over normal epidermal Mata-01. cells, smooth, unornamented. Guard cell pair out- line ovate, outlined by a well-defined anticlinal wall, Stomatal complexes. Stomatal complexes in are- at same level as subsidiary cells (exposed on sur- oles, isolated, randomly oriented, anomocytic or face), length 38–43 m (large), with prominent T- with occasional thin subsidiary cells produced by piece thickenings at polar ends. Outer stomatal tangential divisions (developmentally incomplete ledge sub circular, thicker than normal epidermal stephanocytic bicyclic), size range unimodal. Con- cells, extending over most of the stoma, with a tact cells (4–6) irregularly-shaped, periclinal walls broad, sub-circular pore. thicker than over normal epidermal cells, smooth, ornamentation sometimes a single discontinuous Epidermal Cells. Epidermal cell flanges clearly visi- ridge around the outer stomatal ledge (periclinal ble using TLM, normal cells isodiametric (cells over wall thickness near this rim is much thinner than major veins vaguely or inconsistently more elon- over epidermal cells). Guard cell pair outline circu- gate), approximately the same size as the stomata, lar, or ovate, outlined by a well-defined anticlinal anticlinal walls straight, unbuttressed, smoothly wall, at same level as subsidiary cells (exposed on textured, unornamented. surface), length 37–45 m (large), with prominent Indumentum. Attachment scars of deciduous tri- polar rods. Outer stomatal ledge sub circular, much chomes common, probably restricted to regions thicker than normal epidermal cells, extending over over veins (unclear), inserted between epidermal whole stoma, with a broad, sub-circular pore. cells modified only slightly, forming a thickened rim, Epidermal Cells. Epidermal cell flanges clearly visi- scar diameter similar in size to a normal epidermal ble using TLM, normal cells isodiametric (cells over cell. both major and fine venation more elongate), Distinguishing features. Differing from the other approximately the same size as the stomata, anti- taxa in the group in having very thick outer sto- clinal walls curved to wavy, unbuttressed, smoothly matal ledges, and stomatal complexes with clear textured, unornamented. evidence of tangential divisions, and epidermal Indumentum. Attachment scars of deciduous tri- cells distinctly smaller than the stomata. chomes common, scattered over venal and non- venal regions, inserted between (5–7) epidermal Monimiaceae Jussieu 1809 cells modified by massive wall thickening to form a Key to Monimiaceae rim with radiating walls, scar diameter much smaller than a normal epidermal cell. 1. Unornamented. CUT-Z-DDG Distinctive features. Comparable to CUT-Z-CGB 1. Ornamented with ridges. 2. but without the buttressed and sinuous epidermal 2. Ornamented with widely-spaced short ridges, cell walls. outer stomatal ledge narrow. CUT-Z-DDH

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Figure 10. 1-8. Fossil and extant Monimiaceae. 1. CUT-Z-DDG, TLM view showing stomatal complexes (SB1388, scale-bar = 50 µm); 2. CUT-Z-DDG, TLM detail of single stomatal complex (SB1388, scale-bar = 20 µm); 3. CUT-Z- DDG, SEM view of inner cuticular surface showing two stomatal complexes (S-0327, scale-bar = 10 µm); 4. CUT-Z- DDG, SEM view of outer cuticular surface showing several stomatal complexes (S-0327, scale-bar = 0.05 mm); 5. Extant , TLM view showing stomatal complexes (OPH2624, scale-bar = 50 µm); 6. Extant H. arborea, TLM detail of single stomatal complex (OPH2624, scale-bar = 20 µm); 7. Extant H. dorstenioides, TLM view showing stomatal complexes (AQ063921, scale-bar = 50 µm); 8. Extant H. dorstenioides, TLM detail of single sto- matal complex (AQ063921, scale-bar = 20 µm).

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Figure 11. 1-8. Fossil and extant Monimiaceae: CUT-Z-DDH and CUT-Z-JIJ. 1. CUT-Z-DDH, TLM view showing sto- matal complexes (SB1351, scale-bar = 50 µm); 2. CUT-Z-DDH, TLM detail of single stomatal complex (SB1351, scale-bar = 20 µm). Note the highly granular texture; 3. CUT-Z-DDH, SEM view of inner cuticular surface showing a single stomatal complex. Note highly granular texture (S-1227, scale-bar = 20 µm); 4. CUT-Z-DDH, SEM view of outer cuticular surface showing distinctly narrow outer stomatal ledges (S-1227, scale-bar = 10 µm); 5. Extant , TLM view (AQ063593, scale-bar = 50 µm); 6. Extant H. angustifolia, TLM view (AQ063593, scale-bar = 20 µm); 7. CUT-Z-JIJ, TLM view showing two stomatal complexes (SB1329, scale-bar = 50 µm); 8. CUT-Z-JIJ, TLM detail of single stomatal complex (SB1329, scale-bar = 20 µm).

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2. Ornamented with prominent, closely-spaced thinner than normal epidermal cells, extending over long ridges, outer stomatal ledge prominent. inner edge of stoma. CUT-Z-JIJ Epidermal Cells. Epidermal cell flanges somewhat diffuse, normal cells highly variable from isodiamet- CUT-Z-DDG (Hedycarya) ric to elongate (cells over veins not distinguished Figure 10.1-10.4 by shape), distinctly larger than the stomata, anti- Reference Specimen and locality: SB1388, BL-33. clinal walls straight, unbuttressed, inner surface Stomatal complexes. Stomatal complexes evenly texture granular, outer surface mostly very smooth spread, isolated, randomly oriented, brachypara- but with some ornamentation of short ridges per- cytic or amphibrachyparacytic, size range unimo- pendicular or parallel to some stomata. dal. Subsidiary cells typically elongate, tangential Indumentum. Glabrous. to the guard cells, periclinal walls of same thick- Distinguishing features. Distinguished by the rect- ness as normal epidermal cells, smooth, unorna- angular outline of the stomatal complexes, the mented. Guard cell pair outline elongate, outlined granular texture of the epidermal periclinal walls, by a well-defined anticlinal wall, at same level as the ridges of cuticle running at right angles to the subsidiary cells (exposed on surface), length 20– stomatal pore on some stomatal complexes, and 25 m (medium). Outer stomatal ledge elliptical, the obscure contact between guard cells and sub- same thickness as normal epidermal cells, extend- sidiary cells. CUT-Z-JIJ differs by having distinct ing over inner edge of stoma, with a narrowly ellip- and broad outer stomatal ledges. tic pore. Identification. The similarity to extant Hedycarya Epidermal Cells. Epidermal cell flanges clearly visi- angustifolia A. Cunningham (1838) (AQ063593) is ble using TLM, normal cells highly variable from taken to indicate the genus (Figs 11.5-11.6). isodiametric to elongate (cells over veins not distin- guished by shape), distinctly larger than the sto- CUT-Z-JIJ mata, anticlinal walls curved to wavy, unbuttressed, Figure 11.7-11.8 inner surfaces slightly granular, outer surfaces smooth and undulose, unornamented (though may Reference Specimen and locality: SB1329, BL-32. have some faint ridging). Stomatal complexes. Stomatal complexes evenly Indumentum. Glabrous. spread, isolated, randomly oriented, brachypara- cytic, size range unimodal. Subsidiary cells with Identification. The similarity to extant Hedycarya periclinal walls of same thickness as normal epi- arborea Forster, J. R., and Forster, G. (1776) and dermal cells, granular, ornamented with ridges pro- H. dorstenoides A. Gray (1866) (Figs 10.5-10.8) is jecting at right angles to the stomatal axis. Guard taken to indicate the genus. cell pair outline circular, or ovate, not outlined by a clear anticlinal wall, at same level as subsidiary CUT-Z-DDH (Hedycarya sp.) cells (exposed on surface), length 26–30 m Figure 11.1-11.4 (medium). Outer stomatal ledge elliptical, thicker Reference Specimen and locality: SB1351, BL-32. than normal epidermal cells, extending over whole Referred specimens and occurrence: SL2105, GL- stoma, with a narrowly elliptic pore. 07; SL2145, GL-08. Epidermal Cells. Epidermal cell flanges not clearly CUT-L-DDH, SL1193, GL-02; SL2927, GL-22; visible under TLM because of surface thickenings, SL3293, Mata-03. normal cells elongate, cells over veins not distin- guished by shape, anticlinal walls probably wavy, Stomatal complexes. Stomatal stomatal complexes unbuttressed, inner surfaces coarsely granular, evenly spread, isolated, randomly oriented, brac- outer surfaces ornamented with bands of promi- hyparacytic, size range unimodal. Subsidiary cells nent, irregular ridges parallel to the stomata and with periclinal walls thinner than that over normal radiating from trichomes. epidermal cells, granular, unornamented normally, or with short ridges perpendicular to the stomatal Indumentum. Glabrous. axis. Guard cell pair outline elongate, not outlined Distinguishing features. Distinguished by the brac- by a clear anticlinal wall, at same level as subsid- hyparacytic stomatal complexes with broad and iary cells (exposed on surface), length 25–30 m distinct outer stomatal ledges in combination with (medium). Outer stomatal ledge narrow-elliptical, highly granular cuticle. Specimen SL2647, which

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Figure 12. 1-8. Fossil Proteaceae: CUT-P-EDA (Placospermum sp.) and CUT-P- EFB (Lomatia sp.). 1. CUT-P-EDA, TLM view showing aligned stomatal complexes (SL1867, scale-bar = 100 µm); 2. CUT-P-EDA, TLM detail of a single stomatal complex (SL1867, scale-bar = 20 µm). Note very narrow outer stomatal ledge; 3. CUT-P-EDA, SEM view of outer cuticular surface showing a single stomatal complex (S-1037, scale-bar = 20 µm); 4. CUT-P-EDA, SEM view of inner cuticular surface showing a single stomatal complex (S-1037, scale-bar = 20 µm); 5. CUT-P- EFB, TLM view showing stomatal complexes (SL1828, scale-bar = 50 µm); 6. CUT-P- EFB, TLM detail of single stomatal complex (SL1828, scale-bar = 20 µm); 7. CUT-P- EFB, SEM view of inner cuticular surface showing two stomatal complexes. Note finely granular texture (S-1106, scale-bar = 20 µm); 8. CUT-P- EFB, SEM view of outer cuticular surface showing two stomatal complexes with prominent outer stomatal ledges, and striae over the subsidiary cells (S-1106, scale-bar = 20 µm).

32 PALAEO-ELECTRONICA.ORG lacks the strong surficial ornamental of ridges Epidermal Cells. Epidermal cell flanges clearly visi- found in the Reference Specimen, is included in ble using TLM, normal cells isodiametric (cells over this taxon. It comes from the same sample and veins not distinguished by shape), approximately agrees in the other major characters. CUT-Z-DDH the same size as the stomata; anticlinal walls differs in not having the broad outer stomatal markedly sinuous; buttressed; smoothly textured, ledges and narrow, slit-like stomatal apertures. unornamented. Indumentum. Glabrous. Proteaceae Jussieu 1789 Non-stomatal surface. Epidermal cells highly vari- Family identification: The majority of Proteaceae able from isodiametric to elongate; sinuous; cells have brachyparacytic stomatal complexes. They over veins not distinguished; trichomes absent. often have trichome scars associated with one to several epidermal cells (Lange 1978; Carpenter Identification. There are no multicelled trichome 1994). attachment scars which would make a stronger case for identification as Proteaceae. However, the Key to Proteaceae morphology is comparable with Placospermum 1. Stomatal complexes aligned. 2. coriaceum (White and Francis 1924) (the sole member of the genus, and which also lacks or has 1. Stomatal complexes randomly oriented. 3. very rare trichome attachment scars) in its gener- 2. Outer stomatal ledge narrowly elliptic, epidermal ally aligned stomatal complexes, a strikingly dis- cell anticlinal walls markedly sinuous, but- tinctive elongate stomatal pore (cf. Carpenter tressed. CUT-P-EDA 1994, figs 5-7) and sinuous epidermal cell walls. 2. Outer stomatal ledge subcircular, epidermal cell There are obvious differences in that the epidermal anticlinal walls straight, unbuttressed. CUT- cells of the fossil are also buttressed, and the sto- P-FEE matal complexes of the extant species are often linked by large epidermal cells. The fossil is sug- 3. Cuticle glabrous. CUT-P-EFB gested to be an extinct Placospermum. 3. Attachment scars of trichomes present. 4. 4. Trichome attachment scar with a thickened rim, CUT-P-EFB (Lomatia sp.) subsidiary cells ornamented with many fine Figure 12.5-12.8 ridges. CUT-P-EHA Reference Specimen and locality: SL1828, Sthd- 4. Trichome attachment scar thickened into a 095. raised platform, subsidiary cells ornamented Referred specimens and occurrence: SL1556, with 2-3 ridges. CUT-P-EHJ Sthd-004; SL1766, Sthd-011; SL1843, Sthd-086. Stomatal complexes. Stomatal complexes evenly CUT-P-EDA (Placospermum sp.) spread, isolated, randomly oriented, brachypara- Figure 12.1-12.4 cytic, size range unimodal. Subsidiary cells isodia- Reference Specimen and locality: SL1867, Sthd- metric, periclinal walls of same thickness as normal 089. epidermal cells, smooth, ornamented with 3–4 Stomatal complexes. Stomatal distribution over leaf ridges parallel with the stomatal axis, and slightly surfaces hypostomatic, stomatal complexes evenly longer than the stoma. Guard cell pair outline circu- spread, isolated, aligned more or less parallel to lar, outlined by a well-defined anticlinal wall, at long axis of leaf, brachyparacytic, size range uni- same level as subsidiary cells (exposed on sur- modal. Subsidiary cells elongate parallel to sto- face), length 18–23 m (medium), with prominent T- matal axis, with periclinal walls thinner than that piece thickenings at polar ends. Outer stomatal over normal epidermal cells, smooth, unorna- ledge sub circular, same thickness as normal epi- mented. Guard cell pair outline narrow and elon- dermal cells, extending over whole stoma, with a gate, outlined by a well-defined anticlinal wall, at broad elliptical or sub-circular pore. same level as subsidiary cells (exposed on sur- Epidermal Cells. Epidermal cell flanges clearly visi- face). Outer stomatal ledge narrowly elliptic, ridge- ble using TLM, normal cells isodiametric (cells over like, much thinner than normal epidermal cells major veins more elongate), approximately the (often broken away), extending over inner edge of same size as the stomata, anticlinal walls sinuous, stoma, with a narrowly elliptic pore. slightly buttressed, inner surfaces slightly granular,

33 POLE: MIOCENE LEAF CUTICLE

Figure 13. 1-8. Fossil Proteaceae: CUT-P-EHA and CUT-P-EHJ. 1. CUT-P-EHA, TLM view showing stomatal com- plexes and (upper left) a multi-cellular trichome attachment scar. Note how epidermal cells are partially obscuring parts of some stomatal complexes (SL1680, scale-bar = 50 µm); 2. CUT-P-EHA, TLM detail of single stomatal com- plex (SL1680, scale-bar = 20 µm); 3. CUT-P-EHA, SEM view of inner cuticular surface showing a single stomatal complex (S-1056, scale-bar = 20 µm); 4. CUT-P-EHA, SEM view of outer cuticular surface showing a single stomatal complex with striae over the subsidiary cells (S-1056, scale-bar = 20 µm); 5. CUT-P-EHJ, TLM view showing sto- matal complexes and multi-cellular trichome attachment scars (SL1682, scale-bar = 50 µm); 6. CUT-P-EHJ, TLM detail of single stomatal complex and two multi-cellular trichome attachment scars (SL1682, scale-bar = 20 µm); 7. CUT-P-EHJ, SEM view of inner cuticular surface showing stomatal complexes and (top right) a trichome attachment scar (S-1057, scale-bar = 20 µm); 8. CUT-P-EHJ, SEM view of outer cuticular surface showing stomatal complexes and (upper left) a trichome attachment scar (S-1057, scale-bar = 20 µm).

34 PALAEO-ELECTRONICA.ORG

outer surfaces smooth and undulose, unorna- mostly 1-2 epidermal cells and lack the central mented. pore). Indumentum. Glabrous. CUT-P-EHJ Identification. There are no multicelled trichome Figure 13.5-13.8 attachment scars which would make a stronger case for identification as Proteaceae. In addition, Reference Specimen and locality: SL1682, Sthd- the subsidiary cells are rather large, while in most 033. extant Proteaceae they are relatively narrow. How- Referred specimens and occurrence: SL0291, BL- ever, there is a general similarity with some extant 01; SL2433, Sthd-051; SL2420, Sthd-078; SL2022, species, such as Lomatia fraxinifolia F. Muell. ex Sthd-087; SL1651, Sthd-110; SL3298. Bentham (1870) (cf. Carpenter 1994, figs 65-66) in Stomatal complexes. Stomatal complexes evenly which trichome attachment scars may also be rare spread, isolated, randomly oriented, brachypara- or absent. cytic, size range unimodal. Subsidiary cells isodia- metric, periclinal walls of same thickness as normal CUT-P-EHA epidermal cells, smooth, ornamented with 3–4 Figure 13.1-13.4 ridges parallel with, and on either side of the Reference Specimen and locality: SL1680, Sthd- stoma. Guard cell pair outline ovate, outlined by a 033. well-defined anticlinal wall, at same level as sub- Stomatal complexes. Stomatal complexes evenly sidiary cells (exposed on surface), length 20–22 m spread, isolated, randomly oriented, brachypara- (medium). Outer stomatal ledge sub circular, thin- cytic, size range unimodal. Subsidiary cells isodia- ner than normal epidermal cells, extending over metric, periclinal walls of same thickness as normal whole stoma, with an elliptical pore. epidermal cells, ornamented with many fine ridges Epidermal Cells. Epidermal cell flanges clearly visi- parallel with the stoma. Guard cell pair outline ble using TLM, normal cells isodiametric (cells over ovate, not outlined by a clear anticlinal wall, at veins not distinguished by shape), approximately same level as subsidiary cells (exposed on surface the same size as the stomata, anticlinal walls although sometimes partially obscured by adjacent markedly sinuous, unbuttressed, inner surfaces epidermal cells), length 30–37 m (medium), little smooth, outer surfaces smooth and irregularly polar development between guard cells (guard ridged with bands of irregular ridges parallel to sto- cells appear as continuous ring). Outer stomatal matal complexes and radiating from trichomes. ledge sub circular, thinner than normal epidermal Indumentum. Attachment scars of deciduous tri- cells, extending over centre of stoma, with a broad, chomes common, inserted over several (3–4) epi- sub-circular pore. dermal cells modified to form a thick, raised circular Epidermal Cells. Epidermal cell flanges clearly visi- platform, on top of which sits a smooth, thick hollow ble using TLM, normal cells isodiametric (cells over collar, scattered over venal and non-venal regions veins not distinguished by shape), approximately but slightly more common over veins. Pairs of the same size as the stomata, anticlinal walls scars may be joined by a narrow extension of the straight, unbuttressed, smoothly textured, unorna- platform. Scar diameter similar in size to a normal mented. epidermal cell. Indumentum. Attachment scars of deciduous tri- Identification. General morphology suggests the chomes common, scattered over venal and non- taxon has affinities with Tribe Macadamieae venal regions, inserted over several (2–5) modified of Weston and Barker (2006) (Carpenter 1994). epidermal cells selectively thickened to form a The surface ridging, more prominent on the subsid- thickened rim, with a prominent central pore. Scar iary cells and the morphology of the trichome scars diameter similar in size to a normal epidermal cell. is similar to Sleumerodendron, Hicksbeachia and Identification. General morphology suggests the Turrillia vitensis (Turrill) A.C. Smith (1985) (or Ker- taxon has affinities with Tribe Macadamieae madecia vitensis Turrill 1915). of Weston and Barker (2006) (see illustrations in Carpenter 1994). The surface ridging restricted to CUT-P-FEE the subsidiary cells is similar to Carnarvonia Figure 14 although the morphology of the trichome scar is dif- Reference Specimen and locality: SL3221, Sthd- ferent (in Carnarvonia they are associated with 166.

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Figure 14. 1-4. Fossil Proteaceae: CUT-P-FEE. 1. TLM view showing stomatal complexes and a trichome attachment scar over a single basal cell at upper left (SL3221, scale-bar = 50 µm); 2. TLM detail of single stomatal complex. Note the characteristic round outline and strong polar rod thickenings (SL3221, scale-bar = 20 µm); 3. SEM view of outer cuticular surface showing a single stomatal complex with prominent outer stomatal ledges (S-1363, scale-bar = 10 µm); 4. SEM view of inner cuticular surface showing two stomatal complexes (S-1363, scale-bar = 10 µm).

Stomatal complexes. Stomatal complexes in are- Identification. Identification with the Proteaceae oles, isolated, generally aligned, brachyparacytic, seems clear, although further identification is size range unimodal. Subsidiary cells narrow, peri- uncertain. The generally aligned stomata and tri- clinal walls of same thickness as normal epidermal chome attachment scars over only 1-2 epidermal cells, unornamented. Guard cell pair outline difficult cells are the most important characters and sug- or impossible to see in TLM view, outer margin gest affinities lie with Tribe Embothrieae of Weston obscured under TLM by surface ornamentation, at and Barker (2006). same level as subsidiary cells, length 28–35 m (medium), with strong polar rod thickenings. Outer Menispermaceae Jussieu 1789 stomatal ledge prominent, circular to sub circular, Family identification: Based on the reference col- thicker than normal epidermal cells, extending over lection the Menispermanceae typically have dis- whole stoma, with a broad, sub-circular pore. tinct robust and very densely staining trichome Epidermal Cells. Epidermal cell flanges clearly visi- attachment scars, although some species are gla- ble using TLM, normal cells isodiametric (cells over brous. They also tend to have both guard and sub- major veins more elongate), approximately the sidiary cells with cuticle thinner than that over same size as the stomata, anticlinal walls straight, normal epidermal cells. Hill (1989) described a fos- unbuttressed, smoothly textured, unornamented. sil genus of the family, Menispermaphyllum, Indumentum. Attachment scars of deciduous tri- although it is unclear how this genus differs from chomes common, but not noted over veins. posi- some extant genera, for instance the Legnephora tioned over 1–2 epidermal cells modified to form an with which it was compared. Hill compared the cuti- irregular platform (by thickening of the periclinal cle of his Eocene M, tomentosum with extant L. walls) which contains the trichome scar. Scar diam- moorei (F.Muell.) Miers 1867, but the papillate cuti- eter similar in size to a normal epidermal cell. cle of L. moorei in his fig. 2E looks nothing like any L. moorei specimen prepared for this database

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Figure 15. 1-8. Fossil and extant Menispermaceae. 1. CUT-Z-FCA, TLM view showing stomatal complexes (with cuti- cle over guard cell and subsidiary cells thinner than over normal epidermal cells) and very densely staining trichome attachment scars (SL2877, scale-bar = 50 µm); 2. CUT-Z-FCA, TLM view showing two stomatal complexes. Note the ovate outline and each has a distinct lateral subsidiary cell on the left hand side (SL2877, scale-bar = 20 µm); 3. CUT- Z-FCA, SEM view of inner cuticular surface showing two stomatal complexes (S-1384, scale-bar = 10 µm); 4. CUT-Z- FCA, SEM view of outer cuticular surface showing a single stomatal complex (S-1384, scale-bar = 10 µm); 5. CUT-Z- FEC, TLM view showing stomatal complexes (the lower of a pair is arrowed) and densely staining trichome attach- ment scars. Note the cuticle over guard cell and subsidiary cells is thinner than over normal epidermal cells (SB1321, scale-bar = 50 µm); 6. CUT-Z-FEC, TLM detail of single stomatal complex, note ovate outline (SB1321, scale-bar = 20 µm); 7. Extant Pleiogyne australis, TLM view showing stomatal complexes (with cuticle over guard cell and subsid- iary cells thinner than over normal epidermal cells) and densely staining trichome attachment scars (AQ568296, scale-bar = 50 µm); 8. Extant P. australis, TLM view showing two stomatal complexes. Note the distinct lateral subsid- iary cell on the left of the left hand complex, and lower side of the right hand complex (AQ568296, scale-bar = 20 µm).

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Figure 16. 1-8. Fossil and extant Notothixos sp. (Viscaceae). 1. CUT-Z-FGJ, TLM view showing stomatal complexes and branching trichomes (SL1202, scale-bar = 50 µm); 2. CUT-Z-FGJ, TLM detail of single stomatal complex (SL1202, scale-bar = 20 µm); 3. CUT-Z-FGJ, TLM detail of single stomatal complex (SL1202, scale-bar = 20 µm); 4. CUT-Z-FGJ, SEM view of inner cuticular surface showing a single stomatal complex. Note W-shaped depressions at either end of guard cell cuticle which reflect where the guard cells turned sharply outwards (S-1379, scale-bar = 10 µm); 5. CUT-Z-FGJ, SEM view of outer cuticular surface showing a depressed stomatal complex in the centre. The outer stomatal ledges do not project above the surrounding epidermal cells. Branched trichomes are to right and below. (S-1205, scale-bar = 20 µm); 6. CUT-Z-FGJ, SEM view of outer cuticular surface showing many branched tri- chomes (S-1205, scale-bar = 100 µm); 7. Extant Notothixos subaureus, SEM view of inner surface showing a single stomatal complex (AQ609212, scale-bar = 10 µm); 8. Extant N. subaureus, SEM view of outer surface showing branched trichomes and a rare instance of unobscured stomatal complexes (AQ609212, scale-bar = 20 µm).

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(which are not papillate), nor any other Menisper- iary cells, and the scattered presence of elongate maceae in the database. lateral subsidiary cells (Figs 15.5-15.6). Key to Menispermaceae CUT-Z-FEC 1. Epidermal cell anticlinal walls curved to wavy, Figure 15.5-15.6 cuticle over stomatal complexes much thin- Reference Specimen and locality: SB1321, BL-32. ner than normal epidermal cells (often bro- ken away). CUT-Z-FCA Stomatal complexes. Stomatal complexes in are- oles, isolated, randomly oriented, anisocytic, size 1. Epidermal cell anticlinal walls sinuous, cuticle range unimodal. Subsidiary cells (3–4) isodiamet- over stomatal complexes thinner than normal ric, with periclinal walls thinner than normal epider- epidermal cells. CUT-Z-FEC mal cells, granular, unornamented. Guard cell pair outline ovate, outlined by a well-defined anticlinal CUT-Z-FCA wall, at same level as subsidiary cells (exposed on Figure 15.1-15.4 surface), length 22–23 m (medium). Outer stomatal Reference Specimen and locality: SL2877, GL-20. ledge sub circular, thinner than normal epidermal Stomatal complexes. Stomatal complexes evenly cells, extending over whole stoma, with an elliptical spread, isolated, randomly oriented, cyclocytic, pore. size range unimodal. Subsidiary cells (5–6) irregu- Epidermal Cells. Epidermal cell flanges clearly visi- larly shaped, but often with apparently distinct elon- ble using TLM, normal cells elongate (cells over gate lateral or polar cells, periclinal walls thinner veins not distinguished by shape), distinctly larger than normal epidermal cells, smooth, unorna- than the stomata, anticlinal walls sinuous and mented. Guard cell pair outline ovate, outlined by a sometimes rather sharply so, unbuttressed, texture well-defined anticlinal wall, at same level as sub- granular, unornamented. sidiary cells (exposed on surface), length 20–23 m Indumentum. Bases of thick (densely staining), (medium), some wall development between guard probably poral (with a very narrow pore) trichomes cells, and sometimes development of T-piece thick- common, scattered over venal and non-venal enings at the poles. Outer stomatal ledge sub cir- regions (but slightly more common over veins), the cular, much thinner than normal epidermal cells bases persist, but in all cases the trichome has (often broken away), extending over whole stoma, broken off near base, perhaps along a predefined with an elliptical pore. zone of weakness, base diameter similar in size to Epidermal Cells. Epidermal cell flanges clearly visi- a normal epidermal cell. Epidermal cells around tri- ble using TLM, normal cells highly variable from chome attachment scar (4–6) unmodified. isodiametric to elongate (cells over major veins Identification. Compare with extant Pleiogyne aus- more elongate), approximately the same size as tralis (Figs 15.7-15.8) in terms of very thick tri- the stomata, anticlinal walls curved to wavy, unbut- chome attachment scars, shape of stoma, thin tressed, smoothly textured, unornamented. guard and subsidiary cell cuticle. However, it does Distinguishing features. The most distinct charac- not have the occasional elongate subsidiary cells. ter is the massively-thickened trichome attachment scar.Indumentum. Bases of thick (densely stain- Santalaceae Brown 1814 ing), probably poral (with a very narrow pore) tri- CUT-Z-FGJ (Notothixos sp.) chomes common, scattered over venal and non- Figure 16.1-16.6 venal regions, the bases persist, but in all cases Reference Specimen and locality: SL1202, GL-01. the trichome has broken off near base, perhaps along a predefined zone of weakness, scar diame- Stomatal complexes. Stomatal complexes evenly ter similar in size to a normal epidermal cell. Epi- spread, isolated, randomly oriented, brachypara- dermal cells around trichome attachment scar cytic, size range unimodal (but with wide range of modified into (6–9) radially elongate foot cells, with stomatal size). Subsidiary cells length 32–40 m no distinct thickening of walls. (large), or (medium), periclinal walls thinner than normal epidermal cells. Guard cell pair very nar- Identification. Broadly comparable with extant row, poles sharply reflexed outwards, overarched Pleiogyne australis Endlicher (1843) with respect by the subsidiary cells (not exposed on the sur- to darkly staining trichome attachment scars, ovate face). shape of stoma, thin cuticle over guard and subsid-

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Figure 17. 1-4. Fossil Gymnostoma sp. 1. TLM view of stomatal rows. Note brachyparacytic stomatal complexes ori- ented perpendicular to the long axis of the article (SL2427, scale-bar = 50 µm); 2. SEM view of outer cuticular surface showing stomatal complexes (S-1058, scale-bar = 20 µm); 3. SEM view of inner cuticular surface showing stomatal complexes (S-1058, scale-bar = 20 µm); 4. SEM view of article showing node with whorl of reduced leaves (arrowed) (S-1023, scale-bar = 200 µm).

Epidermal Cells. Epidermal cell flanges clearly visi- Casuarinaceae Brown 1814 ble using TLM, normal cells highly variable from Gymnostoma Johnson 1980 isodiametric to elongate (cells over veins not distin- Gymnostoma sp. guished by shape), approximately the same size Figure 17.1-17.4 as the stomata, anticlinal walls straight, unbut- Reference specimen and locality: SL2427, Sthd- tressed, smoothly textured, unornamented. 055. Indumentum. Hirsute, densely covered with stellate Referred specimens and occurrence: SL3232, GL- trichomes, inserted over, or the outgrowth of, a sin- 30; SL2705, Sthd-046; SL2702, Sthd-047, SL2703, gle modified epidermal cell, similar in size to a nor- Sthd-094; SL2706, Sthd-097; SL2704, Sthd-099. mal epidermal cell. Description. Branchlets with four-sided articles, Distinguishing features. Distinguished by the approximately square in cross-section, with four unique combination of brachyparacytic stomatal highly reduced leaves in a whorl at each junction. complexes (a pair of narrow guard cells flanked by Leaves with acute apices, sharp sinuses, margin a broad subsidiary cell on either side) and stellate/ entire, length c. 400 m, width c. 200 m. Stomatal multibranched trichomes. complexes in two compact zones within a shallow Identification. The basic stomatal morphology is furrow on each side of the article, 4-7 files per comparable with the Lauraceae. However, the stel- zone, complexes brachyparacytic, oriented per- late trichomes indicate it clearly belongs to pendicular to the long axis of the branchlet, length Notothixos (Viscaceae), a mistletoe (see Barlow c 20 m. Mostly with 1-2 epidermal cells between 1983 for more details on the trichomes). It may be each stomatal complex in a file. Glabrous. Epider- compared closely with extant N. subaureus Oliver mal cells isodiametric, walls distinctly thickened (1863) (Fig. 16.7-16.8).

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Figure 18. 1-6. Fossil and extant Strasburgeria (Strasburgeriaceae). 1. CUT-Z-FFH, TLM view showing two stomatal complexes (SL3224, scale-bar = 50 µm); 2. CUT-Z-FFH, TLM detail of single stomatal complex (SL3224, scale-bar = 20 µm); 3. CUT-Z-FFH, SEM view of inner cuticular surface showing two stomatal complexes (S-1362, scale-bar = 20 µm); 4. CUT-Z-FFH, SEM view of outer cuticular surface showing a single stomatal complex with prominent outer sto- matal ledges (S-1362, scale-bar = 10 µm); 5. Extant Strasburgeria robusta, TLM view showing a group of networked stomatal complexes (AQ391571, scale-bar = 50 µm). The details are partially hindered by cutinisation around the internal walls of the epidermal cells; 6. Extant S. robusta, TLM view showing a single stomate (AQ391571, scale-bar = 20 µm). compared with the thinner cells of the stomatal nostoma is based on the four-sided articles and rel- zones. atively surficial stomatal complexes (Scriven and Identification. Clearly belongs to the Casuari- Hill 1995). The stomatal length of c. 20 m also sup- naceae on the basis of gross morphology of article ports Gymnostoma, as Scriven and Hill noted that branchlets with highly reduced leaves in whorls, stomatal length in Casuarina and Allocasuarina and transversely oreiented stomata essentially was usually more than 35 m. Based on Scriven restricted to the articles (terminology follows and Hill’s summary, the shape and thickness of the Scriven and Hill 1995). The identification as Gym- epidermal cells is unusual for Gymnostoma.

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Strasburgeriaceae Solereder 1908 pollen from the Gore Lignite Measures (Jarzen and CUT-Z-FFH (Strasburgeria sp.) Pocknall 1993). Figure 18.1-18.4 Reference Specimen and locality: SL3224, Sthd- Cunoniaceae (Brown 1814) 166. Elaeocarpaceae (de Candolle 1816) Stomatal complexes. Stomatal distribution over leaf Family identification: The guard cells of almost all surfaces hypostomatic; stomatal complexes evenly Elaeocarpaceae – Cunoniaceae appear essentially spread, mostly isolated although sometimes net- ‘naked’ under TLM. The outer stomatal ledges are worked, possibly slightly aligned with one another, so thin and smooth that they leave an unobstructed anomocytic (although there are 4–6 contact cells, view of classic ‘paired kidney’ stomata. The typical and sometimes there have been tangential divi- narrow and darkly-staining ring around the sto- sions around a few stomata which produce what matal pore may be an inner ledge, or a thickened could be called subsidiary cells), size range unimo- rim of the outer stomatal ledges. There is often a dal. Guard cell pair outline ovate, outlined by a distinct notch at either pole of the guard cell pair. well-defined anticlinal wall, at same level as sub- These characters are a good first step in suspect- sidiary cells (exposed on surface), length 50–53 m ing Elaeocarpaceae – Cunoniaceae. Pole (1996) (large), with prominent T-piece thickenings at polar wrote “Presently, no cuticular characters are recog- ends. Outer stomatal ledge sub circular, much nised which can separate these two, unrelated thicker than normal epidermal cells, extending over families" (DNA work has since shown them, to be whole stoma, with an elliptical pore. sister taxa). Carpenter et al. (2004) identified an Australian fossil (“Weinmannia sp. 1”) based on Epidermal Cells. Epidermal cell flanges clearly visi- “areolar cyclocytic stomata with 5 or 6 subsidiary ble using TLM, normal cells highly variable from cells, hydathodes, numerous small, trichome base isodiametric to elongate (cells over veins not distin- pores along veins that are surrounded by usually 7 guished by shape), approximately the same size or 8 radially modified cells, and occasional large as the stomata, anticlinal walls straight, or curved diameter cavities presumably left by detached tri- to wavy, unbuttressed, smoothly textured, unorna- chomes”. They noted “This combination of charac- mented. ters is only known in Weinmannia (Barnes et al. Indumentum. Attachment scars of deciduous tri- 2001)”. Barnes et al’s study was restricted to chomes sparse, scattered over venal and non- Cunoniaceae, so it was not possible to make this venal regions, inserted between (6–7) epidermal distinction, and unfortunately, this combination is cells modified only slightly, with a thickened rim. also found in the Elaeocarpaceae. The author’s Fine ridges of cuticle radiate out from the trichome reference collection contains 140 species of Elaeo- attachment scars, scar diameter much smaller than carpus, of which 23 have hydathodes or lenticels a normal epidermal cell. (following Roth 1989) among which include E. alti- Distinguishing features. Distinguished by the sectus Schlechter (1916), E grandis von Mueller strong thickening over the guard cells and the ano- (1861), E. kaniensis Schlechter (1916), and E. seri- mocytic stomatal complexes. copetalus von Mueller (1868). The “large diameter cavities” are a feature of many species in both fam- Identification: The large size and general form of ilies. In some instances they may have had a tri- the stoma, the epidermal cells, and the anomocytic chome in their centre, in others it appears to have stomatal construction find a good match with extant been a giant stomata, but in others the cavity New Caledonian Strasburgeria robusta (Vieill. ex appears primary. The number of subsidiary cells is Panch. and Sebert ) Guillaumin (1942) (Figs 18.5- more variable. The other characters are very wide- 18.6) The differences include scattered trichome spread and variable throughout the two families attachment scars on the fossil, which the extant (Even the “Weinmannia sp. 1” described by Car- species appears to lack, and while stomata are penter et al. (2004) shows several stomata with 4, common in this species and frequently networked, whereas the extant W. paitensis Schlechter (1916) their density is much lower in the fossil although illustrates has at least one stomata with 3 subsid- networking is still apparent. The modern specimen iary cells). It is still my opinion that differentiation of also shows much cuticularisation of the stomatal the two families on cuticular characters alone is not chamber which is absent on the fossil. The differ- possible (although may be for a few genera). The ences would be consistent with a different species following 20 fossil taxa display a broad syndrome of Strasburgeria. The genus has been recorded as of characters that suggest Elaeocarpaceae–

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Figure 19. 1-8. Fossil Cunoniaceae-Elaeocarpaceae: CUT-Z-AJC, and CUT-Z-CEB. 1. CUT-Z-AJC, TLM view show- ing stomatal complexes (SB1343, scale-bar = 50 µm); 2. CUT-Z-AJC, TLM view showing two stomatal complexes (SB1343, scale-bar = 20 µm); 3. CUT-Z-AJC, TLM view showing TLM view showing stomatal complexes and a cork wart (SB1343, scale-bar = 50 µm); 4. CUT-Z-AJC, TLM view showing stomatal complexes and a cork wart (SB1343, scale-bar = 50 µm); 5. CUT-Z-AJC, SEM view of inner cuticular surface showing a single stomatal complex (S-1126, scale-bar = 20 µm); 6. CUT-Z-AJC, SEM view of outer cuticular surface showing stomatal complexes (S-1126, scale- bar = 20 µm); 7. CUT-Z-CEB, TLM view showing stomatal complexes and a cork wart (SL0235, scale-bar = 50 µm); 8. CUT-Z-CEB, TLM detail of single stomatal complex (SL0235, scale-bar = 20 µm).

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Cunoniaceae. In my opinion, the stomatal form 11. Subsidiary cells narrow to isodiametric. CUT-Z- described above, along with giant stomata (hydath- FAJ odes), simple trichome attachment scars, and cork- 12. Epidermal cell anticlinal walls sinuous. CUT-Z- warts, are highly likely to indicate Elaeocarpaceae AJJ –Cunoniaceae. A variety of subsidiary cell arrange- ments are possible. 12 Epidermal cell anticlinal walls straight, curved or wavy. 13. Key to Cunoniaceae-Elaeocarpaceae 13. Epidermal cell periclinal walls thicker than nor- 1. Cuticle glabrous. 2. mal epidermal cells. CUT-Z-FCJ 1. Attachment scars of trichomes present. 5. 13. Epidermal cell periclinal walls same thickness 2. Epidermal cell anticlinal walls sinuous. CUT-Z- as normal epidermal cells. 14. ADF 14. Trichome attachment sites massively thickened 2. Epidermal cell anticlinal walls straight, curved or CUT-Z-FAA wavy. 3. 14. Trichome attachment sites not massively thick- 3. Lenticels or glands absent. 4. ened. 15. 3. Lenticels or glands present. 9. 15. Trichomes inserted over several cells. CUT-Z- FBC 4. Stomatal complexes anomocytic, size range bimodal, 'giant stomatal complexes' present. 15. Trichomes inserted between several cells. 16. CUT-Z-FBG 16. Foot cells around trichome attachment site with 4. Stomatal complexes cyclocytic (developmentally a distinctly circular outer margin, and thick- tangenticytic) 'giant stomatal complexes' ened periclinal walls. CUT-Z-EDG absent. CUT-Z-EFA 16. Foot cells around trichome attachment site 5. Epidermal cell anticlinal walls sinuous. 6. without a distinctly circular outer margin, and without thickened periclinal walls. 17. 5. Epidermal cell anticlinal walls straight, curved or wavy. 12. 17. Cuticle ornamented with flowing fine ridges. CUT-Z-EDJ 6. Bases of the trichomes are persistent, and bal- loon-out from the insertion hole. CUT-Z-ACC 17. Cuticle unornamented. 18. 6. Trichomes purely deciduous. 7. 18. Trichome attachment sites distinctly frilled. CUT-Z-CCE 7. Epidermal cell flanges clearly visible around sto- matal complexes but become unclear further 18. Trichome attachment sites not frilled. 19. away. CUT-Z-CBI 19. Stoma typically distinctly subcircular. CUT-Z- 7. Epidermal cell flanges clearly visible every- EGA where. 8. 19. Stoma typically distinctly broader than long. 8. Subsidiary cell periclinal walls thicker than nor- CUT-Z-CDJ mal epidermal cells, outer stomatal ledge thin. CUT-Z-FBE CUT-Z-AJC Figure 19.1-19.6 8. Subsidiary cell periclinal walls same thickness as normal epidermal cells, outer stomatal Reference Specimen and locality: SB1343, BL-32. ledge distinct. CUT-Z-CFJ Stomatal complexes. Stomatal complexes clus- 9. Stomatal complexes evenly spread. CUT-Z-CEB tered in areoles, typically isolated though network- ing may occur, randomly oriented, cyclocytic, size 9. Stomatal complexes clustered in areoles. 10. range bimodal, with distinct 'giant stomatal com- 10. Subsidiary cell periclinal walls same thickness plexes' present. Subsidiary cells (6–8) irregularly- as normal epidermal cells. CUT-Z-AJC shaped, periclinal walls of same thickness as nor- 10. Subsidiary cell periclinal walls thicker than nor- mal epidermal cells, smooth, unornamented. mal epidermal cells. 11. Guard cell pair outline circular, outlined by a well- defined anticlinal wall, at same level as subsidiary 11. Subsidiary cells elongate and tangential to cells (exposed on surface), length 19–20 m stoma. CUT-Z-CAI (medium), with prominent T-piece thickenings at

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Figure 20. 1-8. Fossil Cunoniaceae-Elaeocarpaceae: CUT-Z-CCE, and CUT-Z-FBG. 1. CUT-Z-CCE, TLM view show- ing stomatal complexes and a trichome attachment scar (SL0099, scale-bar = 50 µm); 2. CUT-Z-CCE, TLM detail of two stomatal complexes (SL0099, scale-bar = 20 µm); 3. CUT-Z-CCE, SEM view of inner cuticular surface showing stomatal complexes. Note granular texture (S-1110, scale-bar = 20 µm); 4. CUT-Z-CCE, SEM view of outer cuticular surface showing stomatal complexes with simple outer stomatal ledges in an otherwise almost smooth surface (S- 1110, scale-bar = 20 µm); 5. CUT-Z-FBG, TLM view showing stomatal complexes (SB0853, scale-bar = 50 µm); 6. CUT-Z-FBG, TLM view showing two stomatal complexes (SL0853, scale-bar = 20 µm); 7. CUT-Z-FBG, TLM view showing stomatal complexes (SL3058, scale-bar = 50 µm); 8. CUT-Z-FBG, TLM view showing two stomatal com- plexes (SL3058, scale-bar = 20 µm).

45 POLE: MIOCENE LEAF CUTICLE polar ends. Outer stomatal ledge sub circular, thin- SL0384, Mata-01; SL2575, Mata-06; SL1754, ner than normal epidermal cells, extending over Sthd-012; SL3216, Sthd-158. whole stoma. Stomatal complexes. Stomatal distribution over leaf Epidermal Cells. Epidermal cell flanges clearly visi- surfaces hypostomatic, stomatal complexes in are- ble using TLM, normal cells isodiametric (cells over oles, isolated, randomly oriented, cyclocytic, devel- both major and fine venation more elongate), opmentally incomplete stephanocytic bicyclic, all approximately the same size as the stomata, anti- some, or none of the subsidiary cells have been clinal walls straight, unbuttressed, inner and outer modified by a tangential division, size range bimo- surfaces smooth, unornamented. dal, with distinct 'giant stomatal complexes' Indumentum. Glabrous. present. Subsidiary cells (4–8) irregularly-shaped, periclinal walls of same thickness as normal epi- Other structures. Lenticels present, with a distinct dermal cells, smooth, unornamented. Guard cell outer zone where the periclinal walls are thicker pair outline circular, or ovate, outlined by a well- than those of normal epidermal cells, and a central defined anticlinal wall, at same level as subsidiary zone where they are thinner. cells (exposed on surface), length 15–20 m (medium), at same level as subsidiary cells CUT-Z-CEB (exposed on surface), little polar development Figure 19.7-19.8 between guard cells (guard cells appear as contin- Reference Specimen and locality: SL0235, BL-04. uous ring). Outer stomatal ledge sub circular, thin- Referred specimens and occurrence: SL3191, BL- ner than normal epidermal cells, extending over 32. whole stoma and just thickened along inner edge, with a broad, sub-circular pore. Stomatal complexes. Stomatal complexes evenly spread, isolated, randomly oriented, cyclocytic, Epidermal Cells. Epidermal cell flanges clearly visi- developmentally stephanocytic bicyclic size range ble using TLM, normal cells isodiametric (cells over unimodal. Subsidiary cells (6–8) irregularly- both major and fine venation more elongate), shaped, periclinal walls of same thickness as nor- approximately the same size as the stomata, anti- mal epidermal cells, smooth, unornamented. clinal walls straight, or curved to wavy, unbut- Guard cell pair outline circular, outlined by a well- tressed, inner surfaces slightly granular, outer defined anticlinal wall, clearly separated by polar surfaces smooth, unornamented. walls, at same level as subsidiary cells (exposed Indumentum. Attachment scars of deciduous tri- on surface). Outer stomatal ledge sub circular, chomes sparse, restricted to regions over veins, same thickness as normal epidermal cells, extend- inserted between (7–9) epidermal cells modified to ing over centre of stoma, with a broad, sub-circular form a ring of more or less isodiametric foot-cells pore. (sometimes with a distinctly circular outer margin) Epidermal Cells. Epidermal cell flanges clearly visi- and with a 'frilly' thickened rim and radial walls, ble using TLM, normal cells isodiametric (cells over scar diameter similar in size to a normal epidermal veins not distinguished by shape), distinctly smaller cell. than the stomata, anticlinal walls straight, unbut- Distinguishing features. Clear and naked guard tressed, smoothly textured, unornamented. cells and distinctly frilled trichome attachment Indumentum. Glabrous. scars.

Other structures. Lenticels present, as large circu- CUT-Z-FBG lar areas of very thinly cutinised cells (often broken Figure 20.7-20.8 away), surrounded by radiating files of normal- thickness epidermal cells. Reference Specimen and locality: SL3058, BL-25. Stomatal complexes. Stomatal complexes in are- CUT-Z-CCE oles, isolated, randomly oriented, anomocytic, size Figure 20.1-20.6 range bimodal, with distinct 'giant stomatal com- Reference Specimen and locality: SL0099, BL-08. plexes' present (restricted to venal regions). Sub- sidiary cells (5–7). Guard cell pair outline circular, Referred specimens and occurrence: CUT-Z-CCE, or ovate, outlined by a well-defined anticlinal wall, SL0353, BL-05; SL3044, BL-22; SL1220, BL-30; at same level as subsidiary cells (exposed on sur- SL3190, BL-32; SL2683, GL-04; SL2982, GL-26; face), length 19–27 m (medium), little polar devel-

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Figure 21. 1-8. Fossil Cunoniaceae-Elaeocarpaceae: CUT-Z-ADF, CUT-Z-EDG, and CUT-Z-FCJ. 1. CUT-Z-ADF, TLM view showing stomatal complexes (SB0844, scale-bar = 50 µm); 2. CUT-Z-ADF, TLM detail of single stomatal complex (SB0844, scale-bar = 20 µm); 3. CUT-Z-ADF, SEM view of outer cuticular surface showing stomatal com- plexes with simple, raised outer stomatal ledges in an otherwise smooth surface (S-1383, scale-bar = 10 µm); 4. CUT-Z-ADF, SEM view of inner cuticular surface showing a single stomatal complex. Note granular texture (S-1383, scale-bar = 10 µm); 5. CUT-Z-EDG, TLM view showing stomatal complexes and a trichome attachment scar (SL2026, scale-bar = 50 µm); 6. CUT-Z-EDG, TLM view showing stomatal complexes (SL2026, scale-bar = 20 µm); 7. CUT-Z-FCJ, TLM view showing stomatal complexes and a giant complex at lower left (SL3085, scale-bar = 50 µm); 8. CUT-Z-FCJ, TLM detail of single stomatal complex (SL3085, scale-bar = 20 µm).

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opment between guard cells (guard cells appear as major veins more elongate), approximately the continuous ring). Outer stomatal ledge sub circular, same size as the stomata, anticlinal walls straight, thinner than normal epidermal cells, with an ellipti- unbuttressed, smoothly textured, unorna- cal pore. mented.Stomatal complexes. Stomatal complexes Epidermal Cells. Epidermal cell flanges clearly visi- evenly spread, isolated, randomly oriented, cyclo- ble using TLM, normal cells highly variable from cytic, all some, or none of the subsidiary cells may isodiametric to elongate (cells over both major and have been modified by a tangential division (devel- fine venation more elongate), approximately the opmentally stephanocytic bicyclic), size range uni- same size as the stomata, anticlinal walls straight, modal (but with wide range of stomatal size). or curved to wavy, unbuttressed, smoothly tex- Subsidiary cells (6–7) irregularly-shaped, periclinal tured, unornamented. walls of same thickness as normal epidermal cells, smooth, unornamented. Guard cell pair outline cir- Indumentum. Glabrous. cular, outlined by a well-defined anticlinal wall, length 14–20 m (small – medium), underthrust by CUT-Z-ADF subsidiary cells (subsidiary cell anticlinal walls Figure 21.1-21.4 bisect outline of the guard cells), with prominent T- Reference Specimen and locality: SB0844, Mata- piece thickenings at polar ends. Outer stomatal 06. ledge sub circular, same thickness or slightly thin- Stomatal complexes. Stomatal complexes in are- ner than normal epidermal cells, extending over oles, isolated, randomly oriented. Subsidiary cells whole stoma, with a broad, sub-circular pore. (5–7) typically elongate, or wedge-like, often with Indumentum. Attachment scars of deciduous tri- distinct polar cells (at right angles to the stomatal chomes common, scattered over venal and non- axis) which may 'enclose' the lateral cells, develop- venal regions, inserted between epidermal cells mentally incomplete stephanocytic bicyclic pericli- modified into a distinct ring of foot cells with a dis- nal walls of same thickness as normal epidermal tinctly circular outer margin, with thickened pericli- cells, smooth, unornamented. Guard cell pair out- nal walls, scar diameter much larger than a normal line circular, outlined by a well-defined anticlinal epidermal cell. wall, at same level as subsidiary cells (exposed on surface), length 20–22 m (medium), at same level CUT-Z-FCJ as subsidiary cells (exposed on surface). Outer Figure 21.7-21.8 stomatal ledge sub circular, thinner than normal Reference Specimen and locality: SL3085, BL-28. epidermal cells, extending over whole stoma, with a broad, sub-circular pore. Stomatal complexes. Stomatal complexes in are- oles, isolated, randomly oriented, cyclocytic, size Epidermal Cells. Epidermal cell flanges clearly visi- range bimodal, with distinct 'giant stomatal com- ble using TLM, normal cells highly variable from plexes' present (restricted to venal regions). Sub- isodiametric to elongate (unclear if cells over veins sidiary cells (6–8) of varying size, curved and are distinguished), approximately the same size as generally tangential to the guard cells, develop- the stomata, anticlinal walls markedly sinuous, mentally incomplete stephanocytic bicyclic, pericli- unbuttressed, inner surface moderately granular, nal walls thicker than over normal epidermal cells, outer surfaces smooth, unornamented. smooth, unornamented. Guard cell pair outline Indumentum. Glabrous. ovate, outlined by a well-defined anticlinal wall, at Distinguishing features. OSL thin and approxi- same level as subsidiary cells (exposed on sur- mates the guard cell outlines very closely, so face), length 23–38 m (medium), some wall devel- appears almost invisible under TLM. Sinuous epi- opment between guard cells, and sometimes dermal cells. development of T-piece thickenings at the poles. Outer stomatal ledge sub circular, thinner than nor- CUT-Z-EDG mal epidermal cells, extending over centre of Figure 21.5-21.6 stoma, with a broad, sub-circular pore. Reference Specimen and locality: SL2026, Sthd- Epidermal Cells. Epidermal cell flanges clearly visi- 087. ble using TLM, normal cells isodiametric (cells over both major and fine venation more elongate), Epidermal Cells. Epidermal cell flanges clearly visi- approximately the same size as the stomata, anti- ble using TLM, normal cells isodiametric (cells over

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Figure 22. 1-8. Fossil Cunoniaceae-Elaeocarpaceae: CUT-Z-ACC, CUT-Z-FAA, and CUT-Z-EGA. 1. CUT-Z-ACC, TLM view showing stomatal complexes and trichome attachment scars (upper right and lower centre) (SB1302, scale-bar = 50 µm); 2. CUT-Z-ACC, TLM detail of single stomatal complex (SB1302, scale-bar = 20 µm); 3. CUT-Z- ACC, SEM view of inner cuticular surface showing a single stomatal complex (S-1406, scale-bar = 10 µm); 4. CUT- Z-ACC, SEM view of outer cuticular surface showing three stomatal complexes with simple outer stomatal ledges, but surrounded by a thin, discontinuous rim (S-1406, scale-bar = 10 µm); 5. CUT-Z-FAA, TLM view showing stomatal complexes and a trichome attachment scar with massive thickening (SL2574, scale-bar = 50 µm); 6. CUT-Z-FAA, TLM view showing two stomatal complexes (SL2574, scale-bar = 20 µm); 7. CUT-Z-EGA, TLM view showing sto- matal complexes (SL1887, scale-bar = 50 µm); 8. CUT-Z-EGA, TLM detail of single stomatal complex (SL1887, scale-bar = 20 µm).

49 POLE: MIOCENE LEAF CUTICLE clinal walls straight, unbuttressed, smoothly tex- reality only becomes clear when a trichome base is tured, unornamented. found which has been flattened obliquely. Indumentum. Attachment scars of deciduous tri- chomes sparse, restricted to regions over major CUT-Z-FAA veins, inserted between (6–8) epidermal cells mod- Figure 22.5-22.6 ified into radially elongate foot cells, scar diameter Reference Specimen and locality: SL2574, Mata- much smaller than a normal epidermal cell. 06. Stomatal complexes. Stomatal distribution over leaf CUT-Z-ACC surfaces hypostomatic, stomatal complexes in are- Figure 22.1-22.4 oles, sometimes networked, randomly oriented, lat- Reference Specimen and locality: SL2660, GL-04. erocytic, developmentally incomplete Referred specimens and occurrence: SL1201, GL- stephanocytic bicyclic, size range unimodal (but 01; SB1302, Mata-03. with wide range of stomatal size). Subsidiary cells (4–7) with periclinal walls of same thickness as Stomatal complexes. Stomatal stomatal complexes normal epidermal cells, smooth, unornamented. in areoles, isolated, randomly oriented; ano- Guard cell pair outline ovate, outlined by a well- mocytic; size range bimodal, with distinct 'giant sto- defined anticlinal wall, at same level as subsidiary matal complexes' present. Subsidiary cells (4–6, cells (exposed on surface), length 20–25 m difficult to tell under TLM) with periclinal walls of (medium), clearly separated by polar walls. Outer same thickness as normal epidermal cells; smooth; stomatal ledge sub circular, thinner than normal unornamented. Guard cell pair outline circular; out- epidermal cells, extending over whole stoma, with lined by a well-defined anticlinal wall; at same level a broad, sub-circular pore. as subsidiary cells (exposed on surface), length 20–31 m (medium); with prominent T-piece thick- Epidermal Cells. Epidermal cell flanges clearly visi- enings at polar ends. Outer stomatal ledge sub cir- ble using TLM, normal cells highly variable from cular, same thickness as normal epidermal cells, isodiametric to elongate (cells over both major and extending over whole stoma, with a broad, sub-cir- fine venation more elongate), approximately the cular pore. same size as the stomata, anticlinal walls straight, unbuttressed, smoothly textured, unornamented. Epidermal Cells. Epidermal cell flanges clearly visi- ble using TLM; normal cells highly variable from Indumentum. Attachment scars of deciduous tri- isodiametric to elongate (cells over veins not distin- chomes common, scattered over venal and non- guished by shape); approximately the same size venal regions, inserted between (6–10) epidermal as the stomata; anticlinal walls markedly sinuous; cells modified by many tangential divisions and unbuttressed; inner surfaces very finely granular, massively thickened to form a frilly rim, scar diame- outer surfaces smooth, although ornamented with ter smaller than a normal epidermal cell. a few bands of irregular ridges parallel to stomatal Distinguishing features. Distinguished from CUT-Z- complexes and radiating from trichomes. EDC in having straight epidermal walls (even distal Indumentum. Attachment scars of deciduous tri- to stomatal complexes), subsidiary cells that do not chomes common; scattered over venal and non- staining darker than normal epidermal cells, and venal regions; inserted between (5–9) epidermal common trichome scars which have massively cells modified into radially elongate foot cells, with thickened poral rims. no distinct thickening of walls; scar diameter much smaller than a normal epidermal cell, but the tri- CUT-Z-EGA chome expanded abruptly above the insertion Figure 22.7-22.8 point. Trichome-bases over the midvein are distinct Reference Specimen and locality: SL1887, Sthd- in having a massively-thickened rim. 073. Distinguishing features. This is reasonably thin Referred specimens and occurrence: SL2245, BL- cuticle, where the guard cells are raised above the 14; SL2109, GL-07; SL2165, GL-09; SL1963, general surface, and have distinct polar T-pieces. Sthd-054; SL1538, Sthd-067. The very bases of the trichomes are persistent, Stomatal complexes. Stomatal complexes in are- and balloon-out from the insertion hole. In TLM this oles, isolated, randomly oriented, anomocytic (and means the insertion hole appears surrounded by a sometimes incompletely cyclocytic), developmen- ring, or perhaps having a circular cap to it. The tally tangenticytic, size range unimodal, (but with

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Figure 23. 1-8. Fossil Cunoniaceae-Elaeocarpaceae: CUT-Z-CDJ, and CUT-Z-EDJ. 1. CUT-Z-CDJ, TLM view show- ing stomatal complexes (SL0279, scale-bar = 50 µm); 2. CUT-Z-CDJ, TLM view showing stomatal complexes (SL0279, scale-bar = 20 µm); 3. CUT-Z-CDJ, SEM view of inner cuticular surface showing a single stomatal complex (S-1221, scale-bar = 10 µm); 4. CUT-Z-CDJ, SEM view of outer cuticular surface showing a single stomatal complex (S-1221, scale-bar = 10 µm); 5. CUT-Z-EDJ, TLM view showing stomatal complexes (SL1902, scale-bar = 50 µm); 6. CUT-Z-EDJ, TLM detail of single stomatal complex (SL1902, scale-bar = 20 µm); 7. CUT-Z-EDJ, SEM view of inner cuticular surface showing two stomatal complexes (S-1370, scale-bar = 20 µm); 8. CUT-Z-EDJ, SEM view of outer cuticular surface showing probable positions of stomatal pores, obscured by cuticular ridges (two are arrowed) (S- 1370, scale-bar = 10 µm).

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wide range of stomatal size). Subsidiary cells (4–5) major veins more elongate), distinctly smaller than irregularly-shaped, (hard to count as radial flanges the stomata, anticlinal walls straight, unbuttressed, of subsidiary cells are not well developed), pericli- inner and outer surfaces smooth, unornamented nal walls of same thickness as normal epidermal except for a few subdued ridges. cells, smooth, unornamented. Guard cell pair out- Indumentum. Scars of deciduous trichomes line circular, not outlined by a clear anticlinal wall, sparse, scattered over venal and non-venal at same level as subsidiary cells (exposed on sur- regions but slightly more common over veins, sim- face), length 16–22 m (medium), at same level as ple, inserted between (5–8) epidermal cells modi- subsidiary cells (exposed on surface), little polar fied to form a broadly thickened rim, scar diameter development between guard cells (guard cells similar in size to a normal epidermal cell. appear as continuous ring). Outer stomatal ledge sub circular, thinner than normal epidermal cells, Distinguishing features. The presence of broadly extending over whole stoma, with a broad, sub-cir- elliptical (wider than long) stomata and the rela- cular pore. tively broad thickening of the trichome scar are both distinctive characters. Epidermal Cells. Epidermal cell flanges clearly visi- ble using TLM, normal cells isodiametric (cells over CUT-Z-EDJ major veins more elongate), approximately the Figure 23.5-23.8 same size as the stomata, anticlinal walls curved to wavy, unbuttressed, smoothly textured, unorna- Reference Specimen and locality: SL1902, Sthd- mented. 073. Indumentum. Attachment scars of deciduous tri- Referred specimens and occurrence: SL1647, chomes common, restricted to regions over veins, Sthd-109. inserted between epidermal cells modified by tan- Stomatal complexes. Stomatal complexes evenly gential divisions to form a sub-circular zone of foot spread, isolated, randomly oriented, cyclocytic, cells, scar diameter similar in size to a normal epi- size range unimodal. Subsidiary cells (4–5) irregu- dermal cell. larly-shaped, with periclinal walls thicker than over normal epidermal cells, smooth, unornamented. CUT-Z-CDJ Guard cell pair outline ovate, at same level as sub- Figure 23.1-23.4 sidiary cells, outer margin obscured under TLM by Reference Specimen and locality: SL0279, BL-01. surface ornamentation, length 35–40 m (medium - large). Outer stomatal ledge elliptical, thinner than Referred specimens and occurrence: SL0354, BL- normal epidermal cells, extending over whole 05. stoma, with a narrowly elliptic pore. Stomatal complexes. Stomatal distribution over leaf Epidermal Cells. Epidermal cell flanges clearly visi- surfaces hypostomatic, stomatal complexes in are- ble using TLM, irregularly extended, normal cells oles, isolated, randomly oriented, cyclocytic, devel- isodiametric (unclear if cells over veins are distin- opmentally incomplete stephanocytic bicyclic, size guished), approximately the same size as the sto- range bimodal, with distinct 'giant stomatal com- mata, anticlinal walls straight, unbuttressed, inner plexes' present. Subsidiary cells (5–8) irregularly- surfaces smooth, outer surfaces smooth and orna- shaped, periclinal walls of same thickness as nor- mented with a flowing pattern of many fine ridges. mal epidermal cells, smooth, with a narrow, discon- tinuous peristomatal ridge, otherwise Indumentum. Deciduous trichome scars sparse, unornamented. Guard cell pair outline varies from not clear. broadly elliptic (wider than long), to circular, to nar- rowly elliptic, outlined by a well-defined anticlinal CUT-Z-FBC wall, length 12–18 m (small), at same level as sub- Figure 24.1-24.4 sidiary cells (exposed on surface), clearly sepa- Reference Specimen and locality: SL3022, GL-29. rated by polar walls. Outer stomatal ledge sub Stomatal complexes. Stomatal complexes in are- circular, thinner than normal epidermal cells, oles, isolated, randomly oriented, cyclocytic, There extending over centre of stoma, with a broad, sub- are usually two rings of subsidiary cells, an inner circular pore. with periclinal walls thinner than epidermal cells, Epidermal Cells. Epidermal cell flanges clearly visi- and an outer with periclinal walls thicker than epi- ble using TLM, normal cells isodiametric (cells over dermal cells, developmentally stephanocytic bicy-

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Figure 24. 1-8. Fossil Cunoniaceae-Elaeocarpaceae: CUT-Z-FBC, and CUT-Z-FBE. 1. CUT-Z-FBC, TLM view show- ing stomatal complexes (SL3022, scale-bar = 50 µm); 2. CUT-Z-FBC, TLM detail of single stomatal complex (SL3022, scale-bar = 20 µm); 3. CUT-Z-FBC, TLM view of a trichome attachment scar (SL3022, scale-bar = 20 µm); 4. CUT-Z-FBC, SEM view of outer cuticular surface showing a single stomatal complex (S-1388, scale-bar = 10 µm); 5. CUT-Z-FBE, TLM view showing stomatal complexes (SL3007, scale-bar = 50 µm); 6. CUT-Z-FBE, TLM view show- ing two stomatal complexes (SL3007, scale-bar = 20 µm); 7. CUT-Z-FBE, SEM view of inner cuticular surface show- ing a single stomatal complex (S-1389, scale-bar = 10 µm); 8. CUT-Z-FBE, SEM view of outer cuticular surface showing two stomatal complexes (S-1389, scale-bar = 10 µm).

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Figure 25. 1-8. Fossil Cunoniaceae-Elaeocarpaceae: CUT-Z-AJJ, CUT-Z-CFJ, CUT-Z-EFA, and CUT-Z-CAI. 1. CUT- Z-AJJ, TLM view showing stomatal complexes (SB1369, scale-bar = 50 µm); 2. CUT-Z-AJJ, TLM detail of single sto- matal complex (SB1369, scale-bar = 20 µm); 3. CUT-Z-CFJ, TLM view showing stomatal complexes (SL0347, scale- bar = 50 µm); 4. CUT-Z-CFJ, TLM detail of single stomatal complex (SL0347, scale-bar = 20 µm); 5. CUT-Z-EFA, TLM view showing (SL1899, scale-bar = 50 µm); 6. CUT-Z-EFA, TLM detail of single stomatal complex (SL1899, scale-bar = 20 µm); 7. CUT-Z-CAI, TLM view showing stomatal complexes (SL3294, scale-bar = 50 µm); 8. CUT-Z- CAI, TLM detail of single stomatal complex (SL3294, scale-bar = 20 µm).

54 PALAEO-ELECTRONICA.ORG clic, size range unimodal (but with wide range of the stomata, anticlinal walls markedly sinuous, stomatal size). Subsidiary cells (4–7) typically elon- unbuttressed, inner surfaces slightly granular, gate, or wedge-like, often with distinct polar cells outer surfaces smooth, unornamented. (at right angles to the stomatal axis) which may Indumentum. Attachment cars of deciduous tri- 'enclose' the lateral cells, smooth, unornamented. chomes sparse, restricted to regions over major Guard cell pair outline circular, or ovate, outlined by veins, inserted between (5–7) epidermal cells mod- a well-defined anticlinal wall, at same level as sub- ified into a distinct ring of foot cells, with thickened sidiary cells (exposed on surface), length 30–37 m periclinal walls, scar diameter similar in size to a (medium), with prominent T-piece thickenings at normal epidermal cell. polar ends. Outer stomatal ledge sub circular, thin- ner than normal epidermal cells, extending over CUT-Z-AJJ whole stoma, with a broad, sub-circular pore. Figure 25.1-25.2 Epidermal Cells. Epidermal cell flanges clearly visi- Reference Specimen and locality: SB1369, BL-32. ble using TLM, normal cells isodiametric (cells over major veins more elongate), approximately the Stomatal complexes. Stomatal complexes isolated, same size as the stomata, anticlinal walls curved to randomly oriented, cyclocytic, all some, or none of wavy, unbuttressed, inner and outer surfaces the subsidiary cells have been modified by a tan- smooth, unornamented, but with sharp grooves gential division, size range unimodal. Subsidiary over the anticlinal walls. cells (4–5) sometimes forming 2 distinct polar cells (right angles to stomatal orientation) and 2 lateral Indumentum. Attachment scars of deciduous tri- cells (parallel to stomatal orientation), developmen- chomes common, scattered over venal and non- tally incomplete stephanocytic bicyclic, periclinal venal regions, inserted between epidermal cells, walls of same thickness as normal epidermal cells, with the distinctly circular and thickened base smooth, unornamented. Guard cell pair outline cir- expanding over (6–7) surrounding cells and similar cular, outlined by a well-defined anticlinal wall, in size to a normal epidermal cell. length 35–40 m (medium), at same level as subsid- Distinguishing features. The multicellular trichome iary cells (exposed on surface), some wall develop- attachment scar with its central insertion point is ment between guard cells, and sometimes distinctive. development of T-piece thickenings at the poles, slightly granular. Outer stomatal ledge sub circular, CUT-Z-FBE same thickness as normal epidermal cells, extend- Figure 24.5-24.8 ing over whole stoma, with a broad, sub-circular Reference Specimen and locality: SL3007, GL-28. pore. Stomatal complexes. Stomatal complexes in are- Epidermal Cells. Epidermal cell flanges clearly visi- oles, isolated, randomly oriented, cyclocytic, size ble using TLM, normal cells elongate (cells over range bimodal, with distinct 'giant stomatal com- major veins more elongate), approximately the plexes' present (restricted to venal regions). Sub- same size as the stomata, anticlinal walls curved to sidiary cells (5–7) irregularly-shaped, periclinal wavy, unbuttressed, smoothly textured, unorna- walls thicker than over normal epidermal cells, mented. developmentally incomplete stephanocytic bicyclic, Indumentum. Attachment scars of deciduous tri- smooth, unornamented. Guard cell pair outline cir- chomes sparse, restricted to regions over major cular, or ovate, outlined by a well-defined anticlinal veins, simple, inserted between (c. 8) epidermal wall, at same level as subsidiary cells (exposed on cells modified into radially elongate foot cells, scar surface), length 22–25 m (medium), with T-piece diameter much smaller than a normal epidermal thickenings at polar ends (often just the cross-bar cell. of the T). Outer stomatal ledge sub circular to ellip- Distinguishing features. Subsidiary cells having tical, thinner than normal epidermal cells, extend- distinctly straight tangential walls and with clear ing over whole stoma, with a broad, sub-circular polar and lateral forms. Epidermal cells highly sinu- pore. ous. Epidermal Cells. Epidermal cell flanges clearly visi- ble using TLM, normal cells highly variable from CUT-Z-CFJ isodiametric to elongate (cells over both major and Figure 25.3-25.4 fine venation more elongate), distinctly larger than Reference Specimen and locality: SL0347, BL-05.

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Figure 26. 1-8. Fossil Cunoniaceae-Elaeocarpaceae: CUT-Z-CBI and CUT-Z-FAJ. 1. CUT-Z-CBI, TLM view showing stomatal complexes and (lower right) a trichome attachment scar. Note how outlines of epidermal cells become obscure away from stomatal complexes (SL3049, scale-bar = 50 µm); 2. CUT-Z-CBI, TLM view showing stomatal complexes and (centre) a trichome attachment scar (SL3049, scale-bar = 50 µm); 3. CUT-Z-CBI, TLM detail of single stomatal complex with clear brachyparacytic structure (SL3049, scale-bar = 20 µm); 4. CUT-Z-CBI, TLM detail of sin- gle stomatal complex with no apparent brachyparacytic structure (SL3049, scale-bar = 20 µm); 5. CUT-Z-CBI, SEM view of inner cuticular surface showing a single stomatal complex. Note very shallow anticlinal walls of subsidiary cells, and essential absence of walls else where (S-1122, scale-bar = 20 µm); 6. CUT-Z-CBI, SEM view of outer cuticular surface showing a single stomatal complex (S-1122, scale-bar = 20 µm); 7. CUT-Z-FAJ, TLM view showing stomatal complexes. Note giant stomatal complex (lower right) (SL2106, scale-bar = 20 µm); 8. CUT-Z-FAJ, TLM view showing stomatal complexes. The narrow subsidiary cells are well-defined by their thicker periclinal walls (SL2108, scale-bar = 50 µm).

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Stomatal complexes. Stomatal complexes in are- Indumentum. Glabrous. oles, isolated, randomly oriented, anomocytic, size range unimodal. Subsidiary cells (4–7) when dis- CUT-Z-CAI tinct are typically elongate, or wedge-like, often Figure 25.7-25.8 with distinct polar cells (at right angles to the sto- Reference Specimen and locality: SL3294, BL-04. matal axis) which may 'enclose' the lateral cells, developmentally incomplete stephanocytic bicyclic, Referred specimens and occurrence: SL3280, BL- periclinal walls of same thickness as normal epi- 05; SL2848, GL-18. dermal cells, smooth, unornamented. Guard cell Stomatal complexes. Stomatal complexes in are- pair outline ovate, outlined by a well-defined anticli- oles, isolated, randomly oriented, cyclocytic, devel- nal wall, length 25–30 m (medium); at same level opmentally tangenticytic or stephanocytic bicyclic, as subsidiary cells (exposed on surface); with size range unimodal. Subsidiary cells (5–8) isodia- prominent T-piece thickenings at polar ends. Outer metric or elongate and tangential to the guard cells, stomatal ledge sub circular, same thickness as nor- periclinal walls thicker than over normal epidermal mal epidermal cells, extending over centre of cells, smooth, unornamented. Guard cell pair out- stoma, with a broad, sub-circular pore. line circular, outlined by a well-defined anticlinal Epidermal Cells. Epidermal cell flanges clearly visi- wall, at same level as subsidiary cells (exposed on ble using TLM; normal cells elongate (cells over surface), length 23–28 m (medium), clearly sepa- major veins more elongate); approximately the rated by polar walls. Outer stomatal ledge sub cir- same size as the stomata; anticlinal walls markedly cular, thicker than normal epidermal cells, sinuous; unbuttressed; smoothly textured, unorna- extending over centre of stoma, with a broad, sub- mented. circular pore. Indumentum. Attachment scars of deciduous, poral Epidermal Cells. Epidermal cell flanges clearly visi- trichomes sparse; restricted to regions over veins; ble using TLM, normal cells elongate (cells over inserted between (c. 7) epidermal cells modified major veins more elongate), approximately the into radially elongate foot cells; scar diameter same size as the stomata, anticlinal walls curved to much smaller than a normal epidermal cell. wavy, unbuttressed, smoothly textured, unorna- mented. CUT-Z-EFA Indumentum. Glabrous. Figure 25.5-25.6 Other structures. Lenticels present, formed of large Reference Specimen and locality: SL1899, Sthd- circular areas of very thinly cutinised cells (often 073. broken away), surrounded by radiating files of epi- Stomatal complexes. Stomatal complexes evenly dermal cells with relatively thick periclinal walls. spread, isolated, randomly oriented, cyclocytic, developmentally tangenticytic or stephanocytic CUT-Z-CBI bicyclic, size range unimodal. Subsidiary cells (4– Figure 26.1-26.6 5) elongate, periclinal walls thicker than over nor- Reference Specimen and locality: SL3049, BL-22. mal epidermal cells, smooth, unornamented. Stomatal complexes. Stomatal complexes small Guard cell pair outline ovate, not outlined by a clusters, isolated, randomly oriented, anomocytic, clear anticlinal wall, length c. 32 m (medium), at or brachyparacytic, developmentally incomplete same level as subsidiary cells (exposed on sur- stephanocytic bicyclic or amphibrachyparacytic, face), little polar development between guard cells size range unimodal. Subsidiary cells (5–7 contact (guard cells appear as continuous ring). Outer sto- cells) irregularly-shaped, or typically elongate, tan- matal ledge elliptical, much thinner than normal gential to the guard cells, periclinal walls of same epidermal cells (often broken away), extending thickness as normal epidermal cells, smooth, unor- over whole stoma, with a narrowly elliptic pore. namented. Guard cell pair outline circular, outlined Epidermal Cells. Epidermal cell flanges clearly visi- by a well-defined anticlinal wall, at same level as ble using TLM, normal cells isodiametric (cells over subsidiary cells (exposed on surface), length 15– veins not distinguished by shape), approximately 25 m (medium). Outer stomatal ledge sub circular, the same size as the stomata, anticlinal walls thicker than normal epidermal cells, extending over straight, or curved to wavy, unbuttressed, smoothly inner edge of stoma, with a broad, sub-circular textured, unornamented. pore.

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Figure 27. 1-8. Fossil and extant Meliaceae. Note the tendency to have well-defined outer stomatal ledges “floating” within a poorly–outlined stoma: 1. Extant Dysoxylum spectabile, TLM view (OPH2595, scale-bar = 50 µm); 2. Extant Reinwardtiodendron celebicum, TLM view (AQ062423, scale-bar = 50 µm); 3. CUT-Z-AAB, TLM view showing sto- matal complexes (SB1344, scale-bar = 50 µm); 4. CUT-Z-AAB, TLM detail of single stomatal complex (SB1344, scale-bar = 20 µm); 5. CUT-Z-CCH, TLM view showing stomatal complexes (SL0330, scale-bar = 50 µm); 6. CUT-Z- CCH, TLM view showing three stomatal complexes. (SL0330, scale-bar = 20 µm); 7. CUT-Z-FFC, TLM view showing stomatal complexes (SL3009, scale-bar = 50 µm); 8. CUT-Z-FFC, TLM detail of single stomatal complex (SL3009, scale-bar = 20 µm).

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Epidermal Cells. Epidermal cell flanges clearly visi- zone between guard cells and subsidiary cells is ble around stomatal complexes but become often covered by very thin cuticle which leads to unclear further away (cells over veins not distin- the appearance under TLM of the outer stomatal guished by shape), approximately the same size ledge seeming to ‘float’. The pattern of subsidiary as the stomata, anticlinal walls sinuous, unbut- cells is often complex, the result of tangential cell tressed, inner surfaces slightly granular, outer sur- division (see Figs 27.1-27.2 for two examples of faces smooth, unornamented. extant Meliaceae cuticle, Dysoxylum spectabile Indumentum. Attachment scars of deciduous tri- (G.Forst.) Hooker (1864) and Reinwardtiodendron chomes common, epidermal cells around trichome celebicum Koorders (1898). The following five taxa attachment scar (6–7) modified into radially elon- are offered as likely examples of fossil Meliaceae. gate foot cells. Key to Meliaceae

CUT-Z-FAJ 1. Cuticle glabrous. CUT-Z-CCH Figure 26.7-26.8 1. Trichome attachment sites present. 2. Reference Specimen and locality: SL2108, GL-07. 2. Ornamented with 'flowing' pattern of many fine Referred specimens and occurrence: SL2252, BL- ridges. CUT-Z-CDD 14; SL1242, BL-32; SL2166, GL-09; SL2175, GL- 2. Cuticle unornamented. 3. 10. 3. Periclinal walls of epidermal cells distinctly thin- Epidermal Cells. Epidermal cell flanges clearly visi- ner than normal epidermal cells. CUT-Z-FFC ble using TLM, normal cells elongate (cells over 3. Periclinal walls of epidermal cells same thick- both major and fine venation more elongate), dis- ness (or sometimes slightly thicker) than normal tinctly larger than the stomata, anticlinal walls epidermal cells. 4. straight, unbuttressed, smoothly textured, unorna- mented.Stomatal complexes. Stomatal complexes 4. Subsidiary cells (4-8) typically isodiametric, anti- in areoles, isolated (very rarely networked), ran- clinal walls typically slight indistinct. CUT-Z-CDE domly oriented, cyclocytic, developmentally tan- 4. Subsidiary cells (3-5) very irregularly shaped, genticytic or stephanocytic bicyclic, size range anticlinal walls typically distinct. CUT-Z-AAB bimodal, with distinct 'giant stomatal complexes' present. Subsidiary cells (4–7) with periclinal walls CUT-Z-AAB thicker than over normal epidermal cells, smooth, Figure 27.3-27.4 unornamented, shape varying from narrow to isodi- Reference Specimen and locality: SB1344, BL-32. ametric. Guard cell pair outline circular, outlined by a well-defined anticlinal wall, at same level as sub- Stomatal complexes. Stomatal complexes in are- sidiary cells (exposed on surface), length 20–32 m oles, isolated, randomly oriented, anisocytic, size (medium), with prominent T-piece thickenings at range unimodal. Subsidiary cells (3–5) with pericli- polar ends. Outer stomatal ledge sub circular, thin- nal walls of same thickness as normal epidermal ner than normal epidermal cells, extending over cells, smooth, unornamented. Guard cell pair out- whole stoma, with a broad, sub-circular pore. line ovate, not outlined by a clear anticlinal wall, length 17–20 m (medium), at same level as subsid- Indumentum. Glabrous. iary cells (exposed on surface), little polar develop- ment between guard cells (guard cells appear as Glands. Glands present, distinguished by patches continuous ring). Outer stomatal ledge sub circular, of thin cuticle (often broken away) located over same thickness as normal epidermal cells, with an veins, surrounded by relatively small epidermal elliptical pore. cells with thickly cutinised walls. Epidermal Cells. Epidermal cell flanges clearly visi- Meliaceae Jussieu 1789 ble using TLM, normal cells elongate (cells over major veins more elongate), approximately the Family identification: Based on the reference col- same size as the stomata, anticlinal walls curved to lection, one Meliaceae “syndrome” that stands out wavy, unbuttressed, smoothly textured, unorna- is a tendency to have stomata in which the margin mented. of the guard cell pair is indistinct, while the outer stomatal ledge is well-defined but narrow, covering Indumentum. Glabrous. just the inner part of the guard cells. The contact

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Figure 28. 1-8. Fossil Meliaceae. Note the tendency to have well-defined outer stomatal ledges “floating” within a poorly–outlined stoma: CUT-Z-CDE, and CUT-Z-CDD. 1. CUT-Z-CDE, TLM view showing stomatal complexes (SL0302, scale-bar = 50 µm); 2. CUT-Z-CDE, TLM detail of single stomatal complex (SL0302, scale-bar = 20 µm); 3. CUT-Z-CDE, SEM view of outer surface showing stomatal complexes in a slightly papillate surface (S-1347, AQ 441329, scale-bar = 50 µm); 4. CUT-Z-CDE, SEM view of inner cuticular surface showing four stomatal complexes. Note clearly different texture of subsidiary cells than ordinary epidermal cells, and that by this criteria, not all subsid- iary cells are in direct contact with the guard cells (S-1346, B. Hyland 8650, scale-bar = 50 µm); 5. CUT-Z-CDD, TLM view showing stomatal complexes (SL0342, scale-bar = 50 µm); 6. CUT-Z-CDD, TLM view showing two stomatal complexes (SL0342, scale-bar = 20 µm); 7. CUT-Z-CDD, SEM view of inner cuticular surface showing a single sto- matal complex (S-1201, scale-bar = 20 µm); 8. CUT-Z-CDD, SEM view of outer cuticular surface showing a single stomatal complex with radiating striae (S-1201, scale-bar = 20 µm).

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Distinguishing features. Distinguished by having Indumentum. Attachment scars of deciduous tri- highly variable subsidiary cells (although generally chomes sparse, mostly over, or close to, major on an anisocytic pattern). veins, inserted between (5–7) epidermal cells mod- ified only slightly, with a thickened rim, scar diame- CUT-Z-CCH ter much smaller than a normal epidermal cell. Figure 27.5-27.6 Reference Specimen and locality: SL0330, Harli- CUT-Z-CDE wich-3. Figure 28.1-28.4 Stomatal complexes. Stomatal surfaces, stomatal Reference Specimen and locality: SL0302, BL-01. complexes in areoles, isolated, randomly oriented, Referred specimens and occurrence: SL2281, BL- cyclocyctic, size range unimodal, (but with a wide 18; SL3070, BL-26; SL2131, GL-08; SL2151, GL- range of stomatal size). Subsidiary cells (5–9), iso- 09; SL2197, GL-11; SL2853, GL-18; SL2894, GL- diametric, periclinal walls thicker than over normal 20; SL2938, GL-23; SL3099, GL-24; SL2961, GL- epidermal cells, smooth, unornamented. Guard cell 25; SL2984, GL-27; SL2990, GL-27. pair outline ovate, outlined by a well-defined anticli- Stomatal complexes. Stomatal complexes in are- nal wall, length 15–20 m (medium), at same level oles, isolated, randomly oriented, cyclocytic, all as subsidiary cells (exposed on surface). Outer some, or none of the subsidiary cells may have stomatal ledge narrowly elliptical, thinner than nor- been modified by a tangential division, size range mal epidermal cells, extending over inner edge of unimodal. Subsidiary cells (4–8) irregularly- stomate. Pore narrowly elliptic. shaped, periclinal walls same thickness or some- Epidermal Cells. Epidermal cell flanges clearly visi- times thicker than over normal epidermal cells, with ble using TLM, normal cells isodiametric, distinctly a slightly more granular inner surface than normal smaller than the stomata, cells over major veins epidermal cells, unornamented. Guard cell pair more elongate, anticlinal walls curved - wavy, outline ovate, not outlined by a clear anticlinal wall, unbuttressed, texture smooth, unornamented. length 23–25 m (medium), at same level as subsid- Indumentum. Glabrous. iary cells (stomatal complex slightly sunken so that outer stomatal ledges are at same level as surface Distinguishing features. Distinguished from CUT-Z- of epidermal cells), little polar development CDE and CUT-Z-EID by the absence of trichome between guard cells (guard cells appear as contin- attachment scars, by subsidiary cells which are not uous ring). Outer stomatal ledge elliptical, much clearly defined, and by a simple and thin OSL. thicker than normal epidermal cells, extending over inner edge of stoma. Pore narrowly elliptic. CUT-Z-FFC Figure 27.7-27.8 Epidermal Cells. Epidermal cell flanges clearly visi- ble using TLM, normal cells isodiametric (cells over Reference Specimen and locality: SL3009, GL-28. both major and fine venation more elongate), dis- Stomatal complexes. Stomatal complexes evenly tinctly smaller than the stomata, anticlinal walls spread, isolated, randomly oriented, actinocytic, or straight, unbuttressed, inner surfaces smooth, staurocytic, size range unimodal. Subsidiary cells outer surfaces irregular, slightly rough and some (3–5) irregularly-shaped, periclinal walls thicker cells raised up in a slightly papillose manner, other- than over normal epidermal cells, unornamented. wise unornamented. Guard cell pair outline difficult or impossible to see Indumentum. Attachment scars of deciduous tri- in TLM view, not outlined by a clear anticlinal wall, chomes sparse, scattered over venal and non- at same level as subsidiary cells (exposed on sur- venal regions but slightly more common over face), length 15–25 m (medium). Outer stomatal veins, simple, inserted between (5–8) epidermal ledge elliptical, thinner than normal epidermal cells, cells modified only slightly, with a thickened rim, extending over whole stoma. Pore narrowly elliptic. scar diameter much smaller than a normal epider- Epidermal Cells. Epidermal cell flanges clearly visi- mal cell. ble using TLM, normal cells highly variable from Distinguishing features. Distinguished from CUT-Z- isodiametric to elongate (cells over veins not distin- CCH by having clearly defined subsidiary cells guished by shape), approximately the same size (based on texture), prominent OSL, and stomatal as the stomata, anticlinal walls curved to wavy, complexes which are about twice as large. CUT-Z- slightly buttressed, smoothly textured, unorna- mented.

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Figure 29. 1-8. Fossil Sapindaceae: CUT-Z-EDD and CUT-Z- FJI. 1. CUT-Z-EDD, TLM view showing stomatal com- plexes (SL1818, scale-bar = 50 µm); 2. CUT-Z-EDD, TLM view showing three stomatal complexes. Note that the walls of the subsidiary cells abut or slightly bisect the outline of the guard cells (SL1818, scale-bar = 20 µm); 3. CUT- Z-EDD, SEM view of inner cuticular surface showing two stomatal complexes. Note buttressing of epidermal cell walls (S-1036, scale-bar = 20 µm); 4. CUT-Z-EDD, SEM view of outer cuticular surface showing two stomatal complexes. Note striae radiating from each complex (S-1036, scale-bar = 20 µm); 5. CUT-Z- FJI, TLM view showing stomatal complexes (SL2539, scale-bar = 50 µm); 6. CUT-Z- FJI, TLM detail of single stomatal complex (SL2539, scale-bar = 20 µm); 7. CUT-Z- FJI, SEM view of inner cuticular surface showing a single stomatal complex (S-1393, scale-bar = 10 µm); 8. CUT-Z- FJI, SEM view of outer cuticular surface showing stomatal complexes. Note guard cell pair are slightly sunken and the very small outer stomatal ledges do not protrude above the surrounding subsidiary cells (S- 1393, scale-bar = 10 µm).

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CDE differs in having many subsidiary cells and no indaceae based on broad resemblance to extant peristomatal ridge. taxa. Key to Sapindaceae CUT-Z-CDD Figure 28.5-28.8 1. Epidermal cell flanges buttressed. CUT-Z-EDD Reference Specimen and locality: SL0342, Mata- 1. Epidermal cell flanges unbuttressed. 2. 23. 2. Epidermal cell walls sinuous. CUT-Z-GDC Referred specimens and occurrence: SL3130, GL- 2. Epidermal cell walls straight or wavy. 3. 02; SL2208, GL-12; SL2930, GL-22; SL2992, GL- 27. 3. Cuticle unornamented, subsidiary cell number up to 5. CUT-Z-FJI Stomatal complexes. Stomatal complexes evenly spread, isolated, randomly oriented, cyclocytic, 3. Cuticle ornamented with ridges, subsidiary cell size range unimodal (but with wide range of sto- number up to 6. 4. matal size). Subsidiary cells (4–5), irregularly- 4. Subsidiary cells elongate tangential to stoma, shaped, difficult to count under TLM; periclinal often wedge-like. CUT-Z-CEI walls thicker than over normal epidermal cells; 4. Subsidiary cells typically isodiametric, not smooth; ornamented with groups of ridges radiat- wedge-like. CUT-Z-ABF ing from the stoma. Guard cell pair outline ovate; not outlined by a clear anticlinal wall; at same level CUT-Z-EDD as subsidiary cells (exposed on surface), length Figure 29.1-29.4 20–22 m (medium). Outer stomatal ledge elliptical, thinner than normal epidermal cells, extending over Reference Specimen and locality: SL1818, Sthd- inner edge of stoma. Pore narrowly elliptic. 091. Epidermal Cells. Epidermal cell flanges clearly visi- Referred specimens and occurrence: SL2540, GL- ble using TLM; normal cells isodiametric (cells over 02; SL1733, Sthd-017; SL1713, Sthd-018; SL1965, major veins more elongate); distinctly larger than Sthd-054; SL1993, Sthd-072; SL1622, Sthd-106; the stomata; anticlinal walls markedly sinuous; but- SL1595, Sthd-108. tressed, inner surfaces smooth, outer surfaces Stomatal complexes. Stomatal complexes evenly smooth and ornamented with 'flowing' pattern of spread, sometimes networked, randomly oriented, many fine ridges, areas over anticlinal walls staurocytic, developmentally tangenticytic, size marked by a groove flanked on either side by a range bimodal, with distinct 'giant stomatal com- ridge. plexes' present. Subsidiary cells (4–5) typically Indumentum. Attachment scars of deciduous tri- elongate, or wedge-like, or skewed tangentially, chomes sparse, probably restricted to regions over often forming distinct polar cells (at right angles to veins (unclear), inserted between (5–6) epidermal the stomatal axis) which may 'enclose' the lateral cells modified into a distinct ring of foot cells, with cells, periclinal walls thicker than over normal epi- thickened periclinal walls, scar diameter much dermal cells, smooth, unornamented. Guard cell smaller than a normal epidermal cell. pair outline ovate, not outlined by a clear anticlinal wall, at same level as subsidiary cells (stomatal Sapindaceae Jussieu 1789 complexes sunken so that outer stomatal ledges are at same level as surface of epidermal cells), Family identification: The extant Sapindaceae have length 22–30 m (medium), little polar development a very wide range of epidermal morphology. The between guard cells (guard cells appear as contin- epidermis of the Australasian species is under uous ring). Outer stomatal ledge sub circular, thin- review by the author and although some genera ner than normal epidermal cells, extending over and groups of genera are distinct, there is no sim- inner edge of stoma, with an elliptical pore. ple set of epidermal features which will identify the family. However, stomata which are at the small Epidermal Cells. Epidermal cell flanges clearly visi- end of the medium size class, have thin-walls, and ble using TLM, normal cells isodiametric (cells over subsidiary cells with flanges which abruptly end or major veins more elongate), approximately the even overlap the outline of the stomata appear to same size as the stomata, anticlinal walls markedly be common and distinctive in the Sapindaceae. sinuous, buttressed, inner surfaces smooth, outer The following five parataxa are suggested as Sap- surfaces smooth except ornamented with narrow bands of fine ridges joining some stomatal com-

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Figure 30. Extant Sapindaceae. 1. Cupaniopsis anacardioides, TLM view of stomatal complexes (AQ380045, scale- bar = 20 µm); 2. Diploglottis diphyllostegia, TLM view of stomatal complexes (AQ011960, scale-bar = 20 µm); 3. Rhys- otoechia bifoliolata, TLM view of stomatal complexes (AQ504166, scale-bar = 50 µm); 4. Mischocarpus stipitatus, TLM view of stomatal complexes (AQ389174, scale-bar = 20 µm).

plexes and radiating from trichome attachment iary cells which have prominent flanges terminating scars. abruptly at the stomatal outline, and in some cases Indumentum. Scars of deciduous trichomes bisect the outline. It is compared with extant sparse, scattered over venal and non-venal Cupaniopsis anacardioides (A.Rich.) Radlk. (1879) regions but slightly more common over veins, in terms of stoma shape, thin outer stomatal inserted between epidermal cells modified into ledges, thickened subsidiary cell periclinal walls radially elongate foot cells, scar diameter much and flanges of the subsidiary cells which abruptly smaller than a normal epidermal cell. abut against the stomatal outline (Fig. 30.1). How- ever, the subsidiary cell flanges tend not to project Distinguishing features. One of a group of taxa over the guard cell outlines and the subsidiary cells which have relatively small stomatal complexes are typically elongate, which is not a feature of C. without a distinct rim around the guard cells, and anacardioides. where the flanges of the subsidiary cells some- times project over the stomatal outline. CUT-Z- CUT-Z-FJI EDD is distinct in having buttressed epidermal cells Figure 29.5-29.8 and straight subsidiary cell anticlinal walls. CUT-Z- FJI differs in having unbuttressed epidermal cell Reference Specimen and locality: SL2539, GL-02. walls, and curved subsidiary cell anticlinal walls. Referred specimens and occurrence: SL2260, BL- CUT-Z-ABF differs in having large diameter, hollow 15; SL2270, BL-16. trichome attachment scars, which are surrounded Stomatal complexes. Stomatal complexes in are- by a ring of many foot cells, each distinctly smaller oles, isolated, randomly oriented, actinocytic, or than the trichome diameter. staurocytic, size range unimodal. Subsidiary cells Identification. Suggested as Sapindaceae based (4–5) with periclinal walls of same thickness as on the thin-walled stomata surrounded by subsid- normal epidermal cells, smooth, unornamented.

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Figure 31. 1-6. Fossil Sapindaceae: CUT-Z-ABF, CUT-Z-CEI and CUT-Z-GDC. 1. CUT-Z-ABF, TLM view showing stomatal complexes and (just to lower right of centre) a trichome attachment scar (SB1382, scale-bar = 50 µm); 2. CUT-Z-ABF, TLM detail of single stomatal complex. Note flanges of subsidiary cells projecting over the outline of the guard cells (SB1382, scale-bar = 20 µm); 3. CUT-Z-ABF, SEM view of inner cuticular surface showing three stomatal complexes (S-1345, NE 043791A, scale-bar = 10 µm); 4. CUT-Z-ABF, SEM view of outer cuticular surface with a clear stomatal complex at upper right. Note short bands of striae radiating from complex (S-1345, NE 043791A, scale-bar = 20 µm); 5. CUT-Z-CEI, TLM view showing stomatal complexes and trichome attachment scars (SL0364, scale-bar = 50 µm); 6. CUT-Z-CEI, TLM detail of single stomatal complex and trichome attachment scar (SL0364, scale-bar = 20 µm); 7. CUT-Z-GDC. TLM view showing stomatal complexes (SL1222, scale-bar = 50 µm); 8. CUT-Z- GDC. TLM detail of single stomatal complex (SL1222, scale-bar = 20 µm).

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Guard cell pair outline ovate, not outlined by a Stomatal complexes. Stomatal complexes evenly clear anticlinal wall, at same level as subsidiary spread, isolated, randomly oriented, actinocytic, cells (stomatal complexes sunken so that outer size range bimodal, with distinct 'giant stomatal stomatal ledges do not protrude above normal epi- complexes' present. Subsidiary cells (4–6) irregu- dermal cells), length 18–20 m (medium), little polar larly-shaped, typically isodiametric, periclinal walls development between guard cells (guard cells thicker than over normal epidermal cells, smooth, appear as continuous ring). Outer stomatal ledge ornamented with 5-6 bands of fine ridges radiating sub circular, thinner than normal epidermal cells, from the pore. Guard cell pair outline circular, out- extending over whole stoma, with an elliptical pore. lined by a well-defined anticlinal wall, length c. 20 Epidermal Cells. Epidermal cell flanges clearly visi- m (medium), underthrust by subsidiary cells (sub- ble using TLM, normal cells elongate (cells over sidiary cell anticlinal walls bisect outline of the both major and fine venation more elongate), guard cells in TLM view), little polar development approximately the same size as the stomata, anti- between guard cells (guard cells appear as contin- clinal walls curved to wavy, unbuttressed, inner uous ring). Outer stomatal ledge sub circular, thin- and outer surfaces smooth, unornamented. ner than normal epidermal cells, extending over whole stoma, with a broad, sub-circular pore. Indumentum. Scars of deciduous trichomes sparse, simple, inserted between (4–7) epidermal Epidermal Cells. Epidermal cell flanges clearly visi- cells modified into a distinct ring of foot cells, with ble using TLM, normal cells isodiametric (cells over thickened periclinal walls, scar diameter much veins not distinguished by shape), approximately smaller than a normal epidermal cell. the same size as the stomata; anticlinal walls curved to wavy, unbuttressed, inner and outer sur- Distinguishing features. One of a group of taxa faces smooth, unornamented. which have relatively small stomatal complexes without a distinct rim around the guard cells, and Indumentum. Attachment scars of deciduous tri- where the flanges of the subsidiary cells some- chomes common, scattered over venal and non- times project over the stomatal outline. CUT-Z-FJI venal regions, inserted between (8–11) epidermal is distinct in having unbuttressed epidermal cell cells modified by tangential divisions to form a sub- walls, and curved subsidiary cell anticlinal walls. circular zone of foot cells, scar diameter similar in CUT-Z-EDD differs in having buttressed epidermal size to a normal epidermal cell. cells and straight subsidiary cell anticlinal walls. Distinguishing features. One of a group of taxa CUT-Z-ABF differs in having large diameter, hollow which have relatively small stomatal complexes trichome attachment scars, which are surrounded without a distinct rim around the guard cells, and by a ring of many foot cells, each distinctly smaller where the flanges of the subsidiary cells some- than the trichome diameter. times project over the stomatal outline. CUT-Z-ABF Identification. Suggested as Sapindaceae based is distinct in having large diameter, hollow trichome on the thin-walled stomata surrounded by subsid- attachment scars, which are surrounded by a ring iary cells which have prominent flanges which ter- of many foot cells, each distinctly smaller than the minate abruptly at the stomatal outline, and in trichome diameter. some cases bisect the outline. It may be compared Identification. Suggested as Sapindaceae based in a very broad way with extant Cupaniopsis anac- on the thin-walled stomata surrounded by subsid- ardioides (Fig. 30.1), although it differs in shape of iary cells which have prominent flanges which ter- the subsidiary cells (which are more elongate in the minate abruptly at the stomatal outline, and in fossil) and thickness of the subsidiary cell flanges some cases bisect the outline. It compares closely (thinner in the fossil). with extant Cupaniopsis anacardioides in terms of stoma shape, thin outer stomatal ledges, flanges of CUT-Z-ABF the subsidiary cells abruptly abutting against and Figure 31.1-31.4 sometimes project over the stomatal outline, and Reference Specimen and locality: SB1382, BL-32. thickened subsidiary cell periclinal walls (Fig. 30.1).

Referred specimens and occurrence: SB0879, GL- CUT-Z-CEI 01; SL2233, GL-12; SL2875, GL-20; SL3106, GL- Figure 31.5-31.6 24; SL2951, GL-25; SL2995, GL-27; SL3019, GL- 29; SL1218, BL-30; SL1855, Sthd-088; SL1861, Reference Specimen and locality: SL0364, BL-09. Sthd-089.

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Stomatal complexes. Stomatal complexes evenly CUT-Z-GDC spread, isolated, randomly oriented, cyclocytic, all Figure 31.7-31.8 some, or none of the subsidiary cells have been Reference Specimen and locality: SL1222, BL-30. modified by a tangential division, size range unimo- dal. Subsidiary cells (5–6) typically elongate tan- Stomatal complexes. Stomatal complexes in are- gential to stoma, or wedge-like, often with distinct oles, isolated (but very rare networking noted), ran- polar cells (at right angles to the stomatal axis) domly oriented, actinocytic, or staurocytic, size which may 'enclose' the lateral cells, periclinal range unimodal. Subsidiary cells (4–5) isodiamet- walls thicker than over normal epidermal cells, ric, periclinal walls thicker than over normal epider- smooth, ornamented with groups of ridges radiat- mal cells, smooth, unornamented. Guard cell pair ing from the stoma. Guard cell pair outline circular, outline circular, or ovate, outlined by a well-defined outlined by a well-defined anticlinal wall, length 20– anticlinal wall, at same level as subsidiary cells 25 m (medium), underthrust by subsidiary cells (exposed on surface), length 28–32 m (medium), (subsidiary cell anticlinal walls bisect outline of the with prominent T-piece thickenings at polar ends. guard cells). Outer stomatal ledge sub circular, Outer stomatal ledge sub circular, much thicker same thickness as normal epidermal cells, extend- than normal epidermal cells, extending over whole ing over centre of stoma, with a broad, sub-circular stoma, with a broad, sub-circular pore. pore. Epidermal Cells. Epidermal cell flanges clearly visi- Epidermal Cells. Epidermal cell flanges clearly visi- ble using TLM, normal cells elongate (cells over ble using TLM; normal cells elongate (cells over major veins more elongate, but those over finer major veins more elongate); approximately the veins inconsistently so), approximately the same same size as the stomata; anticlinal walls straight, size as the stomata, anticlinal walls markedly sinu- unbuttressed, smoothly textured except for an ous, unbuttressed, smoothly textured, unorna- irregular ridge on either side of stoma. mented. Indumentum. Attachment scars of deciduous tri- Indumentum. Attachment scars of deciduous tri- chomes common, scattered over venal and non- chomes common, scattered over venal and non- venal regions, inserted between (6–10) epidermal venal regions, inserted between (5–7) epidermal cells modified into radially elongate foot cells, with cells modified by several tangential divisions, with no distinct thickening of walls, scar diameter much a massively thickened rim, scar diameter much smaller than a normal epidermal cell. smaller than a normal epidermal cell. Distinguishing features. CUT-Z-CEI can be com- Identification. Suggested as Sapindaceae based pared with CUT-Z-ABF, CUT-Z-EDD, and CUT-Z- on broad similarity with extant species such as FJI, where the subsidiary cell flanges sometimes Rhysotoechia bifoliolata inn terms of stomatal and intersect the outline of the guard cells. However, in subsidiary cell shape and degree of thickening those cases, the guard cell outline is not rimmed, (Fig. 30.3). and the outer stomatal ledges are much thinner. CUT-Z-CEI has clearly rimmed guard cells, and Ericaceae Jussieu 1789 well-developed outer stomatal ledges. The sto- Family identification: The fossil record of the Epac- matal complexes also have distinctly more subsid- ridaceae, which is now regarded as included within iary cells. the Ericaceae (Angiosperm Phylogeny Group Identification. Suggested as Sapindaceae based 2003) was summarized by Jordan and Hill (1996) on subsidiary cells which have prominent flanges and leaf cuticle of some fossil and extant species which terminate abruptly at the stomatal outline, was illustrated by Jordan and Hill (1995). The sto- and in some cases bisect the outline. The stomata matal complexes range from cyclocytic, to para- have a more developed outer stomatal ledge than cytic to anomocytic and are generally aligned. The CUT-Z-EDD, CUT-Z-FJI, or CUT-Z-ABF, but may epidermal cells are often (not always) highly sinu- still be compared with some extant Sapindaceae, ous. The tree-heath Trochocarpa has generally for instance Diploglottis diphyllostegia Mueller, aligned anomocytic stomata and highly sinuous F.J.H. von ex Bailey (1885), (Fig. 30.2), Rhysotoe- epidermal cells. For comparison two species of chia bifoliolata Radlk. (1879) (Fig. 31.3), and Mis- extant Trochocarpa, T. laurina Brown (1814) and T. chocarpus stipitatus S.T. Reynolds (1985) (Fig. bellendenkerensis Domin (1930) are illustrated 30.4). (Figs 32.5 -32.6).

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Figure 32. 1-4. Fossil and extant Ericaceae. 1. CUT-Z-ECD, TLM view showing generally stomatal complexes (SL1929, scale aligned -bar = 50 µm); 2. CUT-Z-ECD, TLM detail of single stomatal complex (SL1929, scale-bar = 20 µm); 3. CUT-Z-ECD, SEM view of outer cuticular surface showing (upper centre) a single stomatal complex with prominent outer stomatal ledges, and 4-5 smaller structures which are probably trichome attachment scars (S-1077, scale-bar = 20 µm); 4. CUT-Z-ECD, SEM view of inner cuticular surface showing three stomatal complexes. Note buttressing of walls right up to the margins of the guard cells (S-1077, scale-bar = 20 µm); 5. Extant Trochocarpa lau- rina, TLM view showing aligned stomata and highly sinuous epidermal cell walls (AQ610328, scale-bar = 50 µm); 6. Extant T. bellendenkerensis, TLM view showing aligned stomata but little sinuosity of epidermal cells (AQ188409, scale-bar = 50 µm).

CUT-Z-ECD under TLM, periclinal walls of same thickness as Figure 32.1-32.4 normal epidermal cells, smooth, unornamented. Reference Specimen and locality: SL1929, Sthd- Guard cell pair outline ovate, outlined by a well- 058. defined anticlinal wall, at same level as subsidiary cells (exposed on surface), length 20–25 m Stomatal complexes. Stomatal distribution over leaf (medium), with prominent T-piece thickenings at surfaces hypostomatic, stomatal complexes evenly polar ends. Outer stomatal ledge sub circular, thin- spread, isolated, showing a clear trend towards ner than normal epidermal cells, extending over alignment, cyclocytic, size range unimodal. Subsid- whole stoma, with a broad, sub-circular pore. iary cells (5–7) irregularly-shaped, difficult to count

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Figure 33. 1-8. Extant and fossil (CUT-Z-AJG, CUT-Z-CDH) Myrsinaceae. 1. Extant Ardisia quinquegona, TLM view showing stomatal complexes and (upper left) a trichome attachment scar (AQ433196, scale-bar = 50 µm); 2. Extant A. humilis, TLM view showing stomatal complexes and (upper left) a trichome attachment scar (AQ452160, scale-bar = 50 µm); 3. Extant A. brevipedata, TLM view showing stomatal complexes (AQ366276, scale-bar = 50 µm); 4. Extant A. bakeri, TLM detail of single stomatal complex. OPH3334, AQ440726, scale-bar = 20 µm); 5. CUT-Z-AJG, TLM view showing stomatal complexes and (upper left) a trichome attachment scar (SB1290, scale-bar = 50 µm); 6. CUT-Z- AJG, TLM detail of single stomatal complex (SB1290, scale-bar = 20 µm); 7. CUT-Z-CDH, TLM view showing stomatal complexes and (upper right) a trichome attachment scar (SL0241, scale-bar = 50 µm); 8. CUT-Z-CDH, TLM detail of two stomatal complexes (SL0241, scale-bar = 20 µm).

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Epidermal Cells. Epidermal cell flanges often indis- Stomatal complexes. Stomatal complexes evenly tinct using TLM, normal cells isodiametric (cells spread, isolated, randomly oriented, anisocytic, over veins not distinguished by shape), approxi- heliocyctic, size range unimodal. Subsidiary cells mately the same size as the stomata, anticlinal (3–4) irregularly-shaped, periclinal walls of same walls markedly sinuous, buttressed, inner surfaces thickness as normal epidermal cells, smooth, orna- slightly granular, outer surfaces strongly granular, mented with many ridges which tend to flow around unornamented. the stoma. Guard cell pair outline ovate, outlined by a well-defined anticlinal wall, at same level as Indumentum. Glabrous. subsidiary cells (exposed on surface), length 22– Glands. Possible sparse multicellular glands 23 m (medium). Outer stomatal ledge sub circular, present. same thickness as normal epidermal cells, extend- ing over inner edge of stoma, with an elliptical Distinguishing features. Distinguished from CUT- pore. M-EFJ in having stomatal complexes which are approximately twice as large. Lid cells, which are Epidermal Cells. Epidermal cell flanges clearly visi- present in CUT-M-EFJ, have not been found in ble using TLM, normal cells highly variable from CUT-Z-ECD. isodiametric to elongate (cells over veins not distin- guished by shape), distinctly larger than the sto- Myrsinaceae Brown 1814 mata, anticlinal walls curved to wavy, unbuttressed, smoothly textured, unornamented. Family identification: The typical morphology is an anisocytic stomata associated with peltate tri- Indumentum. Attachment scars of deciduous tri- chomes (like Myricaceae). There is often an orna- chome common, not clearly rimmed with thicker mentation of fine ridging, and this is often restricted cuticle. to the subsidiary cells. For comparison four spe- Distinguishing features. Distinguished from CUT-Z- cies of extant Ardisia are illustrated; A. quinqueg- CDH by the absence of a thick rim around the tri- ona Blume, A. humilis Vahl (1794), A. brevipedata chome scars. Mueller (1868), and A. bakeri White (1942) (Figs 33.1-33.4). CUT-Z-CDH Figure 33.7-33.8 Key to Mysinaceae Reference Specimen and locality: SL0241, BL-04. 1. Cuticle unornamented. CUT-Z-AJG Stomatal complexes. Stomatal complexes evenly 1. Cuticle ornamented. 2. spread, isolated, randomly oriented, anisocytic, 2. Ornamentation consisting of a single ridge on heliocyctic, size range unimodal. Subsidiary cells either side of the stoma. CUT-Z-CGG (3–4) irregularly-shaped, periclinal walls of same 2. Ornamentation consisting of multiple ridges. 3. thickness as normal epidermal cells, smooth, orna- mented with ridges concentric about the stoma. 3. Ornamentation of fine ridges extending all over Guard cell pair outline ovate, outlined by a well- cuticle (epidermal and subsidiary cells). 4. defined anticlinal wall, at same level as subsidiary 3. Ornamentation of fine ridges restricted to sub- cells (exposed on surface), length 20–25 m sidiary cells. CUT-Z-CDH (medium). Outer stomatal ledge sub circular, thin- 4. Stomatal complexes slightly depressed, portions ner than normal epidermal cells, extending over of subsidiary cell cuticle extending over parts of the inner edge of stoma, with an elliptical pore. outer stomatal ledge. CUT-Z-EDF Epidermal Cells. Epidermal cell flanges clearly visi- 4. Stomatal complexes not depressed, not ble using TLM, normal cells highly variable from obscured by subsidiary cells. CUT-Z-CGC isodiametric to elongate (cells over veins not distin- guished by shape), distinctly larger than the sto- CUT-Z-AJG mata, anticlinal walls curved to wavy, unbuttressed, Figure 33.5-33.6 smoothly textured ornamented with fine ridges over subsidiary cells which flow around the stoma. Reference Specimen and locality: SB1290, Mata- 03. Indumentum. Attachment scars of deciduous tri- chomes common, restricted to regions over major Referred specimens and occurrence: SL3303, veins, inserted between (5–7) epidermal cells mod- Sthd-078. ified into radially elongate foot cells, and with a dis-

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Figure 34. 1-8. Fossil Myrsinaceae (CUT-Z-CGG, CUT-Z-CGC, and CUT-Z-EDF). 1. CUT-Z-CGG, TLM view showing three stomatal complexes (SL0097, scale-bar = 50 µm); 2. CUT-Z-CGG, TLM detail of single stomatal complex (SL0097, scale-bar = 20 µm); 3. CUT-Z-CGG, SEM view of inner cuticular surface showing a single stomatal complex (S-1109, scale-bar = 20 µm); 4. CUT-Z-CGG, SEM view of outer cuticular surface showing a single stomatal complex with sunken outer stomatal ledges and a thin, discontinuous rim (S-1109, scale-bar = 20 µm); 5. CUT-Z-CGC, TLM view showing stomatal complexes and (upper right) a trichome attachment scar (SL0324, scale-bar = 50 µm); 6. CUT- Z-CGC, TLM detail of single stomatal complex (SL0324, scale-bar = 20 µm); 7. CUT-Z-EDF, TLM view showing sto- matal complexes and (centre left) a trichome attachment scar (SL2070, scale-bar = 50 µm); 8. CUT-Z-EDF, TLM detail of single stomatal complex and trichome attachment scar (SL2070, scale-bar = 20 µm).

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tinct rim, scar diameter similar in size to a normal heliocyctic, size range unimodal. Subsidiary cells epidermal cell. (3) irregularly-shaped, periclinal walls of same Distinguishing features. Distinguished from CUT-Z- thickness as normal epidermal cells, smooth, orna- EDF in that the stomatal complexes not depressed mented with many ridges which tend to flow around (so outer stomatal ledge is not obscured by por- the stoma. Guard cell pair outline ovate, outlined tions of subsidiary cell cuticle), and both anticlinal by a well-defined anticlinal wall, length 22–23 m and radial walls of foot cells are thickened. Distin- (medium); at same level as subsidiary cells guished from CUT-Z-AJG by the presence of a dis- (exposed on surface). Outer stomatal ledge sub tinct poral rim around the trichome scar. circular, thinner than normal epidermal cells, extending over inner edge of stoma, with an ellipti- CUT-Z-CGG cal pore. Figure 34.1-34.4 Epidermal Cells. Epidermal cell flanges clearly visi- Reference Specimen and locality: SL0097, BL-08. ble using TLM; normal cells elongate (cells over veins not distinguished by shape); distinctly larger Referred specimens and occurrence: SB0870, BL- than the stomata; anticlinal walls markedly sinu- 30; SL3171, BL-33; SL1962, Sthd-054. ous; unbuttressed; smooth, ornamented with 'flow- Stomatal complexes. Stomatal complexes evenly ing' pattern of many fine ridges. spread, isolated, randomly oriented, anisocytic, Indumentum. Attachment scars of deciduous tri- heliocyctic, size range unimodal. Subsidiary cells chomes common; scattered over venal and non- (3–4) of varying size, curved and generally tangen- venal regions, inserted over a single epidermal cell tial to the guard cells, periclinal walls thicker than with a strongly thickened periclinal wall and a cen- over normal epidermal cells, smooth, ornamented tral pore, epidermal cells around trichome attach- with 1–2 ridges parallel with, and on either side of ment scar (6–7) unmodified, scar diameter similar the stoma. Guard cell pair outline ovate, outer mar- in size to a normal epidermal cell. gin obscured under TLM by surface ornamentation, at same level as subsidiary cells (exposed on sur- Distinguishing features. Distinguished from CUT-Z- face), length 20–30 m (medium). Outer stomatal EDF and CUT-Z-CDH in having an ornamentation ledge sub circular, thinner than normal epidermal of fine ridges extending all over cuticle (both epi- cells, extending over whole stoma, with a broad, dermal and subsidiary cells). sub-circular pore. CUT-Z-EDF Epidermal Cells. Epidermal cell flanges clearly visi- Figure 34.7-34.8 ble using TLM, normal cells elongate (cells over major veins more elongate), distinctly larger than Reference Specimen and locality: SL2070, Sthd- the stomata, anticlinal walls markedly sinuous, 047. unbuttressed, inner surfaces very finely granular, Referred specimens and occurrence: SL1875, outer surfaces smooth, ornamented with an irregu- Sthd-078. lar ridge on either side of the stoma. Stomatal complexes. Stomatal complexes evenly Indumentum. Glabrous. spread, isolated, possibly slightly aligned with one Distinguishing features. Distinguished from other another, anisocytic, heliocyctic, size range unimo- taxa with anisocytic stomatal complexes in having dal. Subsidiary cells (3) of varying size, curved and sinuous anticlinal walls, and ornamentation generally tangential to the guard cells, periclinal restricted to a ridge or two on either side of the sto- walls of same thickness as normal epidermal cells, mata, and being glabrous. smooth, ornamented with fine ridges, often radiat- ing laterally from guard cells, and partially obscur- CUT-Z-CGC ing them. Guard cell pair outline ovate, outer Figure 34.5-34.6 margin obscured under TLM by surface ornamen- tation, length 25–34 m (medium), overarched irreg- Reference Specimen and locality: SL0324, Harli- ularly by subsidiary cells, little polar development wich-3. between guard cells (guard cells appear as contin- Referred specimens and occurrence: SL3198, BL- uous ring). Outer stomatal ledge sub circular, thin- 32. ner than normal epidermal cells, extending over Stomatal complexes. Stomatal complexes evenly inner edge of stoma, with an elliptical pore. spread, isolated, randomly oriented, anisocytic,

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Figure 35. 1-8. Fossil and extant Grisselinia (Grisseliniaceae). 1. CUT-Z-GBF, TLM view showing two stomatal com- plexes. The cuticle is very thick and stains darkly (SL4860, scale-bar = 50 µm); 2. CUT-Z-GBF, TLM detail of single stomatal complex (SL4860, scale-bar = 20 µm); 3. CUT-Z-GBF, SEM view of outer cuticular surface showing a sin- gle stomatal complex with prominent outer stomatal ledges which are surrounded by sharp and deep grooves (S- 1582, scale-bar = 20 µm); 4. CUT-Z-GBF, SEM view of inner cuticular surface showing a single stomatal complex (S-1582, scale-bar = 20 µm); 5. Extant Griselinia lucida, TLM view showing three stomatal complexes (OTA31637, scale-bar = 50 µm); 6. Extant G. lucida, TLM detail of single stomatal complex (OTA31637, scale-bar = 20 µm); 7. Extant G. l uc ida , SEM view of outer cuticular surface showing stomatal complexes with prominent outer stomatal ledges and rims (S-1588, OTA31637, scale-bar = 20 µm); 8. Extant G. l uc ida , SEM view of inner cuticular surface showing a single stomatal complex (S-1588, OTA31637, scale-bar = 20 µm).

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Figure 36. 1-4. Fossil and extant Argophyllum (Grossulariaceae). 1. CUT-Z-ACD, TLM view showing stomatal com- plexes and papillae or trichome bases (SB1366, scale-bar = 50 µm); 2. CUT-Z-ACD, TLM detail of single stomatal complex (SB1366, scale-bar = 20 µm). Note slight overlapping of cuticle ridge onto stomata; 3. Extant Argophyllum nullamense, TLM view showing several stomata interspersed with darker staining trichomes (AQ268618, scale-bar = 50 µm); 4. Extant A. nullamense, TLM view showing two stomata surrounded and partially sunken below irregular ridges of cuticle and separated by dark trichomes (AQ268618, scale-bar = 20 µm).

Epidermal Cells. Epidermal cell flanges clearly visi- insertion points are thickened to form a poral ring ble using TLM, normal cells highly variable from (no thickening of radial walls). isodiametric to elongate (cells over veins not distin- guished by shape), approximately the same size Grisseliniaceae Cunningham 1839 as the stomata, anticlinal walls curved to wavy, CUT-Z-GBF (Grisselinia sp.) unbuttressed, smoothly textured ornamented with Figure 35.1-35.4 'flowing' pattern of many fine ridges. Reference Specimen and locality: SL4860, BL-32. Indumentum. Attachment scars of deciduous tri- Stomatal complexes. Stomatal complexes evenly chomes common, scattered over venal and non- spread, isolated, randomly oriented, cyclocytic, venal regions, inserted between epidermal cells, developmentally tangenticytic, size range unimo- epidermal cells around trichome attachment scar dal. Subsidiary cells (mostly appears to be 5, but modified into radially-elongate foot-cells (some tan- difficult to count) of varying size, curved and gener- gential divisions result in concentric rings), with a ally tangential to the guard cells, periclinal walls massively thickened rim, scar diameter much slightly thicker than over normal epidermal cells, smaller than a normal epidermal cell. smooth, unornamented. Guard cell pair outline Distinguishing features. Distinguished from CUT-Z- ovate, outlined by a well-defined anticlinal wall, at CDH in that the stomatal complexes are slightly same level as subsidiary cells (exposed on sur- depressed, with portions of subsidiary cell cuticle face), length 68–83 m (large). Outer stomatal ledge extending over parts of the outer stomatal ledge. In elliptical, much thicker than normal epidermal cells, addition, only the anticlinal walls around trichome extending over whole stoma, surrounded by a deep, narrow groove, pore elliptical.

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Figure 37. 1-8. Papillate Cuticle: CUT-Z-ABA and CUT-Z-ABC. 1. CUT-Z-ABA, TLM view showing stomatal com- plexes surrounded by a ring of papillate subsidiary cells. (SB1335, scale-bar = 50 µm); 2. CUT-Z-ABA, TLM detail of single stomatal complex (SB1335, scale-bar = 20 µm); 3. CUT-Z-ABA, SEM view of outer cuticular surface showing stomata surrounded by papillate subsidiary cells as well as papillae on some epidermal cells (S-324, scale-bar = 20 µm); 4. CUT-Z-ABA, SEM view of inner cuticular surface showing a single stomatal complex. Note about 12 subsid- iary cells (S-324, scale-bar = 10 µm); 5. CUT-Z-ABC, TLM view showing stomatal complexes and trichome attach- ment scars (a large one is at lower left). The papillae are restricted to the subsidiary cells and a few other epidermal cells. Note also the position of a giant stomatal complex over a vein at centre left (SB1334, scale-bar = 50 µm); 6. CUT-Z-ABC, TLM detail of stomatal complexes showing papillae projecting over the stomatal pore (SB1334, scale- bar = 20 µm); 7. CUT-Z-ABC, SEM view of outer cuticular surface showing papillae almost blocking the stomatal pore (S-309, scale-bar = 10 µm); 8. CUT-Z-ABC, SEM of inner cuticular surface showing large trichome attachment scars and smaller stomatal complexes (S-309, scale-bar = 20 µm).

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Epidermal Cells. Epidermal cell flanges clearly visi- TAXA OF UNCERTAIN FAMILY ble using TLM, with a smooth junction with the per- Group 1. Papillae present iclinal walls, normal cells isodiametric (cells over veins not distinguished by shape), distinctly smaller Key to papillate cuticle than the stomata, anticlinal walls straight, unbut- 1. Cuticle hisute (trichomes persistent). 2. tressed, inner and outer surfaces smooth, unorna- mented. 1. Trichomes deciduous. 3. Indumentum. Glabrous. 2. Papillae simple and smooth. CUT-Z-ECH Identification: CUT-Z-GBF is identified as Griselinia 2. Papillae peltate. CUT-Z-ABI because of the basic similarity, especially with 3. Stomatal complexes scattered.4. respect to the large stomata and prominent outer 3. Stomatal complexes in clearly defined patches. stomatal ledges (Fig. 35.5-35.8). The extant spe- CUT-Z-ABD cies G. lucida Forst.f. (1786), has peristomatal rings which are not present on the fossil. 4. Papillae restricted to subsidiary cells CUT-Z- ABA Argophyllaceae Takhtajan 1987 4. Papillae on both subsidiary and normal epider- CUT-Z-ACD (Argophyllum sp.) mal cells. 5. Figure 36.1-36.2 5. Papillae much smaller than the epidermal cell. Reference Specimen and locality: SB1366, BL-32. CUT-Z-ABC Stomatal complexes. Stomatal complexes evenly 5. Papillae comparable in size to the epidermal spread, isolated, randomly oriented, number of cell. CUT-Z-ABB subsidiary cells unclear, size range unimodal. Irregularly-shaped, periclinal walls of same thick- CUT-Z-ABA ness as normal epidermal cells, smooth, orna- Figure 37.1-37.4 mented with irregular rims of cuticle, which partially Reference Specimen and locality: SB1335, BL-32. project over the OSL. Guard cell pair outline circu- lar, at same level as subsidiary cells, outer margin Referred specimens and occurrence: SL0281, BL- obscured under TLM by surface ornamentation, 01; SL0240, BL-04; SL1279, BL-05; SL0365, BL- length c. 35 m (medium). Outer stomatal ledge sub 09; SL3088, BL-28; SL3178, BL-33; SL2097, GL- circular, same thickness as normal epidermal cells, 07; SL2140, GL-08; SL2219, GL-12; SL2895, GL- extending over whole stoma, with an elliptical pore. 20; SL2904, GL-21; SL2929, GL-22; SL3109, GL- 24; SL2965, GL-25; SL0383, Mata-01; SB0845, Epidermal Cells. Epidermal cell flanges not clearly Mata-06. visible under TLM because of surface thickenings, approximately the same size as the stomata, (cells Stomatal complexes. Stomatal complexes in are- over major veins more elongate), anticlinal walls oles, isolated, randomly oriented, laterocyclic, size curved to wavy, unbuttressed, ornamented with range unimodal. Subsidiary cells (9–11) isodiamet- irregular, broad concentric ridges which cut irregu- ric, periclinal walls of same thickness as normal larly across the subsidiary cells. epidermal cells, granular. With a single papilla which projects horizontally over the stomatal pore. Indumentum. Raised, thick, peg-like attachment Unornamented. Guard cell pair outline difficult or scars of deciduous trichomes common, scattered impossible to see in TLM view, sunken below sub- over venal and non-venal regions, simple, inserted sidiary cells and epidermal cells, outer margin between (c. 7) unmodified epidermal cells, scar obscured under TLM by surface topography, length diameter similar in size to a normal epidermal cell. 20–35 m (medium). Distinguishing features. Distinguished by the orna- Epidermal Cells. Epidermal cell flanges clearly visi- mentation of irregular ridges and trichome attach- ble using TLM, normal cells isodiametric (cells over ment scars. major veins more elongate), approximately the Identification. CUT-Z-ACD can be compared same size as the stomata, anticlinal walls curved to directly with Argophyllum nullumense R.T.Baker wavy, unbuttressed, inner surfaces very slightly (1899) especially with respect to the thick, irregular granular, outer surfaces smooth, unornamented, ridges of cuticle and prominent trichome attach- occasionally papillate. ment scars (Fig. 36.3-36.4).

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Figure 38. 1-8. Papillate Cuticle: CUT-Z-ABI and CUT-Z-ECH. 1. CUT-Z-ABI, TLM view showing stomatal complexes entirely obscured by papillae and trichomes (SB1299, scale-bar = 50 µm); 2. CUT-Z-ABI, TLM detail showing papillae (SB1299, scale-bar = 20 µm); 3. CUT-Z-ABI, SEM view of outer cuticular surface showing papillae obscuring stomata (S-1217, scale-bar = 20 µm); 4. CUT-Z-ABI, SEM of inner cuticular surface showing stomatal complex (upper left) (S- 1217, scale-bar = 20 µm); 5. CUT-Z-ECH, TLM view showing trichomes, papillae, and obscured stomatal complexes (SB1208, scale-bar = 50 µm); 6. CUT-Z-ECH, TLM view showing a single stomatal complex with papillae overarching the stomatal pore (SL1208, scale-bar = 20 µm); 7. CUT-Z-ECH, SEM view of outer cuticular surface showing papillae and trichomes (S-1209, scale-bar = 20 µm); 8. CUT-Z-ECH, SEM view of inner cuticular surface showing (left) a sin- gle stomatal complex and (right) a trichome attachment scar (S-1209, scale-bar = 20 µm).

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Indumentum. Attachment scars of deciduous tri- papillae are found on subsidiary cells and com- chomes common, scattered over venal and non- monly also on some normal epidermal cells. venal regions but slightly more common over veins, inserted between (7–9) epidermal cells mod- CUT-Z-ABI ified only slightly, with a thickened rim. Scar diame- Figure 38.1-38.4 ter much smaller than a normal epidermal cell. Reference Specimen and locality: SB1299, Mata- Smooth papillae present. 03. Distinguishing features. Distinguished from CUT-Z- Referred specimens and occurrence: SL3167, BL- ABC in that papillae are restricted to subsidiary 33. cells. Stomatal complexes. Stomatal complexes appar- CUT-Z-ABC ently in well-defined areoles. All details of stomata Figure 37.5-37.8 obscured under TLM by surface ornamentation and trichomes. Reference Specimen and locality: SB1334, BL-32. Epidermal Cells. Details of normal epidermal cells Referred specimens and occurrence: SL0084, BL- obscured under TLM because of surface thicken- 08; SL1182, GL-01; SL2654, GL-04; SL2090, GL- ings, but cells over some veins are distinguished 07; SL1760, Sthd-012. by being unornamented. Stomatal complexes. Stomatal complexes in are- Indumentum. Hirsute and with dense peltate exten- oles, isolated, randomly oriented, cyclocytic, size sions of epidermal cells. Trichomes abundant, range bimodal, with distinct 'giant stomatal com- uniseriate, apparently more common over venal plexes' present (restricted to venal regions). Sub- areas. sidiary cells (6–10) isodiametric, periclinal walls of same thickness as normal epidermal cells, smooth. CUT-Z-ECH With a single papilla which projects horizontally Figure 38.5-38.8 over the stomatal pore. Unornamented. Guard cell pair outline difficult or impossible to see in TLM Reference Specimen and locality: SL1208, GL-01. view, sunken below subsidiary cells, outer margin Referred specimens and occurrence: SL3156, GL- obscured under TLM by surface topography, length 01; SB1208, Mata-01. c. 20 m (medium). Stomatal complexes. Stomatal complexes in are- Epidermal Cells. Epidermal cell flanges clearly visi- oles, isolated, number of subsidiary cells unclear ble using TLM, normal cells isodiametric (cells over because of obscuring papillae, size range bimodal, both major and fine venation more elongate), with distinct 'giant stomatal complexes' present. approximately the same size as the stomata, anti- Subsidiary cells (4–6) isodiametric, periclinal walls clinal walls curved to wavy, unbuttressed, inner of same thickness as normal epidermal cells, surfaces slightly granular, outer surfaces slightly smooth, each with a single papilla which projects rough, unornamented. horizontally over the stomatal pore. Guard cell pair Indumentum. Attachment scars of deciduous tri- outline difficult or impossible to see in TLM view as chomes common, restricted to regions over veins, obscured by surface topography, but appears to be inserted between (6–11) epidermal cells modified sunken with respect to subsidiary and epidermal into radially elongate foot cells, insertion ‘peg’ cells. diameter much smaller than a normal epidermal Epidermal Cells. Epidermal cell flanges clearly visi- cell, but the trichome then expands abruptly above ble using TLM, normal cells isodiametric (cells over the insertion. Papillae in patches, generally both major and fine venation more elongate), restricted to regions surrounding stomatal com- approximately the same size as the stomata, anti- plexes, one papilla per cell, formed by the projec- clinal walls straight, unbuttressed, inner surfaces tion of a discrete region within the boundaries of smooth, outer surfaces rough, unornamented. the epidermal cell, smooth, much smaller than the Indumentum. Hirsute and papillate, trichomes com- epidermal cell, oriented vertically. mon, scattered over venal and non-venal regions, Distinguishing features. Distinguished from CUT-Z- simple, inserted between (6-11) unmodified or ABB by the smaller papillae – much smaller than sometimes radially elongate epidermal cells, inser- the epidermal cells, and from CUT-Z-ABA in that tion peg diameter very small, but trichome base similar in size to a normal epidermal cell. The peri-

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Figure 39. 1-8. Papillate Cuticle: CUT-Z-ABB and CUT-Z-ABD. 1. CUT-Z-ABB, TLM view showing stomatal com- plexes obscured by papillae and, at lower centre, a trichome attachment scar (SB1333, scale-bar = 50 µm); 2. CUT-Z- ABB, TLM detail of stomatal complexes obscured by papillae (SB1333, scale-bar = 20 µm); 3. CUT-Z-ABB, SEM view of inner cuticular surface showing stomatal complexes and a trichome attachment scar (S-338, scale-bar = 20 µm); 4. CUT-Z-ABB, SEM view of outer cuticular surface showing papillae surrounding stomatal pores (S-338, scale-bar = 10 µm); 5. CUT-Z-ABD, TLM view showing a patch of stomatal complexes obscured by papillae (SB1332, scale-bar = 50 µm); 6. CUT-Z-ABD, TLM detail of a single stomatal complex (SB1332, scale-bar = 20 µm); 7. CUT-Z-ABD, SEM view of outer cuticular surface showing patches of papillae which are covering stomatal complexes (S-360, scale-bar = 0.1 mm); 8. CUT-Z-ABD, SEM view of inner cuticular surface showing a single stomatal complex. Note the very narrow guard cell pair and the coarsely granular texture of the subsidiary and epidermal cell walls (S-360, scale-bar = 10 µm).

79 POLE: MIOCENE LEAF CUTICLE clinal surface of each epidermal cells projects 06; SL0026, BL-07; SL0098, BL-08; SL2467, BL- upwards as a single smooth papilla. 09; SL2250, BL-14; SL2271, BL-16; SL2292, BL- Distinguishing features. Distinguished from other 18; SL3052, BL-24; SL3076, BL-27; SL3083, BL- taxa with persistent trichomes in that the entire sur- 28; SL3093, BL-29; SL1224, BL-30; SL2517, BL- face of all epidermal and subsidiary cells are raised 31; SL1377, BL-32; SL2537, BL-33; SL1179, GL- up as papillae. 01; SL1187, GL-02; SL2658, GL-04; SL2190, GL- 10; SL2201, GL-11; SL2216, GL-12; SL2823, GL- CUT-Z-ABB 16; SL2838, GL-17; SL2861, GL-18; SL2883, GL- Figure 39.1-39.4 20; SL2899, GL-21; SL2918, GL-22; SL2939, GL- 23; SL3104, GL-24; SL2949, GL-25; SL2993, GL- Reference Specimen and locality: SB1333, BL-32. 27; SL3018, GL-29; SL1057, Mata-01; SL0334, Referred specimens and occurrence: SL1277, BL- Mata-23. 05; SL3081, BL-28; SL2141, GL-08; SL2173, GL- Stomatal complexes. Stomatal complexes in are- 09; SL2180, GL-10; SL2205, GL-11; SL2218, GL- oles, isolated, randomly oriented, cyclocytic, size 12; SL2849, GL-18; SL2890, GL-20; SL2943, GL- range unimodal. Subsidiary cells (4–6) isodiamet- 23; SL3101, GL-24; SL2968, GL-25; SL2975, GL- ric, periclinal walls of same thickness as normal 26. epidermal cells; with peltate scales. Unorna- Stomatal complexes. Stomatal distribution over leaf mented. Guard cell pair outline distinctly narrow surfaces unknown, randomly oriented, actinocytic, and almost rectangular, but difficult or impossible to size range bimodal, with distinct 'giant stomatal see in TLM view; overarched by subsidiary cells, complexes' present. Subsidiary cells (4–6) isodia- outer margin obscured under TLM by surface metric, periclinal walls of same thickness as normal topography; length c. 10 m (small). epidermal cells; smooth, each with a single papilla Epidermal Cells. Epidermal cell flanges clearly visi- that projects horizontally over the stomatal pore. ble using TLM; normal cells highly variable from Guard cell pair outline difficult or impossible to see isodiametric to elongate (cells over both major and in TLM view as obscured by surface topography. fine venation more elongate); distinctly larger than Epidermal Cells. Epidermal cell flanges clearly visi- the stomata; anticlinal walls curved to wavy; unbut- ble using TLM but only away from stomatal areas; tressed; inner surfaces coarsely granular, outer anticlinal walls curved to wavy (cells over major surfaces smooth, ornamented with short ridges veins more elongate); approximately the same size perpendicular or parallel to some stomatal com- as the stomata; unbuttressed; inner and outer sur- plexes. faces moderately granular, unornamented. Indumentum. Attachment scars of deciduous tri- Indumentum. Scars of deciduous trichomes and chomes common, restricted to regions over veins, papillae present. Trichome attachment scars abun- inserted between (9–11) epidermal cells modified dant, scattered over venal and non-venal regions, by tangential divisions to form a sub-circular zone inserted between (6–9) epidermal cells modified of foot cells, scar diameter similar in size to a nor- into radially elongate foot cells, scar diameter mal epidermal cell. much smaller than a normal epidermal cell. Papil- Distinguishing features. Distinguished by having lae in patches, generally restricted to regions sur- stomatal complexes in clearly defined patches and rounding stomatal complexes, with one smooth, which are entirely hidden below peltate cuticular discrete papilla per cell, comparable in size to the extensions. epidermal cel. Group 2. Stomatal complexes generally aligned Distinguishing features. Distinguished from CUT-Z- ABC by the larger papillae – comparable in size to Key to cuticle with Stomatal complexes the epidermal cell. generally aligned 1. Outer stomatal ledge elliptical, Guard cell pair CUT-Z-ABD outline difficult or impossible to see in TLM view. Figure 39.5-39.8 CUT-Z-AAE Reference Specimen and locality: SB1332, Mata- 1. Outer stomatal ledge sub circular, guard cell pair 03. outline ovate. CUT-Z-CGJ Referred specimens and occurrence: SL0286, BL- 01; SL0238, BL-04; SL1275, BL-05; SL2697, BL-

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Figure 40. 1-4. Cuticle with stomatal complexes generally aligned: CUT-Z-AAE and CUT-Z-CGJ. 1. CUT-Z-AAE, TLM view showing stomatal complexes aligned generally left to right (SB1367, scale-bar = 50 µm); 2. CUT-Z-AAE, TLM detail of single stomatal complex (SB1367, scale-bar = 20 µm).Note the narrowly elliptic stomatal pore; 3. CUT-Z- CGJ, TLM view showing stomatal complexes aligned generally left to right (SL0352, scale-bar = 50 µm); 4. CUT-Z- CGJ, TLM detail of single stomatal complex (SL0352, scale-bar = 20 µm).

CUT-Z-AAE straight, unbuttressed, very smoothly textured, Figure 40.1-40.2 unornamented. Reference Specimen and locality: SB1367, BL-32. Indumentum. Glabrous. Referred specimens and occurrence: SL0122, BL- 03. CUT-Z-CGJ Figure 40.3-40.4 Stomatal complexes. Stomatal complexes in are- oles, isolated, showing a clear trend towards align- Reference Specimen and locality: SL0352, BL-05. ment, cyclocytic, size range unimodal. Subsidiary Stomatal complexes. Stomatal complexes evenly cells (5–7) irregularly-shaped, difficult to count spread, isolated, with a tendency to be aligned with under TLM, periclinal walls of same thickness as one another at least in patches, anomocytic, size normal epidermal cells, smooth, unornamented. range bimodal, with distinct 'giant stomatal com- Guard cell pair outline difficult or impossible to see plexes' present. Subsidiary cells (4–5) irregularly- in TLM view, not outlined by a clear anticlinal wall, shaped, periclinal walls of same thickness as nor- at same level as subsidiary cells, length 17–21 m mal epidermal cells, smooth, unornamented. (medium). Outer stomatal ledge elliptical, same Guard cell pair outline ovate, outlined by a well- thickness as normal epidermal cells, extending defined anticlinal wall, length 27–30 m (medium), over inner edge of stoma. Pore narrowly elliptic. at same level as subsidiary cells (exposed on sur- Epidermal Cells. Epidermal cell flanges clearly visi- face), some wall development between guard cells, ble using TLM, normal cells isodiametric (cells over and sometimes development of T-piece thicken- both major and fine venation more elongate), dis- ings at the poles. Outer stomatal ledge sub circular, tinctly smaller than the stomata, anticlinal walls same thickness as normal epidermal cells, extend-

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Figure 41. 1-6. Cuticle with highly sinuous epidermal cells: CUT-Z-AAC and CUT-Z-AAD. 1. CUT-Z-AAC, TLM view showing stomatal complexes (SB1345, scale-bar = 50 µm); 2. CUT-Z-AAC, TLM detail of single stomatal complex (SB1345, scale-bar = 20 µm); 3. CUT-Z-AAC, SEM view of inner cuticular surface showing a single stomatal complex (S-339, scale-bar = 10 µm); 4. CUT-Z-AAC, SEM view of outer cuticular surface showing stomatal complexes with low outer stomatal ledges and slight concentric striae, in an otherwise smooth surface (S-339, scale-bar = 25 µm); 5. CUT-Z-AAD, TLM view showing stomatal complexes (SB1291, scale-bar = 50 µm); 6. CUT-Z-AAD, TLM detail of sin- gle stomatal complex (SB1291, scale-bar = 20 µm). ing over whole stoma, with a broad, sub-circular Group 3. Epidermal cells highly sinuous pore. Key to cuticle with highly sinuous epidermal Epidermal Cells. Epidermal cell flanges clearly visi- cells ble using TLM, normal cells highly variable from isodiametric to elongate (cells over major veins 1. Guard cell pair outline circular, clearly visible more elongate), approximately the same size as under TLM. CUT-Z-AAC the stomata, anticlinal walls straight, unbuttressed, 1. Guard cell pair outline ovate, but obscured by slightly rough, unornamented. the strong outer stomatal ledge. CUT-Z-AAD Indumentum. Glabrous. CUT-Z-AAC Distinguishing features. Glabrous cuticle with virtu- Figure 41.1-41.4 ally naked kidney-shaped guard cells. Reference Specimen and locality: SB1345, BL-32.

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Figure 42. 1-8. Cuticle with prominent peristomatal rings: CUT-Z-ADD, CUT-Z-EIG, and CUT-Z-FGA. 1. CUT-Z-ADD, TLM view showing three stomatal complexes (SB0843, scale-bar = 50 µm); 2. CUT-Z-ADD, TLM detail of single sto- matal complex. Note pronounced peristomatal ring (arrowed) (SB0843, scale-bar = 20 µm); 3. CUT-Z-ADD, SEM view of inner cuticular surface showing a single stomatal complex (S-1198, scale-bar = 50 µm); 4. CUT-Z-ADD, SEM view of outer cuticular surface showing a single stomatal complex with prominent outer stomatal ledges surrounded by a prominent rim (S-1198, scale-bar = 20 µm); 5. CUT-Z-EIG, TLM view showing two stomatal complexes (SL1764, scale-bar = 50 µm); 6. CUT-Z-EIG, TLM detail of single stomatal complex. Note peristomatal ring (arrowed) (SL1764, scale-bar = 20 µm); 7. CUT-Z-FGA, TLM view showing stomatal complexes (SL2016, scale-bar = 50 µm); 8. CUT-Z- FGA, TLM detail of single stomatal complex. Note peristomatal ring (arrowed) (SL2016, scale-bar = 20 µm).

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Referred specimens and occurrence: SL2597, BL- dermal cells, extending over inner edge of stoma, 01; SL2284, BL-18; SL1183, GL-01; SL1198, GL- with an elliptical pore. 02; SL2675, GL-04; SL2187, GL-10; SL2200, GL- Epidermal Cells. Epidermal cell flanges clearly visi- 11; SL2831, GL-17; SL3012, GL-28. ble using TLM, normal cells highly variable from Stomatal complexes. Stomatal complexes evenly isodiametric to elongate (cells over major veins spread, isolated, randomly oriented, cyclocytic, more elongate), approximately the same size as size range unimodal. Subsidiary cells (8–10, diffi- the stomata, anticlinal walls markedly sinuous, cult to count under TLM) isodiametric; periclinal strongly buttressed, smoothly textured ornamented walls of same thickness as normal epidermal cells; with ridges of irregular course. smooth; unornamented. Guard cell pair outline cir- Indumentum. Attachment scars of deciduous tri- cular; outlined by a well-defined anticlinal wall; chomes sparse, not noted over veins, inserted length 25–30 m (medium); at same level as subsid- between (7–8) epidermal cells modified by tangen- iary cells (exposed on surface); with prominent tial divisions to form a sub-circular zone of foot polar rods. Outer stomatal ledge sub circular, thin- cells, scar diameter much smaller than a normal ner than normal epidermal cells, extending over epidermal cell. whole stoma, with a broad, sub-circular pore. Distinguishing features. Distinguished from CUT-Z- Epidermal Cells. Epidermal cell flanges clearly visi- AAC in having ovate stomatal complexes and ble using TLM; normal cells irregular (cells over fewer (about 4) subsidiary cells. both major and fine venation elongate); approxi- mately the same size as the stomata; anticlinal Group 4. Stomatal complexes surrounded by a walls markedly sinuous; strongly buttressed; inner prominent, broad peristomatal ring surfaces slightly granular, outer surfaces smooth, Key to cuticle with prominent peristomal ring unornamented. 1. Peristomal ring margins sharply defined, epider- Indumentum. Attachment cars of deciduous tri- mal cells distinctly small than stoma. CUT-Z-ADD chomes sparse, restricted to regions over veins, simple, inserted between (6–9) epidermal cells 1. Peristomal ring margins smooth, epidermal cells modified by tangential divisions to form a sub-cir- of comparable size to the stoma. 2. cular zone of foot cells, scar diameter much 2. Outer stomatal ledge separated from the peristo- smaller than a normal epidermal cell. matal rim by a narrow groove. Stoma with promi- Distinguishing features. Distinguished from CUT-Z- nent thickenings of polar rods. CUT-Z-EIG AAD in having very round stomatal complexes and 2. Outer stomatal ledge separated from the peristo- many subsidiary cells. matal rim by a broad groove. Stoma without thick- enings of polar rods. CUT-Z-FGA CUT-Z-AAD Figure 41.5-41.6 CUT-Z-ADD Reference Specimen and locality: SB1291, Mata- Figure 42.1-42.4 03. Reference Specimen and locality: SB0843, GL-02. Referred specimens and occurrence: SL3295, BL- Referred specimens and occurrence: SL3137, GL- 04; SL2207, GL-12; SL2842, GL-18; SL3123, 01. Sthd-198. Stomatal complexes. Stomatal complexes evenly Stomatal complexes. Stomatal complexes evenly spread, isolated, randomly oriented, cyclocytic, all spread, isolated, randomly oriented, laterocyclic, or some, or none of the subsidiary cells have been cyclocytic, developmentally tangenticytic, size modified by a tangential division (developmentally range unimodal. Subsidiary cells (4–5) typically stephanocytic bicyclic), size range bimodal, with elongate tangential to the stoma, periclinal walls of distinct 'giant stomatal complexes' present (sepa- same thickness as normal epidermal cells, smooth, rated from other stomatal complexes by a distinct ornamented with groups of ridges radiating from stomatal-free zone). Subsidiary cells (3–5) typically the stoma. Guard cell pair outline ovate, but elongate tangential to stoma, periclinal walls of obscured by the strong outer stomatal ledge, at same thickness as normal epidermal cells, granu- same level as subsidiary cells (exposed on sur- lar, ornamented with a thick, sharply defined and face), length 32–38 m (medium). Outer stomatal slightly lumpy peristomatal rim. Guard cell pair out- ledge sub circular, same thickness as normal epi- line ovate, outer margin obscured under TLM by

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surface topography, length c. 50 m (large). Outer with a thick, smooth edged peristomatal rim. Guard stomatal ledge sub circular, same thickness as nor- cell pair outline difficult or impossible to see in TLM mal epidermal cells, extending over whole stoma, view, outer margin obscured under TLM by surface separated from the peristomatal rim by a deep, ornamentation, length 35–41 m (large-medium). narrow groove. Pore broad, sub-circular. Outer stomatal ledge sub circular, thicker than nor- Epidermal Cells. Epidermal cell flanges clearly visi- mal epidermal cells, extending over whole stoma, ble using TLM, normal cells isodiametric (cells over separated from peristomatal rim by a broad, shal- veins not distinguished by shape), distinctly smaller low groove. Pore broad, sub-circular. than the stomata, anticlinal walls curved to wavy, Epidermal Cells. Epidermal cell flanges somewhat unbuttressed, inner surfaces slightly granular, diffuse, normal cells elongate (unclear if cells over outer surfaces smooth, unornamented. veins are distinguished), approximately the same Indumentum. Glabrous. size as the stomata, anticlinal walls curved to wavy, unbuttressed, slightly granular, flanges reflected on Distinguishing features. Distinguished by the mas- the surface by discontinuous, broad ridges, other- sive peristomatal ring. wise unornamented .

CUT-Z-EIG Indumentum. Glabrous. Figure 42.5-42.6 Group 5. Persistent trichomes present Reference Specimen and locality: SL1764, Sthd- Key to cuticle with persistent trichomes 011. 1. Trichome width multicellular at base. CUT-Z- Stomatal complexes. Stomatal complexes evenly ABG spread, isolated, randomly oriented, cyclocytic, developmentally tangenticytic, size range unimo- 1. Trichome width single-celled at base. 2. dal. Subsidiary cells (4–5) isodiametric, periclinal 2. Cuticle surface ornamented with ridges. 3. walls of same thickness as normal epidermal cells, 2. Cuticle surface unornamented. 4. smooth, ornamented with a thick, smooth-edged peristomatal rim. Guard cell pair outline ovate, out- 3. Normal epidermal cells curved to wavy and lined by a well-defined anticlinal wall, at same level unbuttressed. CUT-Z-CDC as subsidiary cells (exposed on surface), length 3. Normal epidermal cells sinuous and buttressed. 42–43 m (large), with prominent thickenings of CUT-Z-FFE polar rods. Outer stomatal ledge sub circular, thin- 4. Stomatal complexes not anisocytic. 5. ner than normal epidermal cells, extending over whole stoma, with a broad, sub-circular pore, sepa- 4. Stomatal complexes often anisocytic. 6. rated from the peristomatal rim by a narrow groove. 5. Stomatal complexes anomocytic (developmen- Epidermal Cells. Epidermal cell flanges clearly visi- tally tangeticytic), Stomatal pore elliptical. ble using TLM, normal cells isodiametric (cells over CUT-Z-ACJ veins not distinguished by shape), approximately 5. Stomatal complexes cyclocytic, developmen- the same size as the stomata, anticlinal walls tally stephanocytic Stomatal pore with a key- curved to wavy, or markedly sinuous, unbut- hole outline. CUT-Z-ABH tressed, smoothly textured, unornamented. 6. Cuticle over the stoma so thin it often breaks Indumentum. Glabrous. away, or, under TLM view, gives impression of the outer stomatal ledge is “floating”. CUT- CUT-Z-FGA Z-EEJ Figure 42.7-42.8 6. Cuticle over the stoma typically robust, not Reference Specimen and locality: SL2016, Sthd- breaking. 7. 010. 7. Stomatal outline unclear under TLM, no T-thick- Stomatal complexes. Stomatal complexes isolated, enings at poles. CUT-Z-FGB possibly with some general alignment although this 7. Stomatal outline clear under TLM, often T-thick- is not clear, number of subsidiary cells unclear, size enings at poles. CUT-Z-EDI range unimodal. Periclinal walls of same thickness as normal epidermal cells, smooth, ornamented

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Figure 43. 1-8. Cuticle with persistent trichomes: CUT-Z-FGB and CUT-Z-EDI. 1. CUT-Z-FGB TLM view showing sto- matal complexes and a trichome (SL1826, scale-bar = 50 µm); 2. CUT-Z-FGB TLM detail of single stomatal complex (SL1826, scale-bar = 20 µm); 3. CUT-Z-FGB TLM view of a trichome (SL1826, scale-bar = 50 µm); 4. CUT-Z-FGB TLM view of a trichome (SL1826, scale-bar = 50 µm); 5. CUT-Z-EDI. TLM view showing stomatal complexes (lower right) the remaining base of a trichome (SL2002, scale-bar = 50 µm); 6. CUT-Z-EDI. TLM detail of single stomatal complex (SL2002, scale-bar = 20 µm); 7. CUT-Z-EDI. SEM view of outer cuticular surface showing a single stomatal complex with prominent outer stomatal ledges. The light, granular material may be fungal growth after the specimen was mounted (S-1078, scale-bar = 10 µm); 8. CUT-Z-EDI. SEM view of inner cuticular surface showing where two stomata have detached (S-1078, scale-bar = 50 µm).

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Figure 44. 1-8. Cuticle with persistent trichomes: CUT-Z-CDC, CUT-Z-FFE, and CUT-Z-EEJ. 1. CUT-Z-CDC, TLM view showing stomatal complexes and a trichome (SL0332, scale-bar = 50 µm); 2. CUT-Z-CDC, TLM detail of three stomatal complexes (SL0332, scale-bar = 20 µm); 3. CUT-Z-CDC, SEM view of outer cuticular surface showing a sto- matal complex (left) partially obscured by ridges of cutin, and a broken trichome (right) (S-1104, scale-bar = 20 µm); 4. CUT-Z-CDC, SEM view of inner cuticular surface showing a single stomatal complex. Note very granular texture (S-1104, scale-bar = 10 µm); 5. CUT-Z-FFE, TLM view showing stomatal complexes and trichomes (SL2120, scale- bar = 50 µm); 6. CUT-Z-FFE, TLM detail of two stomatal complexes (SL2120, scale-bar = 20 µm); 7. CUT-Z-EEJ, TLM view showing stomatal complexes and broken trichomes (SL1778, scale-bar = 50 µm); 8. CUT-Z-EEJ, TLM detail of single stomatal complex (SL1778, scale-bar = 20 µm).

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CUT-Z-FGB ing over whole stoma, with a broad, sub-circular Figure 43.1-43.4 pore. Reference Specimen and locality: SL1826, Sthd- Epidermal Cells. Epidermal cell flanges clearly visi- 095. ble using TLM, normal cells isodiametric (cells over Stomatal complexes. Stomatal complexes evenly major veins more elongate), approximately the spread, isolated, randomly oriented, anisocytic, same size as the stomata, anticlinal walls curved to size range unimodal. Subsidiary cells (3) with peri- wavy, buttressed, inner and outer surfaces smooth, clinal walls thinner than that over normal epidermal unornamented. cells, smooth, ornamented with ridges concentric Indumentum. Bases of trichomes common, proba- about the stoma. Guard cell pair outline difficult or bly restricted to regions over veins (unclear), per- impossible to see in TLM view, not outlined by a sistent, but all cases the main length of the clear anticlinal wall, at same level as subsidiary trichome has broken off irregularly just above the cells (exposed on surface), length 20–22 m base, inserted between unmodified epidermal (medium), little polar development between guard cells, basal diameter much larger than a normal cells (guard cells appear as continuous ring). Outer epidermal cell. stomatal ledge elliptical, thinner than normal epi- dermal cells, only extending over inner edge of CUT-Z-CDC stoma, with a narrowly elliptic pore. Figure 44.1-44.4 Epidermal Cells. Epidermal cell flanges clearly visi- Reference Specimen and locality: SL0332, Mata- ble using TLM, normal cells highly variable from 23. isodiametric to elongate (cells over veins not distin- Stomatal complexes. Stomatal complexes evenly guished by shape), approximately the same size spread, isolated, randomly oriented, number of as the stomata, anticlinal walls curved to wavy, subsidiary cells unclear (difficult to count under unbuttressed, smoothly textured, unornamented. TLM), size range bimodal, with distinct 'giant sto- Indumentum. Hirsute, trichomes common, scat- matal complexes' present. Subsidiary cell periclinal tered over venal and non-venal regions, uniseriate, walls thicker than over normal epidermal cells, a chain of 4-5 individual cells, inserted between smooth, ornamented with many ridges which tend radially elongate epidermal cells. Insertion diame- to flow around the stoma but also frequently cross ter much smaller than a normal epidermal cell, but portions of the stoma. Guard cell pair outline ovate, then the trichome base expanded abruptly above outer margin obscured under TLM by surface orna- the insertion point. mentation, length 11–12 m (small), at same level Distinguishing features. Clearly distinguished by as subsidiary cells (exposed on surface). Outer the uniseriate trichomes which are inserted over stomatal ledge sub circular, thinner than normal several modified epidermal cells. epidermal cells, extending over whole stoma, with a broad, sub-circular pore. CUT-Z-EDI Epidermal Cells. Epidermal cell flanges poorly visi- Figure 43.5-43.8 ble under TLM and only visible away from stomatal Reference Specimen and locality: SL2002, Sthd- complexes, normal cells highly variable from isodi- 072. ametric to elongate (cells over major veins more elongate), anticlinal walls curved to wavy, unbut- Stomatal complexes. Stomatal complexes evenly tressed; inner and outer surfaces smooth, orna- spread, isolated, randomly oriented, anisocytic, or mented with bands of irregular ridges parallel to unevenly paracytic, size range unimodal. Subsid- stomatal complexes and radiating from trichomes. iary cells (2–4) irregularly-shaped, periclinal walls of same thickness as normal epidermal cells, Distinguishing features. Distinguished from other smooth, unornamented. Guard cell pair outline cir- taxa with persistent trichomes in that the surface is cular, not outlined by a clear anticlinal wall, at same densely ornamented with irregular ridges concen- level as subsidiary cells (exposed on surface), tric or roughly parallel to stomatal complexes and length 22–29 m (medium), little wall development which radiate from trichomes.Indumentum. Tri- between guard cells but often with prominent T- chomes sparse; scattered over venal and non- thickenings at poles. Outer stomatal ledge sub cir- venal regions, simple, mostly broken of irregularly cular, thinner than normal epidermal cells, extend- above the base, trichome diameter similar in size to a normal epidermal cell. Epidermal cells around

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Figure 45. 1-6. Cuticle with persistent trichomes: CUT-Z-ABG and CUT-Z-ACJ. 1. CUT-Z-ABG, TLM view showing stomatal complexes and multi-cellular trichomes (SB1353, scale-bar = 50 µm); 2. CUT-Z-ABG, TLM detail of single stomatal complex (SB1353, scale-bar = 20 µm); 3. CUT-Z-ABG, SEM view of inner cuticular surface showing two sto- matal complexes (S-1123, scale-bar = 20 µm); 4. CUT-Z-ABG, SEM view of outer cuticular surface showing stomatal complexes and (upper right) a trichome (S-1123, scale-bar = 20 µm); 5. CUT-Z-ACJ, TLM view showing stomatal complexes and trichomes (SB1350, scale-bar = 50 µm); 6. CUT-Z-ACJ, TLM detail of two stomatal complexes (SB1350, scale-bar = 20 µm).

trichome base (6–7) modified by tangential divi- size range unimodal. Periclinal walls of same thick- sions to form 4–5 rings of foot cells. ness as normal epidermal cells, smooth; orna- mented with bands of ridges projecting at right CUT-Z-FFE angles to the stomatal axis. Guard cell pair outline Figure 44.5-44.6 difficult or impossible to see in TLM view because Reference Specimen and locality: SL2120, GL-07. of ornamentation; not outlined by a clear anticlinal wall; at same level as subsidiary cells (exposed on Stomatal complexes. Stomatal complexes evenly surface), length 17–18 m (medium). Outer stomatal spread, isolated, randomly oriented, number of ledge elliptical, same thickness as normal epider- subsidiary cells unclear, possibly brachyparacytic,

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Figure 46. 1-4. Cuticle with persistent trichomes: CUT-Z-ABH. 1. TLM view showing stomatal complexes and tri- chomes (SB1372, scale-bar = 50 µm); 2. TLM detail of single stomatal complex. Note distinctive 'keyhole' shape to stomatal pore (SB1372, scale-bar = 20 µm); 3. SEM view of outer cuticular surface showing stomatal complexes and trichomes (S-1091, scale-bar = 100 µm); 4. SEM view of inner cuticular surface showing a single stomatal complex (S-1091, scale-bar = 20 µm).

mal cells, extending over whole stoma, with a nar- Stomatal complexes. Stomatal complexes evenly rowly elliptic pore. spread, isolated, randomly oriented, anisocytic, Epidermal Cells. Epidermal cell flanges clearly visi- size range unimodal. Subsidiary cells (3–4) irregu- ble using TLM; normal cells highly variable from larly-shaped, (hard to count as radial flanges of isodiametric to elongate (cells over veins not distin- subsidiary cells are not well developed), periclinal guished by shape); approximately the same size walls of same thickness as normal epidermal cells, as the stomata; anticlinal walls markedly sinuous; smooth, unornamented. Guard cell pair outline buttressed; smoothly textured, ornamented with ovate, not outlined by a clear anticlinal wall, at ridges of irregular course. same level as subsidiary cells (exposed on sur- face), length 18–23 m (medium), little polar devel- Indumentum. Trichomes common; uniseriate opment between guard cells (guard cells appear as although some may branch from the base, or are continuous ring). Outer stomatal ledge elliptical, perhaps in clusters, trichome diameter similar in much thinner than normal epidermal cells (often size to a normal epidermal cell. broken away), extending over whole stoma. Pore narrowly elliptic. CUT-Z-EEJ Figure 44.7-44.8 Epidermal Cells. Epidermal cell flanges clearly visi- ble using TLM, normal cells isodiametric (cells over Reference Specimen and locality: SL1778, Sthd- veins not distinguished by shape), approximately 030. the same size as the stomata, anticlinal walls Referred specimens and occurrence: SL1731, straight, unbuttressed, smoothly textured, unorna- Sthd-017. mented.

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Indumentum. Bases of trichomes common, appar- Stomatal complexes. Stomatal complexes evenly ently uniseriate although most broken off in the spread, isolated, randomly oriented, anomocytic lower segment, inserted between (6-8) epidermal (developmentally tangeticytic), size range unimo- cells modified into radially elongate foot cells, but dal. Subsidiary cells (4–5) irregularly-shaped, peri- with no distinct thickening of walls, basal diameter clinal walls of same thickness as normal epidermal similar to much larger than a normal epidermal cell. cells, smooth, unornamented. Outlined by a well- Distinguishing features. Distinguished from other defined anticlinal wall, at same level as subsidiary taxa with persistent trichomes in the very thin cuti- cells (exposed on surface), length 13–15 m (small). cle over the stoma which often breaks away, or Outer stomatal ledge elliptical, same thickness as gives impression of the outer stomatal ledge “float- normal epidermal cells, extending over inner edge ing” in a TLM view. of stoma, with an elliptical pore. Epidermal Cells. Epidermal cell flanges clearly visi- CUT-Z-ABG ble using TLM, normal cells highly variable from Figure 45.1-45.4 isodiametric to elongate (cells over at least some Reference Specimen and locality: SB1353, BL-32. veins are more elongate), approximately the same size as the stomata, anticlinal walls markedly sinu- Referred specimens and occurrence: SL3131, GL- ous, unbuttressed, smoothly textured, unorna- 02. mented. Stomatal complexes. Stomatal complexes evenly Indumentum. Hirsute, trichomes common, proba- spread, isolated, randomly oriented, anisocytic, bly restricted to regions over veins (unclear), unise- size range unimodal. Subsidiary cells (3–4, difficult riate, 3–5 cells long, the cells nearest the trichome to count under TLM) irregularly-shaped, periclinal base are sometimes about as long as they are walls of same thickness as normal epidermal cells, wide, more distal cells are 5–6 times as long as smooth, ornamented with ridges concentric about wide, total length about 88–113 m, the base is a the stoma. Guard cell pair outline circular, outlined circular, non-poral cell, but the base may expand by a well-defined anticlinal wall, at same level as over surrounding epidermal cells, basal diameter of subsidiary cells (exposed on surface), length 12– similar or distinctly larger size than normal epider- 16 m (small). Outer stomatal ledge sub circular, mal cells. Epidermal cells around trichome base thinner than normal epidermal cells, extending over unmodified whole stoma, with a broad, sub-circular pore. Epidermal Cells. Epidermal cell flanges clearly visi- CUT-Z-ABH ble using TLM, normal cells highly variable from Figure 46.1-46.4 isodiametric to elongate (cells over veins not distin- Reference Specimen and locality: SB1372, BL-32. guished by shape), approximately the same size as the stomata, anticlinal walls markedly sinuous, Referred specimens and occurrence: SL2101, GL- unbuttressed, inner surface very finely granular, 07; SL2160, GL-09. outer surface smooth, ornamented with ridges of Stomatal complexes. Stomatal complexes evenly irregular course. spread, isolated, randomly oriented, cyclocytic, Indumentum. Trichomes abundant, scattered over developmentally stephanocytic bicyclic, size range venal and non-venal regions, multicellular, unimodal (but with wide range of stomatal size). unbranched, lower part of trichome formed of sev- Subsidiary cells (5–7) typically elongate tangential eral elongate cells packed side by side, which then to stoma, with periclinal walls thinner than that over narrows to a single apical cell, base diameter normal epidermal cells, smooth, unornamented. equivalent to 3–6 epidermal cells. Guard cell pair outline circular, outer margin obscured under TLM by robustness of outer sto- Distinguishing features. Distinguished from other matal ledge, at same level as subsidiary cells (sto- taxa with persistent trichomes in that the trichomes matal complexes sunken so that outer stomatal are multicellular. ledges at same level as surface of epidermal cells), length 20–28 m (medium). Outer stomatal ledge CUT-Z-ACJ sub circular, same thickness as normal epidermal Figure 45.5-45.6 cells, extending over whole stoma. Pore broad, but Reference Specimen and locality: SB1350, BL-32. narrows abruptly towards the poles (a key-hole outline) and may be slightly off-set from one guard cell to the other.

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Figure 47. 1-8. Cuticle with paracytic stomatal complexes: CUT-Z-CCD and CUT-Z-CFF. 1. CUT-Z-CCD, TLM view showing stomatal complexes (SL0297, scale-bar = 50 µm); 2. CUT-Z-CCD, TLM detail of single stomatal complex (SL0297, scale-bar = 20 µm); 3. CUT-Z-CCD, SEM view of inner cuticular surface showing two stomatal complexes. Note granular texture of subsidiary cells (S-1583, scale-bar = 10 µm); 4. CUT-Z-CCD, SEM view of outer cuticular surface showing two stomatal complexes and irregularly raised epidermal cells (S-1583, scale-bar = 10 µm); 5. CUT- Z-CFF, TLM view showing stomatal complexes (SL0359, scale-bar = 50 µm); 6. CUT-Z-CFF, TLM detail of single sto- matal complex (SL0359, scale-bar = 20 µm); 7. CUT-Z-CFF, SEM view of outer cuticular surface showing a single stomatal complex and prominently raised epidermal cells (S-1232, scale-bar = 20 µm); 8. CUT-Z-CFF, SEM view of inner cuticular surface showing a single stomatal complex (S-1232, scale-bar = 20 µm).

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Epidermal Cells. Epidermal cell flanges clearly visi- Epidermal Cells. Epidermal cell flanges clearly visi- ble using TLM, normal cells isodiametric (cells over ble using TLM, normal cells elongate (cells over major veins more elongate), distinctly larger than major veins more elongate), distinctly larger than the stomata, anticlinal walls curved to wavy, unbut- the stomata, anticlinal walls markedly sinuous, tressed, inner surfaces slightly rough, outer sur- unbuttressed, inner surfaces granular, outer sur- faces smooth, unornamented. faces smooth, unornamented. Indumentum. Trichomes common, scattered over Indumentum. Attachment scars of deciduous tri- venal and non-venal regions, persistent, inserted chomes rare, inserted between (c. 6) epidermal between (6–12) epidermal cells modified into radi- cells modified by tangential divisions to form a sub- ally elongate foot cells, with no distinct thickening circular zone of foot cells, scar diameter much of walls, diameter of insertion ‘peg’ much smaller smaller than a normal epidermal cell. than a normal epidermal cell, but then the trichome Distinguishing features. Distinguished by the large expands abruptly to about the same size or larger and unevenly sized brachyparacytic subsidiary than a typical epidermal cell. cells. Distinguishing features. CUT-Z-ABH is distin- guished from CUT-Z-AJD in the presence of tri- CUT-Z-CFF chomes. It is distinguished from other taxa with Figure 47.5-47.8 persistent trichomes in that the subsidiary cell peri- Reference Specimen and locality: SL0359, BL-05. clinal walls are much thinner than either normal epidermal cells or the outer stomatal ledges. Referred specimens and occurrence: SL3181, BL- 01; SL2541, GL-02. Group 6. Stomatal complexes paracytic Stomatal complexes. Stomatal complexes evenly Key to cuticle with paracytic stomatal spread, isolated, randomly oriented, brachypara- complexes cytic, or anisocytic, size range unimodal. Subsid- iary cells (2–4) sometimes forming distinct polar 1. Stomatal complexes large, prominent T-piece cells (right angles to stomatal orientation) and 2 lat- thickening at guard cell poles, epidermal cell anti- eral cells (parallel to stomatal orientation), pericli- clinal walls sinuous. CUT-Z-CDD nal walls of same thickness as normal epidermal 1. Stomatal complexes medium, slight T-piece cells, smooth, unornamented. Guard cell pair out- thickening at guard cell poles, epidermal cell anti- line ovate, outlined by a well-defined anticlinal wall, clinal walls wavy to slightly sinuous. CUT-Z-CFF length 22–30 m (medium), at same level as subsid- iary cells (exposed on surface), with slight T-piece CUT-Z-CCD thickenings at polar ends. Outer stomatal ledge Figure 47.1-47.4 sub circular, thinner than normal epidermal cells, Reference Specimen and locality: SL0297, BL-01. extending over whole stoma, with a broad, sub-cir- cular pore. Referred specimens and occurrence: SL0234, BL- 04; SL3075, BL-27; SL3152, GL-01; SL4791, GL- Epidermal Cells. Epidermal cell flanges clearly visi- 07; SL2128, GL-08; SL2870, GL-20; SL2920, GL- ble using TLM, normal cells highly variable from 22; SL2947, GL-25; SL1235, BL-32. isodiametric to elongate (cells over veins not distin- guished by shape), distinctly larger than the sto- Stomatal complexes. Stomatal complexes evenly mata, anticlinal walls wavy to slightly sinuous, spread, isolated, randomly oriented, brachypara- unbuttressed, outer surfaces smooth, but with peri- cytic, size range unimodal. Subsidiary cells more clinal walls irregularly raised leaving grooves over or less equal to unevenly sized, typically expanding the anticlinal walls. to longer than the guard cells, periclinal walls of same thickness as normal epidermal cells, smooth, Indumentum. Glabrous. unornamented. Guard cell pair outline elongate, Distinguishing features. Although the typical sub- often with flattened poles, outlined by a well- sidiary cell pattern of CUT-Z-CFF is brachypara- defined anticlinal wall, length 38–40 m (large), at cytic, the underlying stomatal pattern is probably same level as subsidiary cells (exposed on sur- anisocytic. A spiralling process of cell division face), with very prominent T-piece thickenings at results in two subsidiary cells of reasonably equal polar ends. Outer stomatal ledge sub circular, size and on opposite sides of the guard cells. A thicker than normal epidermal cells, extending over third subsidiary can sometimes be distinguished. centre of stoma, with a broad, sub-circular pore.

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Figure 48. 1-8. Cuticle with anisocytic stomatal complexes: CUT-Z-AJH, CUT-Z-FCD, and CUT-Z-JII. 1. CUT-Z-AJH, TLM view showing stomatal complexes (SB1300, scale-bar = 50 µm); 2. CUT-Z-AJH, TLM detail of single stomatal complex (SB1300, scale-bar = 20 µm); 3. CUT-Z-FCD, TLM view showing stomatal complexes (SL3016, scale-bar = 50 µm); 4. CUT-Z-FCD, TLM detail of single stomatal complex (SL3016, scale-bar = 20 µm); 5. CUT-Z-JII, TLM view showing stomatal complexes (SB1356, scale-bar = 50 µm); 6. CUT-Z-JII, TLM detail of single stomatal complex (SB1356, scale-bar = 20 µm). Note the prominent polar rods; 7. CUT-Z-JII, SEM view of inner cuticular surface showing a single stomatal complex. Note lumpier texture of subsidiary cells than epidermal cells (S-1092, scale-bar = 10 µm); 8. CUT-Z-JII, SEM view of outer cuticular surface showing a single stomatal complex with low outer sto- matal ledges and flanked by short, low ridges (S-1092, scale-bar = 10 µm).

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This taxon thus differs fundamentally from truly thickness as normal epidermal cells, extending brachyparacytic taxa. over whole stoma, with a broad, sub-circular pore. Group 7. Stomatal complexes anisocytic Epidermal Cells. Epidermal cell flanges clearly visi- ble using TLM; normal cells elongate (cells over Key to cuticle with anisocytic stomatal major veins more elongate); distinctly larger than complexes the stomata; anticlinal walls curved to wavy; unbut- 1. Trichomes present and persistent. CUT-Z-AJH tressed; smoothly textured, unornamented. 1. Cuticle glabrous. 2. Indumentum. Hirsute; trichome bases common; scattered over venal and non-venal regions but 2. Epidermal cell periclinal walls distinctly raised, slightly more common over veins, persistent, but all leaving very narrow, deep grooves over anticlinal cases broken off near base, inserted between walls. CUT-Z-ECE unmodified epidermal cells, basal diameter similar 2. Epidermal cell periclinal walls not distinctly in size to a normal epidermal cell. raised. 3. Distinguishing features. Distinguished from CUT-Z- 3. Epidermal cell anticlinal walls sinuous, promi- JII in the presence of persistent trichomes. nent polar rod thickenings. CUT-Z-JII 3. Epidermal cell anticlinal walls straight to wavy, CUT-Z-FCD without prominent polar rod thickenings. 4. Figure 48.3-48.4 4. Subsidiary cells granular, stomatal complex with Reference Specimen and locality: SL3016, GL-28. distinctive angular outline. CUT-Z-FFB Stomatal complexes. Stomatal complexes in are- 4. Subsidiary cells not granular, stomatal complex oles, isolated, randomly oriented, anisocytic, (this without a distinctive angular outline. 5. appears to be the basic state, but there are often more than the definitive three subsidiary cells), size 5. Peristomatal rim present. CUT-Z-FJG range unimodal. Subsidiary cells (3–4) irregularly- 5. Peristomatal rim absent. 6. shaped, periclinal walls of same thickness as nor- 6. Normal epidermal cells elongate, outer stomatal mal epidermal cells, smooth, unornamented. ledge distinctly thinner than normal epidermal cells, Guard cell pair outline ovate, outlined by a well- subsidiary cells (3–4). CUT-Z-FCD defined anticlinal wall, at same level as subsidiary cells (exposed on surface), length 22–30 m 6. Normal epidermal cells isodiametric to elongate, (medium), with prominent T-piece thickenings at outer stomatal ledge slightly thinner than normal polar ends. Outer stomatal ledge sub circular, dis- epidermal cells, subsidiary cells (4–5). CUT-Z-EIH tinctly thinner than normal epidermal cells, extend- ing over centre of stoma, with a broad, sub-circular CUT-Z-AJH pore. Figure 48.1-48.2 Epidermal Cells. Epidermal cell flanges clearly visi- Reference Specimen and locality: SB1300, Mata- ble using TLM, normal cells elongate (cells over 03. major veins more elongate), approximately the Referred specimens and occurrence: SL0087, BL- same size as the stomata, anticlinal walls curved to 08; SL1196, GL-02. wavy, unbuttressed, smoothly textured, unorna- Stomatal complexes. Stomatal complexes evenly mented. spread, isolated, randomly oriented, anisocytic, Indumentum. Glabrous. heliocyctic, size range bimodal, with distinct 'giant stomatal complexes' present. Subsidiary cells (3) CUT-Z-JII of varying size, curved and generally tangential to Figure 48.5-48.8 the guard cells, periclinal walls of same thickness Reference Specimen and locality: SB1356, BL-32. as normal epidermal cells; smooth; unornamented. Guard cell pair outline circular; outlined by a well- Stomatal complexes. Stomatal complexes in are- defined anticlinal wall; length 13–15 m (small); at oles, sometimes networked, randomly oriented, same level as subsidiary cells (exposed on sur- anisocytic, size range unimodal. Subsidiary cells face); with prominent T-piece thickenings at polar (3–4) with periclinal walls of same thickness as ends. Outer stomatal ledge sub circular, same normal epidermal cells, inner surfaces granular, unornamented. Guard cell pair outline circular, out-

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Figure 49. 1-6. Cuticle with anisocytic stomatal complexes: CUT-Z-FFB and CUT-Z-FJG. 1. CUT-Z-FFB, TLM view showing stomatal complexes (SL3135, scale-bar = 50 µm); 2. CUT-Z-FFB, TLM detail of single stomatal complex (SL3135, scale-bar = 20 µm); 3. CUT-Z-FFB, SEM view of inner cuticular surface showing two stomatal complexes (S-1397, scale-bar = 10 µm); 4. CUT-Z-FFB, SEM view of outer cuticular surface showing a single stomatal complex (S-1397, scale-bar = 10 µm); 5. CUT-Z-FJG, TLM view showing stomatal complexes (SL2506, scale-bar = 50 µm); 6. CUT-Z-FJG, TLM detail of two stomatal complexes (SL2506, scale-bar = 20 µm). lined by a well-defined anticlinal wall, at same level sinuous, slightly buttressed, inner and outer sur- as subsidiary cells (exposed on surface), length faces smooth, unornamented. 16–20 m (medium), with very prominent polar rods. Indumentum. Glabrous. Outer stomatal ledge sub circular, same thickness as normal epidermal cells, extending over whole Distinguishing features. Distinguished from CUT-Z- stoma, with a broad, sub-circular pore. AJH in being glabrous and having buttressed epi- dermal cell walls. Epidermal Cells. Epidermal cell flanges clearly visi- ble using TLM, normal cells elongate (cells over CUT-Z-FFB major veins more elongate), approximately the Figure 49.1-49.4 same size as the stomata, anticlinal walls markedly Reference Specimen and locality: SL3135, GL-02.

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Figure 50. 1-6. Cuticle with anisocytic stomatal complexes: CUT-Z-ECE and CUT-Z-EIH. 1. CUT-Z-ECE, TLM view showing stomatal complexes (SL1840, scale-bar = 50 µm); 2. CUT-Z-ECE, TLM detail of single stomatal complex (SL1840, scale-bar = 20 µm); 3. CUT-Z-ECE, SEM view of inner cuticular surface showing stomatal complexes. Note massively thickened epidermal cell walls (S-1083, scale-bar = 20 µm); 4. CUT-Z-ECE, SEM view of outer cuticular surface showing stomatal complexes surrounded by epidermal cells with prominently raised outer (pericli- nal) walls (S-1083, scale-bar = 20 µm); 5. CUT-Z-EIH, TLM view showing stomatal complexes (SL1566, scale-bar = 50 µm); 6. CUT-Z-EIH, TLM detail of single stomatal complex (SL1566, scale-bar = 20 µm).

Stomatal complexes. Stomatal complexes in are- same thickness as normal epidermal cells, with a oles, isolated, randomly oriented, size range uni- narrowly elliptic pore. modal. Subsidiary cells (3–4) with periclinal walls Epidermal Cells. Epidermal cell flanges clearly visi- of same thickness as normal epidermal cells, gran- ble using TLM, normal cells isodiametric (cells over ular, unornamented, with angular outline. Guard both major and fine venation more elongate), cell pair outline elongate, outlined by a well-defined approximately the same size as the stomata, anti- anticlinal wall, at same level as subsidiary cells clinal walls straight, unbuttressed, inner surfaces (though may be slightly sunken below surface), smooth, outer surfaces rough, unornamented. length 22–30 m (medium). Outer stomatal ledge Indumentum. Glabrous.

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Identification. The distinctive angular outline to the outline circular, outlined by a well-defined anticlinal stomatal complex is reminiscent of the Oxylobium wall, at same level as subsidiary cells (exposed on illicifolium illustrated by Jordan (1997). This may surface), length 25–35 m (medium). Outer stomatal indicate relationships with the Fabaceae. ledge sub circular, same thickness as normal epi- dermal cells, extending over whole stoma, with a CUT-Z-FJG broad, sub-circular pore. Figure 49.5-49.6 Epidermal Cells. Epidermal cell flanges not clearly Reference Specimen and locality: SL2506, BL-03. visible under TLM because of surface thickenings, Referred specimens and occurrence: SL2238, GL- normal cells very irregular or elongate (cells over 12. veins not distinguished by shape), approximately the same size as the stomata, anticlinal walls Stomatal complexes. Stomatal complexes in are- curved to wavy, very thick, unbuttressed, smoothly oles, isolated, randomly oriented, anisocytic, helio- textured, periclinal walls raised, leaving very nar- cyctic, size range unimodal. Subsidiary cells (3–5) row, deep grooves over anticlinal walls. with periclinal walls of same thickness as normal epidermal cells, smooth, ornamented with a thick Indumentum. Glabrous. peristomatal rim. Guard cell pair outline elongate, Distinguishing features. Distinguished from other outer margin obscured under TLM by surface taxa with anisocytic stomatal complexes in having topography, length 21–30 m (medium). Outer sto- anticlinal walls of epidermal and subsidiary cells matal ledge elliptical, same thickness as normal reflected by strong ridges on cuticle surface. The epidermal cells, extending over whole stoma, with subsidiary and epidermal cell anticlinal walls are a narrowly elliptic pore. also markedly depressed below general surface of Epidermal Cells. Epidermal cell flanges clearly visi- cuticle. ble using TLM, normal cells highly variable from isodiametric to elongate (cells over veins not distin- CUT-Z-EIH guished by shape), approximately the same size Figure 50.5-50.6 as the stomata, anticlinal walls straight, unbut- Reference Specimen and locality: SL1566, Sthd- tressed, smoothly textured, unornamented. 004. Indumentum. Glabrous. Stomatal complexes. Stomatal complexes evenly Distinguishing features. Stomatal complex is elon- spread, isolated, randomly oriented, anisocytic, all gate with rounded ends and with a peristomatal some, or none of the subsidiary cells may have ridge. CUT-Z-CDE differs in having many subsid- been modified by a tangential division, size range iary cells and no peristomatal ridge. bimodal, with distinct 'giant stomatal complexes' present (separated from other stomatal complexes Identification. The distinctive angular outline to the by a distinct stomatal-free zone). Subsidiary cells stomatal complex is reminiscent of the Oxylobium (4–5) irregularly-shaped, difficult to count under ilicifolium (Andrews) Domin illustrated by Jordan TLM, periclinal walls of same thickness as normal (1997). This may indicate relationships within the epidermal cells, smooth, unornamented. Guard cell Fabaceae. pair outline circular, not outlined by a clear anticli- nal wall, at same level as subsidiary cells (exposed CUT-Z-ECE on surface), length 25–32 m (medium), some wall Figure 50.1-50.4 development between guard cells, and sometimes Reference Specimen and locality: SL1840, Sthd- development of T-piece thickenings at the poles. 086. Outer stomatal ledge sub circular, slightly thinner Referred specimens and occurrence: SL2686, than normal epidermal cells, extending over whole Sthd-090. stoma, with a broad, sub-circular pore. Stomatal complexes. Stomatal complexes evenly Epidermal Cells. Epidermal cell flanges clearly visi- spread, isolated, randomly oriented, anisocytic, ble using TLM, normal cells isodiametric to elon- heliocyctic, size range unimodal. Subsidiary cells gate (cells over veins not distinguished by shape), (3) of varying size, curved and generally tangential anticlinal walls curved to wavy, unbuttressed, to the guard cells, periclinal walls of same thick- smoothly textured, unornamented. ness as normal epidermal cells, smooth, orna- Indumentum. Glabrous. mented with a single, irregular rim. Guard cell pair

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Figure 51. 1-8. Cuticle with multicellular trichome attachment scars: CUT-Z-ECC and CUT-Z-EID. 1. CUT-Z-ECC, TLM view showing tightly clustered stomatal complexes and a multi-cellular trichome attachment scar (SL1811, scale-bar = 50 µm); 2. CUT-Z-ECC, TLM view (SL1811, scale-bar = 20 µm); 3. CUT-Z-ECC, SEM view of inner cuticular surface showing a single stomatal complex (S-1366, scale-bar = 10 µm); 4. CUT-Z-ECC, SEM view of outer cuticular surface showing a single stomatal complex depressed with a rough, irregular surface (S-1366, scale-bar = 10 µm); 5. CUT-Z- EID, TLM view showing stomatal complexes and a multi-cellular trichome attachment scar with a thickened rim (SL1811, scale-bar = 20 µm); 6. CUT-Z-EID, TLM view showing stomatal complexes and a multi-cellular trichome attachment scar with a thickened rim (SL1811, scale-bar = 20 µm); 7. CUT-Z-EID, TLM view showing stomatal com- plexes and a thickened multi-cellular trichome attachment scar (SL1718, scale-bar = 50 µm); 8. CUT-Z-EID, TLM detail of three stomatal complexes (SL1718, scale-bar = 20 µm).

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Group 8. Multi Cellular trichome bases present Distinguishing features. Distinct in having stomatal complexes with well-defined (by thickness of anti- Key to cuticle with multi-cellular trichome clinal walls) subsidiary cells which occur in tight bases clusters, within which there is some networking, 1. Trichome attachment scar with thickened, raised and by the many and prominent trichome attach- rim, cells forming base of scar not thickened. CUT- ment scars which are situated over several epider- Z-ECC mal cells. 1. Trichome attachment scar without thickened, raised rim, cells below scar distinctly thickened. CUT-Z-EID CUT-Z-EID Figure 51.7-51.8 Reference Specimen and locality: SL1718, Sthd- CUT-Z-ECC 018. Figure 51.1-51.6 Referred specimens and occurrence: SL3176, BL- Reference Specimen and locality: SL1811, Sthd- 33. 100. Stomatal complexes. Stomatal complexes in are- Referred specimens and occurrence: SL1351, BL- oles, isolated, randomly oriented, cyclocytic, size 05; SL1249, BL-32; SB1308, Mata-03; SL1751, range unimodal. Subsidiary cells (6–8) isodiamet- Sthd-016; SL1704, Sthd-019; SL1871, Sthd-078; ric, periclinal walls of same thickness as normal SL1527, Sthd-111. epidermal cells, smooth, unornamented. Guard cell Stomatal complexes. Stomatal distribution over leaf pair outline ovate, not outlined by a clear anticlinal surfaces hypostomatic, stomatal complexes tightly wall, at same level as subsidiary cells (exposed on clustered in areoles, frequently networked, ran- surface), length 13–18 m (medium), or (small), little domly oriented, anisocytic, size range bimodal, polar development between guard cells (guard with distinct 'giant stomatal complexes' present. cells appear as continuous ring). Outer stomatal Subsidiary cells (3) isodiametric, periclinal walls ledge elliptical, thinner than normal epidermal cells, thicker than over normal epidermal cells, smooth, extending over inner edge of stoma, with a nar- ornamented with many ridges which tend to flow rowly elliptic pore. around the stoma. Guard cell pair outline ovate, Epidermal Cells. Epidermal cell flanges clearly visi- outlined by a well-defined anticlinal wall, length 9– ble using TLM, normal cells isodiametric (cells over 16 m (small), at same level as subsidiary cells major veins vaguely or inconsistently more elon- (exposed on surface), little polar development gate), distinctly smaller than the stomata, anticlinal between guard cells (guard cells appear as contin- walls straight, unbuttressed, smoothly textured, uous ring). Outer stomatal ledge sub circular, thin- unornamented. ner than normal epidermal cells, extending over Indumentum. Attachment scars of deciduous tri- whole stoma, with a broad, sub-circular pore. chomes common, probably restricted to regions Epidermal Cells. Epidermal cell flanges clearly visi- over veins (unclear), inserted over several (4–5) ble using TLM, normal cells isodiametric (cells over modified epidermal cells. Scar diameter much major veins vaguely or inconsistently more elon- larger than a normal epidermal cell. Epidermal gate), approximately the same size as the stomata, cells of the trichome attachment scar strongly anticlinal walls straight, unbuttressed, inner and thickened. outer surfaces smooth, ornamented with many fine Distinguishing features. Distinguished from CUT-Z- ridges over subsidiary cells, running parallel to sto- CCH by the presence of multi-cellular trichome matal axis. attachment scars. Indumentum. Insertion scars of deciduous tri- Group 9. Fine surface ridging or striae around chomes abundant, scattered over venal and non- stomatal complexes venal regions but slightly more common over veins, inserted over several (3–7) modified epider- Key to cuticle with fine surface ridging or striae mal cells. Scar diameter similar in size to a normal around stomatal complexes epidermal cell. Epidermal cells are selectively 1.Cuticle glabrous. 2. thickened to form a slightly raised, thickened rim, although the cells within the rim are not thickened. 1. Attachment scars of trichome present. 3.

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Figure 52. 1-8. Cuticle with prominent ridging or striae: CUT-Z-AAF, CUT-Z-AJE, CUT-Z-EIB, and CUT-Z-JIB. 1. CUT-Z-AAF, TLM view showing stomatal complexes. Most complexes are plugged (SB1354, scale-bar = 50 µm); 2. CUT-Z-AAF, TLM detail of single stomatal complex (SB1354, scale-bar = 20 µm); 3. CUT-Z-AJE, TLM view showing a stomatal complex and a trichome attachment scar (SB1289, scale-bar = 50 µm); 4. CUT-Z-AJE, TLM detail of sin- gle stomatal complex (SB1289, scale-bar = 20 µm); 5. CUT-Z-EIB, TLM view showing two stomatal complexes (SL1762, scale-bar = 50 µm); 6. CUT-Z-EIB, TLM detail of single stomatal complex (SL1762, scale-bar = 20 µm); 7. CUT-Z-JIB, TLM view showing two stomatal complexes within a single patch of striae (SB1364, scale-bar = 50 µm); 8. CUT-Z-JIB, TLM detail of single stomatal complex (SB1364, scale-bar = 20 µm).

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Figure 53. 1-4. Cuticle with stomatal complexes in islands: CUT-Z-EEA. 1. TLM view showing a patch of stomatal complexes on the left and a glandular structure to the right (SL1820, scale-bar = 50 µm). Note granular texture over stomatal patch; 2. TLM detail of a single stomatal complex (SL1820, scale-bar = 20 µm); 3. SEM view of inner cuticu- lar surface showing a single stomatal complex (S-1066, scale-bar = 10 µm); 4. SEM view of outer cuticular surface showing a single stomatal complex covered by granular cuticular material (S-1066, scale-bar = 20 µm).

2. Surface ridging very fine, inconsistent, epidermal extending over whole stoma, with a broad, sub-cir- cells granular. CUT-Z-EIB cular pore which typically appears plugged with 2. Surface ridging distinct, over subsidiary cells, dark material. epidermal cells not granular. CUT-Z-JIB Epidermal Cells. Epidermal cell flanges clearly visi- 3. Ridges over subsidiary and normal epidermal ble using TLM, normal cells highly variable from cells, stomatal pores plugged. CUT-Z-AAF isodiametric to elongate (cells over both major and fine venation more elongate), approximately the 3. Ridges restricted to subsidiary cells, stomatal same size as the stomata, anticlinal walls curved to pores not plugged. CUT-Z-AJE wavy, unbuttressed, smoothly textured, orna- mented with 'flowing' pattern of many fine ridges CUT-Z-AAF over subsidiary and normal epidermal cells. Figure 52.1-52.2 Indumentum. Attachment scars of deciduous tri- Reference Specimen and locality: SB1354, BL-32. chomes sparse, scattered over venal and non- Stomatal complexes. Stomatal complexes in are- venal regions but slightly more common over oles, isolated, randomly oriented, cyclocytic, size veins, simple, inserted between (6–8) epidermal range unimodal. Subsidiary cells (4–6) irregularly- cells modified only slightly, with a thickened poral shaped, difficult to count under TLM, periclinal rim, scar diameter similar in size to a normal epi- walls of same thickness as normal epidermal cells, dermal cell. smooth, ornamented with many ridges which tend to flow around the stoma. Guard cell pair outline CUT-Z-AJE ovate, at same level as subsidiary cells, outer mar- Figure 52.3-52.4 gin obscured under TLM by surface ornamentation, Reference Specimen and locality: SB1289, Mata- length c. 20 m (medium). Outer stomatal ledge sub 03. circular, same thickness as normal epidermal cells,

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Stomatal complexes. Stomatal complexes evenly fine ridging on some subsidiary cells, but generally spread, isolated, randomly oriented, number of unornamented. subsidiary cells unclear, size range unimodal. Sub- Indumentum. Glabrous. sidiary cells irregularly-shaped, periclinal walls of same thickness as normal epidermal cells, smooth, CUT-Z-JIB ornamented with ridges concentric about the stoma Figure 52.7-52.8 (outline not visible under TLM). Guard cell pair out- line ovate, at same level as subsidiary cells, outer Reference Specimen and locality: SB1364, BL-32. margin obscured under TLM by surface ornamen- Stomatal complexes. Stomatal complexes in are- tation, length 29–32 m (medium), at same level as oles, isolated, randomly oriented, number of sub- subsidiary cells (exposed on surface). Outer sto- sidiary cells unclear, size range unimodal. matal ledge sub circular, thinner than normal epi- Subsidiary cells with periclinal walls thinner than dermal cells, extending over whole stoma, with a that over normal epidermal cells, smooth, orna- broad, sub-circular pore. mented with many ridges which tend to flow around Epidermal Cells. Epidermal cell flanges clearly visi- the stoma (outline not visible under TLM). Guard ble using TLM, normal cells highly variable from cell pair outline ovate, at same level as subsidiary isodiametric to elongate (cells over veins not distin- cells, outer margin obscured under TLM by surface guished by shape), approximately the same size ornamentation, length 35–40 m (medium), little as the stomata, anticlinal walls curved to wavy, polar development between guard cells (guard unbuttressed, smoothly textured ornamented with cells appear as continuous ring). Outer stomatal irregular concentric ridges around subsidiary cells. ledge sub circular, thinner than normal epidermal cells, extending over whole stoma, with a broad, Indumentum. Attachment scars of deciduous tri- sub-circular pore. chomes common, scattered over venal and non- venal regions, persistent, but all cases broken off Epidermal Cells. Epidermal cell flanges clearly visi- near base, inserted between (4–7) epidermal cells ble using TLM, normal cells isodiametric (cells over modified only slightly, with a thickened rim, scar veins not distinguished by shape), distinctly smaller diameter much smaller than a normal epidermal than the stomata, anticlinal walls straight, unbut- cell. tressed, smoothly textured ornamented with many fine ridges over subsidiary cells, running parallel to CUT-Z-EIB stomatal axis. Figure 52.5-52.6 Indumentum. Glabrous. Reference Specimen and locality: SL1762, Sthd- Group 10. Stomatal complexes in islands 011. Key to stomatal complexes in islands Stomatal complexes. Stomatal complexes evenly spread, isolated, randomly oriented, number of 1. Glandular structures or lenticels common. CUT- subsidiary cells unclear, size range unimodal. Sub- Z-EEA sidiary cells irregularly-shaped, difficult to count 1. Glandular structures or lenticels absent or indis- under TLM, periclinal walls of same thickness as tinct. 2. normal epidermal cells, granular, ornamented with 2. Subsidiary cells often modified by tangential divi- ridges concentric about the stoma. Guard cell pair sions, attachment scars of trichomes outline circular, not outlined by a clear anticlinal present. CUT-Z-EDC wall, at same level as subsidiary cells (exposed on surface), length c. 35 m (medium), little polar 2. Subsidiary cells not modified by tangential divi- development between guard cells (guard cells sions, glabrous. CUT-Z-EIC appear as continuous ring). Outer stomatal ledge sub circular, thinner than normal epidermal cells, CUT-Z-EEA extending over whole stoma, with a broad, sub-cir- Figure 53.1-53.4 cular pore. Reference Specimen and locality: SL1820, Sthd- Epidermal Cells. Epidermal cell flanges poorly visi- 091. ble under TLM and only visible away from stomatal Referred specimens and occurrence: SL1268, BL- complexes, normal cells isodiametric (unclear if 32; SL1560, Sthd-004, SL2008, Sthd-020; SL1657, cells over veins are distinguished); anticlinal walls Sthd-040; SL2004, Sthd-072; SL1616, Sthd-106; straight; unbuttressed; texture granular; some very SL1598, Sthd-108; SL1650, Sthd-110.

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Figure 54. 1-8. Cuticle with stomatal complexes in islands: CUT-Z-EDC and CUT-Z-EIC. 1. CUT-Z-EDC, TLM view showing stomatal complexes. Note the giant stomatal complex at lower left (SL1908, scale-bar = 50 µm); 2. CUT-Z- EDC, TLM view showing stomatal complexes (SL1908, scale-bar = 100 µm); 3. CUT-Z-EDC, TLM view showing sto- matal complexes. The lower complex has a brachyparacytic subsidiary cell arrangement while the others are less ordered and can be termed cyclocytic (SL1908, scale-bar = 20 µm); 4. CUT-Z-EDC, SEM view of inner cuticular sur- face showing stomatal complexes. Note networking and granular texture (S-1048, scale-bar = 20 µm); 5. CUT-Z- EDC, SEM view of outer cuticular surface showing three stomatal complexes with slight raised outer stomatal ledges and an otherwise smooth surface (S-1048, scale-bar = 20 µm); 6. CUT-Z-EDC, SEM view of inner cuticular surface showing a trichome attachment scar (S-1048, scale-bar = 20 µm); 7. CUT-Z-EIC, TLM view showing stomatal com- plexes. Note common networking (SL2438, scale-bar = 50 µm); 8. CUT-Z-EIC, TLM view showing stomatal com- plexes (SL2438, scale-bar = 20 µm).

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Stomatal complexes. Stomatal complexes in are- ledge sub circular, thinner than normal epidermal oles, crowded, randomly oriented, paracytic, or cells, extending over whole stoma, with a broad, anisocytic, stomatal complexes are in groups of sub-circular pore. typically 2–3 which are located within a group of Epidermal Cells. Epidermal cell flanges clearly visi- cells clearly bounded by a series of anticlinal cell ble using TLM; normal cells isodiametric (cells over walls which form a continuous loop. All cells within major veins more elongate), approximately the this group appear to have divided as part of the same size as the stomata, anticlinal walls curved to stomatal-forming process, size range unimodal. wavy, or markedly sinuous, unbuttressed, inner Subsidiary cells irregularly-shaped, periclinal walls and outer surfaces smooth, unornamented. of same thickness as normal epidermal cells, the exterior of all cells in the stomatal patch are cov- Indumentum. Attachment scars of deciduous tri- ered with granular cuticular material. Guard cell chomes sparse, scattered over venal and non- pair outline difficult or impossible to see in TLM venal regions but slightly more common over view, sunken below subsidiary cells, length 18–23 veins, inserted between epidermal cells modified m (medium). by tangential divisions to form a sub-circular zone of foot cells with thickened periclinal walls. Scar Epidermal Cells. Epidermal cell flanges clearly visi- diameter distinctly smaller, about a third to a quar- ble using TLM, normal cells highly variable from ter of typical epidermal cells. isodiametric to elongate (cells over veins distin- guished by being more regularly oriented, in flow- Non-stomatal surface. Epidermal cells isodiamet- ing files. but not elongate), distinctly larger than the ric, polygonal, cells over major veins distinguished, stomata, anticlinal walls straight, unbuttressed, trichomes present and sparse (same morphology inner surfaces very finely granular, outer surfaces as stomatal surface). smooth, unornamented. Glands. Multi-cell glands possibly present. Indumentum. Glabrous. Distinguishing features. Distinguished from CUT-Z- Glands. Glandular structures or lenticels common, FAA in having epidermal walls which are sinuous visible as large (50–75 mu) circular holes sur- away from the stomatal complexes, subsidiary rounded by rings of highly, tangentially divided cells which stain darker than normal epidermal cells. cells, uncommon trichome scars, and, when present, do not have a massively thickened poral CUT-Z-EDC rim. Figure 54.1-54.6 CUT-Z-EIC Reference Specimen and locality: SL1908, Sthd- Figure 54.7-54.8 074. Reference Specimen and locality: SL2438, Sthd- Referred specimens and occurrence: SL2007, 073. Sthd-020. Referred specimens and occurrence: SL3043, BL- Stomatal complexes. Stomatal distribution over leaf 22; SL3063, BL-25; SL2962, GL-25. surfaces hypostomatic, stomatal complexes in are- oles, frequently networked, randomly oriented, Stomatal complexes. Stomatal complexes in are- cyclocytic or brachyparacytic, all some, or none of oles, frequently networked, randomly oriented, act- the subsidiary cells may have been modified by a inocytic, or staurocytic, size range unimodal. tangential division, developmentally stephanocytic Subsidiary cells (4–6) irregularly-shaped, periclinal bicyclic, size range bimodal, with distinct 'giant sto- walls thicker than over normal epidermal cells, matal complexes' present (separated from other smooth, unornamented. Guard cell pair outline cir- stomatal complexes by a distinct stomatal-free cular, outlined by a well-defined anticlinal wall, zone). Subsidiary cells (4–7) irregularly-shaped, length 15–20 m (medium), at same level as subsid- periclinal walls thicker than over normal epidermal iary cells (exposed on surface), with prominent T- cells, inner surface granular, outer surfaces piece thickenings at polar ends. Outer stomatal smooth, unornamented. Guard cell pair outline cir- ledge sub circular, thinner than normal epidermal cular; outlined by a well-defined anticlinal wall; cells, extending over whole stoma, with a broad, length 20–23 m (medium); at same level as subsid- sub-circular pore. iary cells (exposed on surface); with prominent T- Epidermal Cells. Epidermal cell flanges clearly visi- piece thickenings at polar ends. Outer stomatal ble using TLM, normal cells elongate (cells over

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Figure 55. 1-6. Cuticle with unusually shaped outer stomatal ledges: CUT-Z-FBI and CUT-Z-FCB. 1. CUT-Z-FBI, TLM view showing stomatal complexes and trichome insertion scar (SL2956, scale-bar = 50 µm); 2. CUT-Z-FBI, TLM view showing two stomatal complexes (SL2956, scale-bar = 20 µm); 3. CUT-Z-FCB, TLM view showing sto- matal complex (SL3027, scale-bar = 50 µm); 4. CUT-Z-FCB, TLM detail of single stomatal complex (SL3027, scale- bar = 20 µm); 5. CUT-Z-FCB, SEM view of inner cuticular surface showing a single stomatal complex. Note pro- nounced granular texture (S-1386, scale-bar = 10 µm); 6. CUT-Z-FCB, SEM view of outer cuticular surface showing three stomatal complexes. Note the distinctive key-hole shape of the stomatal pore (S-1386, scale-bar = 10 µm). major veins vaguely or inconsistently more elon- Group 11. Outer stomatal ledges prominent or gate), distinctly larger than the stomata, anticlinal unusually shaped walls curved to wavy, or markedly sinuous, unbut- Key to cuticle with prominent or unusually tressed, smoothly textured, unornamented. shaped outer stomatal ledges Indumentum. Glabrous. 1. Stomatal pore elliptical, outer stomatal ledge Distinguishing features. Distinguished from CUT-Z- possibly complex. CUT-Z-FBI CGE in having stomatal complexes which are clearly in tight clusters, subsidiary cells clearly dis- 1. Stomatal pore typical or often “key-hole” shaped, tinguished by thick periclinal walls, and relatively outer stomatal ledge simple. 2. smaller stomatal complexes.

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Figure 56. 1-8. Cuticle with unusual shaped outer stomatal ledges: CUT-Z-AJD and CUT-Z-FFG. 1. CUT-Z-AJD, TLM view showing stomatal complexes (SB1379, scale-bar = 50 µm); 2. CUT-Z-AJD, TLM detail of single stomatal com- plex (SB1379, scale-bar = 20 µm). Note key-hole shape to the stomatal pore; 3. CUT-Z-AJD, SEM view of inner cutic- ular surface showing a single stomatal complex (S-1200, scale-bar = 10 µm); 4. CUT-Z-AJD, SEM view of outer cuticular surface showing a single stomatal complex with prominent outer stomatal ledges (S-1200, scale-bar = 10 µm); 5. CUT-Z-FFG, TLM view showing stomatal complexes (SL2229, scale-bar = 50 µm); 6. CUT-Z-FFG, TLM detail of single stomatal complex (SL2229, scale-bar = 20 µm); 7. CUT-Z-FFG, SEM view of inner cuticular surface showing (near centre) a reasonably well preserved stomatal complex and (left) a poorly preserved one (S-1396, scale-bar = 10 µm); 8. CUT-Z-FFG, SEM view of inner cuticular surface showing a single stomatal complex (S-1396, scale-bar = 10 µm).

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2. Outer stomatal ledge a distinctive 'lemon' shape. Stomatal complexes. Stomatal complexes evenly CUT-Z-FCB spread, isolated, randomly oriented, size range 2. Outer stomatal ledge not 'lemon' shaped. 3. unimodal (but with wide range of stomatal size). Subsidiary cells (4–7) irregularly-shaped, difficult to 3. Outer stomatal ledges thinner than normal epi- count under TLM, periclinal walls of same thick- dermal cells, epidermal cell anticlinal walls ness as normal epidermal cells, inner surfaces straight. CUT-Z-AJD granular, outer surfaces smooth, ornamented with 3. Outer stomatal ledges thicker than normal epi- 3–4 ridges parallel with, and on either side of the dermal cells, epidermal cell anticlinal walls stoma. Guard cell pair outline difficult or impossible sinuous. CUT-Z-FFG to see in TLM view, at same level as subsidiary cells, outer margin obscured under TLM by surface CUT-Z-FBI ornamentation, length 20–38 m (medium). Outer Figure 55.1-55.2 stomatal ledge a distinctive 'lemon' shape, broad in Reference Specimen and locality: SL2956, GL-25. the middle, and narrowing sharply at either end, thicker than normal epidermal cells, extending over Referred specimens and occurrence: SL2851, GL- whole stoma, with a broad pore, but which narrows 18. abruptly towards the poles (a key-hole outline). Stomatal complexes. Stomatal complexes in are- Epidermal Cells. Epidermal cell flanges clearly visi- oles, isolated, randomly oriented, cyclocytic, size ble using TLM, normal cells isodiametric (cells over range unimodal. Subsidiary cells (4–6, difficult to veins not distinguished by shape), approximately count under TLM) with periclinal walls of same the same size as the stomata, anticlinal walls thickness as normal epidermal cells, smooth, orna- curved to wavy, unbuttressed, inner surfaces finely mented with 3–4 ridges parallel with, and on either granular, outer surfaces smooth, unornamented. side of the stoma. Guard cell pair outline difficult or impossible to see in TLM view, outer margin usu- Indumentum. Glabrous. ally obscured by subsidiary cell ornamentation pro- Glands. Glands present, compound, composed of jecting over the outer stomatal ledge, length 22–33 a cluster of cells with periclinal walls thicker than m (medium), clearly separated by polar walls. normal epidermal cells, and surrounded by a Outer stomatal ledge sub circular, thinner than nor- rosette of cells, 2–3 cycles broad, which otherwise mal epidermal cells, extending over whole stoma, look similar to normal epidermal cells. narrowing towards poles, possibly with a complex Distinguishing features. Clearly distinguished by structure (two ledges may overlap), with a broad, the lemon-shaped outer stomatal ledges, and the sub-circular pore. compound glands. Differs from CUT-Z-FBI in hav- Epidermal Cells. Epidermal cell flanges clearly visi- ing straight, unbuttressed epidermal cell walls ble using TLM, normal cells isodiametric (cells over major veins more elongate), approximately the CUT-Z-AJD same size as the stomata; anticlinal walls markedly Figure 56.1-56.4 sinuous; buttressed; smoothly textured, unorna- Reference Specimen and locality: SB1379, BL-32. mented. Referred specimens and occurrence: SL0350, BL- Indumentum. Attachment scars of deciduous tri- 05; SL3041, BL-22; SL3133, GL-02; SL2144, GL- chomes common; scattered over venal and non- 08; SL2182, GL-10; SL2213, GL-12; SL2822, GL- venal regions; inserted between (5–7) epidermal 16; SL2998, GL-27. cells modified into a distinct ring of foot cells, with thickened periclinal walls; scar diameter similar in Stomatal complexes. Stomatal complexes in are- size to a normal epidermal cell. oles, isolated, randomly oriented, cyclocytic, size range unimodal. Subsidiary cells (6) irregularly- Distinguishing features. Has a similar outer sto- shaped, with periclinal walls thinner than that over matal ledge as CUT-Z-FCB but differs in having normal epidermal cells, smooth, unornamented. sinuous and buttressed epidermal cells and tri- Guard cell pair outline circular, at same level as chome attachment scars. subsidiary cells, outer margin obscured under TLM by surface ornamentation, length 17–25 m CUT-Z-FCB (medium). Outer stomatal ledge sub circular, thin- Figure 55.3-55.6 ner than normal epidermal cells, extending over Reference Specimen and locality: SL3027, GL-29.

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Figure 57. 1-8. Cuticle with sinuous epidermal cells and networked stomatal complexes: CUT-Z-ADJ and CUT-Z- EGB. 1. CUT-Z-ADJ, TLM view showing stomatal complexes and trichome attachment scars (SB0860, scale-bar = 50 µm); 2. CUT-Z-ADJ, TLM detail of single stomatal complex (SB860, scale-bar = 20 µm); 3. CUT-Z-ADJ, SEM view of inner cuticular surface showing stomatal complexes. Note networking, slight buttressing and granular texture (S- 1119, scale-bar = 20 µm); 4. CUT-Z-ADJ, SEM view of outer cuticular surface showing a single stomatal complex with prominent outer stomatal ledges, surrounded by discontinuous rings of striae (S-1119, scale-bar = 10 µm); 5. CUT-Z-EGB, TLM view showing stomatal complexes and a trichome attachment scar (SL1955, scale-bar = 50 µm); 6. CUT-Z-EGB, TLM view showing two stomatal complexes (SL1955, scale-bar = 20 µm); 7. CUT-Z-EGB, SEM view of inner cuticular surface showing a single stomatal complex (S-1368, scale-bar = 10 µm); 8. CUT-Z-EGB, SEM view of outer cuticular surface showing a single stomatal complex with slightly raised outer stomatal ledges in a bumpy surface (S-1368, scale-bar = 10 µm).

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Figure 58. 1-4. Stomatal complexes with two rings of narrow subsidiary cells (CUT-Z-FBD) and tangentic stomatal complexes (CUT-Z-FBH). 1. CUT-Z-FBD, TLM view showing stomatal complexes (SL2946, scale-bar = 50 µm); 2. CUT-Z-FBD, TLM detail of single stomatal complex (SL2946, scale-bar = 20 µm). Note the two rings of very narrow subsidiary cells; 3. CUT-Z-FBH, TLM detail of two stomatal complexes (SL2108, scale-bar = 20 µm); 4. CUT-Z-FBH, TLM view showing three stomatal complexes (SL2981, scale-bar = 20 µm). whole stoma, with a broad pore but which narrows Subsidiary cells with periclinal walls of same thick- abruptly towards the poles (a key-hole outline). ness as normal epidermal cells, smooth, orna- Epidermal Cells. Epidermal cell flanges clearly visi- mented with ridges concentric about the stoma. ble using TLM, normal cells isodiametric (cells over Guard cell pair outline difficult or impossible to see both major and fine venation more elongate), dis- in TLM view, not outlined by a clear anticlinal wall, tinctly larger than the stomata, anticlinal walls at same level as subsidiary cells, length 19–23 m straight, unbuttressed, inner surfaces finely granu- (medium). Outer stomatal ledge sub circular, lar, outer surfaces smooth, unornamented. thicker than normal epidermal cells, extending over whole stoma, with a broad pore, but which often Indumentum. Glabrous. narrows abruptly towards the poles (a “key-hole” Distinguishing features. Similar to CUT-Z-ABH, but outline). glabrous. Epidermal Cells. Epidermal cell flanges clearly visi- ble using TLM, normal cells highly variable from CUT-Z-FFG isodiametric to elongate (cells over major veins Figure 56.5-56.8 more elongate), approximately the same size as Reference Specimen and locality: SL2229, GL-12. the stomata, anticlinal walls markedly sinuous, Stomatal complexes. Stomatal complexes in are- slightly buttressed; smoothly textured, unorna- oles, isolated, randomly oriented, number of sub- mented. sidiary cells unclear, size range unimodal. Indumentum. Glabrous.

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Group 12. Epidermal cells sinuous and ness as normal epidermal cells, smooth, unorna- stomatal complexes highly networked mented. Guard cell pair outline circular, outlined by a well-defined anticlinal wall, length 16–24 m Key to cuticle with sinuous epidermal cells and (medium), at same level as subsidiary cells highly networked stomatal complexes (exposed on surface), with prominent T-piece thick- 1. Stomatal complexes clustered, cuticle glabrous, enings at polar ends. Outer stomatal ledge sub cir- often with 2-3 thin ridges parallel to the long cular, same thickness as normal epidermal cells, axis of stoma. CUT-Z-ADJ extending over whole stoma, with a broad, sub-cir- 1. Stomatal complexes not clustered, cuticle with cular pore. trichome attachment scars, unornamented. Epidermal Cells. Epidermal cell flanges clearly visi- CUT-Z-EGB ble using TLM, normal cells elongate (cells over major veins vaguely or inconsistently more elon- CUT-Z-ADJ gate), approximately the same size as the stomata, Figure 57.1-57.4 anticlinal walls markedly sinuous, unbuttressed, Reference Specimen and locality: SB1298, Mata- smoothly textured, unornamented. 03. Indumentum. Attachment scars of deciduous tri- Referred specimens and occurrence: SL1228, BL- chomes common, probably restricted to regions 30. over veins (unclear), inserted between epidermal cells modified into radially elongate foot cells, scar Stomatal complexes. Stomatal complexes in are- diameter similar in size to a normal epidermal cell. oles, often networked, randomly oriented, ano- mocytic (3–4 contact cells), size range unimodal. Group 13. Stomatal complexes typically having Subsidiary cells with periclinal walls of same thick- two rings of narrow subsidiary cells ness as normal epidermal cells, inner surfaces finely granular, outer surfaces slightly rough, unor- CUT-Z-FBD namented, or with 2-3 thin ridges parallel to the Figure 58.1-58.2 long axis of stoma. Guard cell pair outline ovate, Reference Specimen and locality: SL2946, GL-23. outlined by a well-defined anticlinal wall, at same Stomatal complexes. Stomatal complexes in are- level as subsidiary cells (exposed on surface), oles, isolated, randomly oriented, cyclocytic, There length 19–23 m (medium), clearly separated by are typical two rings of markedly narrow subsidiary polar walls. Outer stomatal ledge sub circular, cells, plus the outer ring of broader and more irreg- same thickness as normal epidermal cells, extend- ular cells that the subsidiaries have been cut off ing over whole stoma, with a broad, sub-circular from, developmentally stephanocytic brachypara- pore. cytic, size range unimodal (but with wide range of Epidermal Cells. Epidermal cell flanges clearly visi- stomatal size). Subsidiary cells (4–7) with periclinal ble using TLM, normal cells elongate (cells over walls of same thickness as normal epidermal cells, major veins more elongate), approximately the smooth, unornamented. Guard cell pair outline cir- same size as the stomata, anticlinal walls markedly cular, or ovate, outlined by a well-defined anticlinal sinuous, unbuttressed, inner surfaces finely granu- wall, at same level as subsidiary cells (exposed on lar, outer surfaces slightly rough, unornamented. surface), length 29–30 m (medium), with prominent Indumentum. Glabrous. T-piece thickenings at polar ends. Outer stomatal ledge sub circular, much thicker than normal epi- CUT-Z-EGB dermal cells, extending over whole stoma, with an Figure 57.5-57.8 elliptical pore. Reference Specimen and locality: SL1955, Sthd- Epidermal Cells. Epidermal cell flanges clearly visi- 051. ble using TLM, normal cells isodiametric (cells over both major and fine venation more elongate), Referred specimens and occurrence: SL2289, BL- approximately the same size as the stomata, anti- 18; SB0860, Mata-03. clinal walls straight, unbuttressed, smoothly tex- Stomatal complexes. Stomatal complexes evenly tured, unornamented. spread, frequently networked, randomly oriented, Indumentum. Glabrous. anomocytic, size range unimodal. Subsidiary cells (4–6) isodiametric, periclinal walls of same thick-

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Figure 59. 1-8. Stomatal complexes tangeticytic: CUT-Z-AJI, CUT-Z-FCC, and CUT-Z-EHI. 1. CUT-Z-AJI, TLM view showing stomatal complexes (SL2411, scale-bar = 50 µm); 2. CUT-Z-AJI, TLM detail of two stomatal complexes (SB1293, scale-bar = 20 µm); 3. CUT-Z-FCC, TLM view showing stomatal complexes (SL3010, scale-bar = 50 µm); 4. CUT-Z-FCC, TLM detail of single stomatal complex (SL3010, scale-bar = 20 µm). Note the wedge-shaped sub- sidiary cells; 5. CUT-Z-FCC, TLM view showing stomatal complexes (SL2531, scale-bar = 50 µm); 6. CUT-Z-FCC, TLM detail of single stomatal complex (SL2531, scale-bar = 20 µm); 7. CUT-Z-EHI, TLM view showing stomatal complexes (SL1576, scale-bar = 50 µm). Note the distinctly raised OSLs; 8. CUT-Z-EHI, TLM detail of two stomatal complexes. The tangenticytic divisions are so close to the prominent outer stomatal ledges that they are usually obscured (SL1576, scale-bar = 20 µm).

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Group 14. Stomatal complexes prominently Stomatal complexes. Stomatal complexes evenly cyclocytic (developmentally tangenticytic) with spread, isolated, randomly oriented, cyclocytic, prominent outer stomatal ledges developmentally tangenticytic, size range unimo- dal. Subsidiary cells (4–7, difficult to count under Key to cuticle with prominently cyclocytic TLM) with periclinal walls thinner than that over complexes normal epidermal cells, smooth, unornamented. 1. Trichome attachment scars present. CUT-Z-EHI Guard cell pair outline circular, or ovate, outlined by 1. Cuticle glabrous. 2. a well-defined anticlinal wall, at same level as sub- sidiary cells (exposed on surface), length 22–27 m 2. Guard cell outlines difficult or impossible to see (medium), clearly separated by polar walls. Outer under TLM, outer stomatal ledges distinct stomatal ledge sub circular, same thickness as nor- but appearing to “float” over very thin guard mal epidermal cells, extending over centre of cell cuticle under TLM. CUT-Z-FCC stoma, with a broad, sub-circular pore. 2. Guard cell outlines generally clearly visible, Epidermal Cells. Epidermal cell flanges clearly visi- outer stomatal ledges not “floating”. 3. ble using TLM, normal cells isodiametric (cells over 3. Periclinal walls of subsidiary cells thicker than veins not distinguished by shape), distinctly smaller normal epidermal cells. CUT-Z-FBH than the stomata, anticlinal walls straight, unbut- 3. Periclinal walls of subsidiary cells thinner than tressed, smoothly textured, unornamented. normal epidermal cells. CUT-Z-AJI Indumentum. Glabrous.

CUT-Z-FBH CUT-Z-FCC Figure 58.3-58.4 Figure 59.3-59.6 Reference Specimen and locality: SL2981, GL-26. Reference Specimen and locality: SL3010, GL-28. Stomatal complexes. Stomatal complexes in are- Referred specimens and occurrence: SL2531, BL- oles, isolated, randomly oriented, cyclocytic, devel- 05; SL2087, GL-07. opmentally tangenticytic, size range unimodal. Stomatal complexes. Stomatal complexes evenly Subsidiary cells (4–8) typically elongate, or wedge- spread, isolated, randomly oriented, cyclocytic, or like, often with distinct polar cells (at right angles to staurocytic (there are two rings of subsidiary cells, the stomatal axis) which may 'enclose' the lateral the inner has granular periclinal walls which are cells, periclinal walls thicker than over normal epi- thinner than normal epidermal cells, while the outer dermal cells, smooth, unornamented. Guard cell are smooth, but thicker than normal epidermal pair outline circular, or ovate, outlined by a well- cells), developmentally tangenticytic, size range defined anticlinal wall, at same level as subsidiary unimodal (but with wide range of stomatal size). cells (exposed on surface), length 17–19 m Subsidiary cells (4–5) sometimes forming 2 distinct (medium), with prominent T-piece thickenings at polar cells (at right angles to stomatal orientation) polar ends. Outer stomatal ledge sub circular, and 2 lateral cells (parallel to stomatal orientation), same thickness as normal epidermal cells, extend- smooth, unornamented. Guard cell pair outline dif- ing over whole stoma, with an elliptical pore. ficult or impossible to see in TLM view, not outlined Epidermal Cells. Epidermal cell flanges clearly visi- by a clear anticlinal wall, at same level as subsid- ble using TLM, normal cells highly variable from iary cells, length 25–30 m (medium), with promi- isodiametric to elongate (cells over major veins nent T-piece thickenings at polar ends. Outer more elongate), approximately the same size as stomatal ledge elliptical, thicker than normal epi- the stomata, anticlinal walls curved to wavy, unbut- dermal cells, extending over whole stoma, typically tressed, smoothly textured, unornamented. appearing under TLM to “float” within the much Indumentum. Glabrous. thinner cuticle over the rest of the guard cells, with a narrowly elliptic pore. CUT-Z-AJI Epidermal Cells. Epidermal cell flanges clearly visi- Figure 59.1-59.2 ble using TLM, normal cells highly variable from Reference Specimen and locality: SB1293, Mata- isodiametric to elongate (cells over major veins 03. more elongate), approximately the same size as the stomata, anticlinal walls curved to wavy, unbut- Referred specimens and occurrence: SL2411, BL- tressed, smoothly textured, unornamented. 32.

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Figure 60. 1-6. Stomatal complexes anomocytic: CUT-Z-ADA and CUT-Z-ECF. 1. CUT-Z-ADA, TLM view showing networked stomatal complexes (SB1285, scale-bar = 50 µm); 2. CUT-Z-ADA, TLM detail of single stomatal complex (SB1285, scale-bar = 20 µm). Note the distinctly circular outline of the guard cell pair; 3. CUT-Z-ECF, TLM view showing stomatal complexes (SL0062, scale-bar = 50 µm); 4. CUT-Z-ECF, TLM detail of single stomatal complex (SL0062, scale-bar = 20 µm). Note the lemon-shaped outline of the OSL; 5. CUT-Z-ECF, SEM view of inner cuticular surface showing a single stomatal complex (S-1051, scale-bar = 20 µm); 6. CUT-Z-ECF, SEM view of outer cuticular surface showing a single stomatal complex (S-1051, scale-bar = 20 µm).

Indumentum. Attachment scars of deciduous tri- CUT-Z-EHI chomes common, more common over major veins, Figure 59.7-59.8 inserted between (4–6) epidermal cells modified by Reference Specimen and locality: SL1576, Sthd- tangential divisions to form a sub-circular zone of 004. foot cells having periclinal walls much thicker than normal epidermal cells, diameter much smaller Stomatal complexes. Stomatal distribution over leaf than a normal epidermal cell. surfaces hypostomatic, stomatal complexes evenly spread, isolated, randomly oriented, cyclocytic, all some, or none of the subsidiary cells may have been modified by a tangential division, develop-

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Figure 61. 1-8. Cuticle apparently non-stomatiferous: CUT-Z-CFE, CUT-Z-EHC, CUT-Z-EIF, and CUT-Z-JIC. 1. CUT-Z-CFE, TLM view showing possible stomatal complexes (SL0321, scale-bar = 50 µm); 2. CUT-Z-CFE, TLM detail of possible single stomatal complex (SL0321, scale-bar = 20 µm). Note the slight thickening around the pit; 3. CUT-Z-EHC, TLM view showing possible stomatal complexes (SL1526, scale-bar = 50 µm); 4. CUT-Z-EHC, TLM detail of possible stomatal complexes (SL1526, scale-bar = 20 µm). Note the very smooth, raised rim; 5. CUT-Z-EIF, TLM view showing possible stomatal complexes (SL2432, scale-bar = 50 µm); 6. CUT-Z-EIF, TLM detail of possible single stomatal complex (SL2432, scale-bar = 20 µm). Note the probable bicyclic nature; 7. CUT-Z-JIC, TLM view showing possible stomatal complexes obscured by papillae (SB1395, scale-bar = 50 µm); 8. CUT-Z-JIC, TLM detail of possible stomatal complexes obscured by papillae (SB1395, scale-bar = 20 µm).

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mentally tangenticytic, size range unimodal. Sub- Epidermal Cells. Epidermal cell flanges clearly visi- sidiary cells (5–7) irregularly-shaped, periclinal ble using TLM, normal cells highly variable from walls of same thickness as normal epidermal cells, isodiametric to elongate (cells over both major and smooth, unornamented. Guard cell pair outline cir- fine venation more elongate); approximately the cular, outlined by a well-defined anticlinal wall, same size as the stomata; anticlinal walls straight; length 23–30 m (medium), underthrust by subsid- unbuttressed; smoothly textured, unornamented. iary cells (subsidiary cell anticlinal walls bisect out- Indumentum. Attachment scars of deciduous tri- line of the guard cells), with prominent T-piece chomes sparse; restricted to regions over veins; thickenings at polar ends. Outer stomatal ledge simple; inserted between (4–6) epidermal cells sub circular, same thickness as normal epidermal modified by tangential divisions to form a sub-cir- cells, distinctly raised, extending over whole stoma, cular zone of foot cells, scar diameter much with a broad, sub-circular pore. smaller than a normal epidermal cell. Epidermal Cells. Epidermal cell flanges clearly visi- Distinguishing features. Distinguished from other ble using TLM, normal cells isodiametric (cells over taxa with networked stomatal complexes in having veins not distinguished by shape), approximately anomocytic stomatal complexes and very thin cuti- the same size as the stomata, anticlinal walls cle. straight, unbuttressed, smoothly textured, unorna- mented. CUT-Z-ECF Indumentum. Glabrous. Figure 60.3-60.6 Glands. Glands present, compound, or large circu- Reference Specimen and locality: SL0360, BL-05. lar areas of very thinly cutinised cells (often broken Referred specimens and occurrence: SL0062, away), surrounded by radiating files of normal- Mata-18; SL1614, Sthd-106; SL1597, Sthd-108. thickness epidermal cells. Stomatal complexes. Stomatal complexes evenly Distinguishing features. Distinguished from CUT-Z- spread, isolated, randomly oriented, anisocytic, or CEF and CUT-Z-CFC by having stomatal com- anomocytic, size range unimodal. Subsidiary cells plexes which are raised well above the level of epi- (4–5) irregularly-shaped, periclinal walls of same dermal cells. thickness as normal epidermal cells, smooth, unor- Group 15. Stomatal complexes anomocytic namented. Guard cell pair outline elliptical, not out- lined by a clear anticlinal wall, at same level as Key to anomocytic cuticle subsidiary cells (exposed on surface), length 28– 1. Stomatal complexes networked, attachment 38 m (medium), at same level as subsidiary cells scars of trichomes present. CUT-Z-ADA (exposed on surface), some wall development between guard cells, and sometimes development 1. Stomatal complexes isolated, glabrous. CUT-Z- of T-piece thickenings at the poles. Outer stomatal ECF ledge elliptical, sometimes lemon-shaped, thinner than normal epidermal cells, extending over whole CUT-Z-ADA stoma, with a narrowly elliptic pore. Figure 60.1-60.2 Epidermal Cells. Epidermal cell flanges clearly visi- Reference Specimen and locality: SB1285, BL-33. ble using TLM, normal cells elongate (cells over Stomatal complexes. Stomatal complexes in are- veins not distinguished by shape), approximately oles, frequently networked, randomly oriented, the same size as the stomata, anticlinal walls anomocytic, size range unimodal, (but with wide curved to wavy, unbuttressed, smoothly textured range of stomatal size). Subsidiary cells (5–6) iso- ornamented with discontinuous ridges, which prob- diametric, periclinal walls of same thickness as ably correspond to epidermal cell walls. normal epidermal cells, smooth, unornamented. Indumentum. Glabrous. Guard cell pair outline circular, outlined by a well- defined anticlinal wall, at same level as subsidiary Group 16. Cuticle either non-stomatiferous or cells (exposed on surface), length 20–30 m stomatal complexes very inconspicuous. (medium), little polar development between guard Key to non-stomatiferous cuticle. cells (guard cells appear as continuous ring). Outer stomatal ledge sub circular, same thickness as nor- 1. Ornamented with prominent ridges or peltate mal epidermal cells, extending over whole stoma, scales. CUT-Z-JIC with a broad, sub-circular pore.

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1. Unornamented. 2. Description. Cuticle with scattered highly thick- 2. Possible stomatal complexes surrounded by a ened, elliptical rings of cuticle, possibly stomata or double ring of subsidiary cells. CUT-Z-EIF trichome attachment sites. Epidermal cells isodia- metric on one surface, but more elongate and 2. Possible stomatal complexes surrounded by dis- organised into loose rows on the other. tinct subsidiary cells. 3. 3. Possible stomatal complexes not surrounded by CUT-Z-EIF raised rings of cuticle. CUT-Z-CFE Figure 61.5-61.6 2. Possible stomatal complexes surrounded by Reference Specimen and locality: SL2432, Sthd- raised rings of cuticle. CUT-Z-EHC 051. Description. Possible stomatal complexes sur- CUT-Z-CFE rounded by a double ring of subsidiary cells, with Figure 61.1-61.2 cuticle thinner than that over normal epidermal Reference Specimen and locality: SL0321, Harli- cells, guard cells probably deeply sunken. Epider- wich-3. mal cell anticlinal walls straight, epidermal cell tex- Description. Cuticle with polygonal depressions or ture smooth, unornamented pits which may contain sunken stoma. The edges of the pits show diffuse thickening. Epidermal cells CUT-Z-JIC irregularly shaped, but typically isodiametric, dis- Figure 61.7-61.8 tinctly straight-walled. Reference Specimen and locality: SB1395, BL-33. Description. Stomatal distribution over leaf sur- CUT-Z-EHC faces unknown, size range difficult to observe. Figure 61.3-61.4 Ornamented with prominent ridges or peltate Reference Specimen and locality: SL1526, Sthd- scales. Epidermal cell flanges not clearly visible 111. under TLM because of surface thickenings, normal Referred specimens and occurrence: SL1775, cells unclear, anticlinal walls unclear, smoothly tex- Sthd-022. tured, unornamented.

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