New Zealand Journal of Botany

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Standardizing names applied to pollen and spores in New Zealand Quaternary palynology

N T Moar , J M Wilmshurst & M S McGlone

To cite this article: N T Moar , J M Wilmshurst & M S McGlone (2011) Standardizing names applied to pollen and spores in New Zealand Quaternary palynology, New Zealand Journal of Botany, 49:2, 201-229, DOI: 10.1080/0028825X.2010.526617 To link to this article: https://doi.org/10.1080/0028825X.2010.526617

Published online: 31 May 2011.

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Full Terms & Conditions of access and use can be found at https://www.tandfonline.com/action/journalInformation?journalCode=tnzb20 New Zealand Journal of Botany Vol. 49, No. 2, June 2011, 201Á229

Standardizing names applied to pollen and spores in New Zealand Quaternary palynology NT Moar, JM Wilmshurst* and MS McGlone Landcare Research, Lincoln, New Zealand (Received 27 April 2010; final version received 21 September 2010)

Pollen analysts often study pollen grains that cannot be easily attributed to a particular taxon. As a consequence, a pollen type may be recorded under various names, leading to uncertainty in their interpretation by others. This limits the usefulness of pollen data for inclusion in databases or meta-analyses. To minimize any such ambiguity, we briefly describe the pollen grains of New Zealand gymnosperms and angiosperms and assign an appropriate name to each. Tree ferns are also included, as they are almost ubiquitous in New Zealand palynomorph records, and are often misidentified and misrepresented. We also provide a review of the occurrence of all taxa in New Zealand pollen records, and of pollination modes, pollen production and dispersal. Keywords: pollen grains; Filicopsida; Pinopsida; Magnoliopsida; pollen record; pollination mode; pollen production; pollen dispersal

Introduction may be obscured or destroyed by corrosion Pollen analysis is a valuable tool in diverse (Wilmshurst & McGlone 2005a). For these fields such as plant ecology, vegetation history reasons, palynologists often differ in the name and climate history, oil exploration and glaciol- they apply to a particular pollen grain or spore. ogy, and melissopalynology to determine the Easily recognized grains may be recorded in nectar source in honey. In all these studies, it is different ways for no apparent reason. Thus, important to identify the pollen grains and pollen derived from a monotypic taxon in New Zealand, e.g. Ascarina lucida, is often recorded spores as accurately as possible. However, not at the generic level in the same way as pollen all pollen grains can be assigned with certainty derived from a genus with several or many to a particular taxon. Factors influencing species, e.g. Metrosideros. Pollen also may be pollen production, pollen morphology, pollen referred to as a family, e.g. Myrtaceae, without dispersal, the habit and range of source plants, any attempt to separate genera, and therefore deposition and preservation of pollen and treated in the same way as those which are so spores (Wilmshurst & McGlone 2005a, b) and similar as to defy easy recognition, e.g. Poaceae. the process of sample preparation can all In some cases, the pollen of genera such as influence the final decision. Access to a regional Leptospermum and Kunzea (Myrtaceae) cannot pollen collection or pollen atlas (Moar 1993) be distinguished; in others, pollen of some may resolve some problems, but there are genera within a family such as Podocar- always grains that cannot easily be identified, paceae can be easily recognized, e.g. Dacrydium either because they have not been encountered cupressinum, whereas others cannot, e.g. before or because their characteristic features Halocarpus. A review of pollen diagrams

*Corresponding author. Email: [email protected]

ISSN 0028-825X print/ISSN 1175-8643 online # 2011 The Royal Society of New Zealand DOI: 10.1080/0028825X.2010.526617 http://www.informaworld.com 202 NT Moar et al. published over the last 30 years shows that as older sediments, particularly since these often pollen grains became progressively better have no known modern plant source to which known, the precision of identification has they can be assigned. In large families, e.g. improved. Despite this, the ambiguities we refer Cyperaceae, or in large genera, e.g. Hebe, not to persist. For example, sedge pollen is usually all taxa have been considered, an admission referred to as Cyperaceae, but as Moar and which extends to recently revised taxa, e.g. Wilmshurst (2003) have shown, it is possible to Montia, which now includes eight species achieve a more precise identification of the (Heenan 2007) rather than the one, Neopaxia sedges by recognizing different pollen types australasica, previously recognized, and to within the group. pollen of the Orchidaceae and Poaceae, which A difficulty for some who use pollen to date have not been studied in detail in New analytical data arises when new taxonomic Zealand. We suggest the following conventions names are adopted, but not explained by the for nominative use in pollen diagrams: palynologist. An example of this problem is the almost universal use of the subgeneric name 1. Use a specific name when there is only one Fuscospora for pollen formerly recorded as species represented in the region or country, Nothofagus fusca type, or the changes for the e.g. Dacrycarpus dacrydioides and Ascarina taxon once known as Dacrydium colensoi, then lucida, or when the pollen or spores of one as Lagarostrobos colensoi, and currently as species of a genus is clearly different from Manoao colensoi. It is important that an author the rest, e.g. Nothofagus menziesii and clearly states the nomenclatural basis for names Drosera arcturi. used in diagrams and any associated discussion. 2. Use a generic or subgeneric name when the The above comments apply as much to fern pollen or spores of different species within a spores as they do to pollen grains. However, it genus or subgenus are so similar that is not intended here to discuss spores in any separation is difficult or impossible under detail (covered by Large & Braggins 1991) routine analysis. Examples are Epilobium, except those of Cyatheaceae and Dicksonia- Fuscospora (a subgenus of Nothofagus that ceae. The spores of these tree ferns are found in includes N. fusca, N. solandri s.l. and N. most sediments because they are highly resis- truncata), Metrosideros and Myrsine. tant to decay, and are often well preserved, as 3. Use a generic name and ‘type’ when two or in peat, or variously degraded, as in lake more genera share the same pollen or spore sediments or in organic soils (Wilmshurst et type. Examples are Leptospermum type to al. 1999; Wilmshurst & McGlone 2005a). include pollen of Leptospermum and Kun- Although they are distinctive enough, it is not zea, and Phyllachne type to include Phyl- always easy to separate them from other lachne and Forstera. In selecting the generic similarly shaped spores such as Adiantum. name in these instances, we follow previous Accordingly, we include the Cyatheaceae and palynological usage. Dicksoniaceae in this discussion; comments are 4. Use specific name followed by ‘type’ when based on Large & Braggins (1991). the pollen or spores of different species The purpose of this review is to suggest how within a genus can be identified into groups best to refer to the pollen types and tree fern based on distinctive characters. For exam- spores recorded in a New Zealand pollen ple, Myosotis pollen can be distinguished diagram so as to avoid ambiguity, and to into Myosotis uniflora type, Myosotis angu- make for a more uniform presentation of the stata type and Myosotis australis type data. We restrict our treatment to late Qua- (Moar 1993); likewise, Cyathea spores can ternary material in order to avoid conflict with be distinguished into Cyathea smithii type form genera proposed by those working with and Cyathea dealbata type. Standardizing names in New Zealand palynology 203

5. Use a tribe or family name for recording Heywood et al. (2007) because of the distinct pollen or spores above generic level as differences in their pollen morphology. For appropriate, e.g. Lactuceae, Poaceae. some genera, alternative familial affinities are indicated in parentheses. We provide brief Although it is hoped that these conventions descriptions of pollen grains based mainly on will lead to a more uniform presentation of Cranwell (1953), Moar (1993) and Pocknall pollen diagrams, it is not suggested that refer- (1981a, b, c), and of the spores of tree ferns ence to species be omitted from any discussion (Large & Braggins 1991), together with the when it is appropriate to do so. Thus a pollen suggested name to be used for each pollen and diagram from Westland may record the pre- spore type recorded. Where relevant, we also sence of Metrosideros, but it needs to be stated provide brief comments. Finally, for each taxon that the probable source is Metrosideros um- we provide a categorical summary for the bellata, especially if it is abundant. When this pollen occurrence, pollination mode, type of pollen is recorded in high numbers from Auck- pollen production and dispersal based on data land Island peat samples, the source can only from published pollen diagrams to assist with be that of Metrosideros umbellata. However, a the interpretation of pollen diagrams. These conservative approach is recommended for categories are summarized for convenience in most standard pollen diagrams and all data- Table 1. Dispersal refers to occurrence of bases. Distinctions should go no lower than the significant quantities of a given pollen type, taxonomic level at which they can be reliably not simply presence. It is noted that for easy and consistently made. comparison, pollen and spore preparations In the following review, the pollen of most from sediments, and from modern reference gymnosperm and most angiosperm genera in- material (either fresh or herbarium specimens), digenous to New Zealand is considered, some are both processed using the standard acid in greater detail than others. Some have never treatment of acetolysis. Finally, we would like been recorded in a pollen analysis, some only to acknowledge the previous work on pollen occasionally, some frequently, while others are distribution by numerous palynologists in New ubiquitous. However, all types are considered Zealand, in particular Macphail & McQueen here because we may have overlooked some (1983). rare records of certain pollen grains, and some presently unrecorded types may be recognized POLYPODIOPSIDA (Filicopsida) in the future. The figure in parentheses following the Cyatheaceae generic name refers to the number of species GENUS: Cyathea (7). in that genus. Generic and specific names SPORES: Trilete; triangular in polar view; exine generally follow the Allan Herbarium thick (2Á3 mm) in all taxa except C. smithii and (2002Á2006), but more recent treatments are C. kermadecensis (1Á2 mm); spores generally followed for family names and taxa at a higher smooth after acetolysis; size variable. taxonomic rank (Heywood et al. 2007; Mab- NAMES: Cyathea: C. smithii type (C. colensoi, berley 2008). For this reason, Dianella, Herpo- C. kermadecensis, C. smithii); C. dealbata type lirion and Phormium are included in (C. cunninghamii, C. dealbata, C. medullaris, C. Hemerocallidaceae and Arthropodium and Cor- milnei). dyline are assigned to Laxmanniaceae because COMMENTS: C. kermadecensis and C. milnei are the pollen in these two groups are quite endemic to the Kermadec group. In most different. It is convenient for palynologists to analyses, the C. smithii type usually refers to retain the traditional families Callitrichaceae, C. smithii. C. smithii spores are often mista- Plantaginaceae and Scrophulariaceae following kenly identified as Pteridium esculentum despite 204 NT Moar et al.

Table 1 Categories used to summarise the occur- tuberculate (D. lanata) or obviously tuberculateÁ rence, pollination mode, production and dispersal reticulate (D. squarrosa); in these last two, taxa mechanism for each pollen type discussed. sculpture may vary between distal and prox- Criteria Category imal surfaces. NAMES: Dicksonia fibrosa; D. lanata; D. squar- Occurrence X the absence of any record in rosa. OXÁ4 pollen analyses COMMENT: Spores of Dicksonia squarrosa are 1 an occasional record most frequently recorded. 2 a regular occurrence SUMMARY: O4, W, P3, D2. 3 a regular occurrence, often in considerable numbers 4 ubiquitous types dependent PINOPSIDA upon the nature of source vegetation Araucariaceae GENUS: Agathis (1). Pollination W wind-pollinated taxa mode POLLEN: Grains spheroidal, inaperturate; exine W, I, V, A I pollination by insects or other thick, minutely scabrate or papillate. invertebrates NAME: Agathis australis. V pollination by vertebrates, COMMENT: Grains may be variously degraded birds, or other animals during preservation, especially through loss of A water pollinated papillae or the tectum. Pollen 1 low pollen production SUMMARY: O2, W, P3, D2. production P1Á32moderate pollen production 3 prolific pollen production Cupressaceae Dispersal 1 restricted distribution GENUS: Libocedrus (2). D1Á32a distribution within a few POLLEN: Grains small, thin walled and spher- kilometres of the source oidal; exine bears scattered papillae. 3 grains that may be widely NAME: Libocedrus. distributed, often over great COMMENTS: Grains are often split into two distances halves and consideration should be given to counting these as half grains. The papillae may be sloughed off. Pollen of introduced Cupres- the fact their exine is more than twice as thick saceae is difficult to separate from Libocedrus. as P. esculentum. Like Pteridium, tree fern In post-European sites, pollen should be re- spores may be over-represented through re- corded as Cupressaceae. working. SUMMARY: O3, W, P2, D3. SUMMARY: O4, W, P3, D2.

Podocarpaceae Dicksoniaceae GENERA: Dacrycarpus (1), Dacrydium (1), Ha- GENUS: Dicksonia (3). locarpus (3), Lepidothamnus (2), Manoao (1), SPORES: Trilete; triangular in polar view; exine Phyllocladus (3), Podocarpus (4), Prumnopitys thick (2Á3 mm), thickens abruptly at angles; (2). under light microscope surface sculpture varies POLLEN: Grains of all taxa are bisaccate with from smooth or scabrate (D. fibrosa), granulateÁ the exception of the trisaccate Dacrycarpus Standardizing names in New Zealand palynology 205 dacrydioides. Of the bisaccate grains, those of MAGNOLIOPSIDA Dacrydium cupressinum are easily recognized by 1. NYMPHAEALES the reduced size of the pollen sacs, which form a shallow frill around the body of the grain. Hydatellaceae The rather larger saccae of Halocarpus with GENUS: Trithuria (1). thick parallel markings allow for easy identifi- POLLEN: Grains monosulcate or monoporate; cation of this genus as do the small, plain, often exine very thin, sparingly perforate and bearing partly folded, saccae of Phyllocladus. Pollen of minute broad-based spinules difficult to re- Podocarpus totara and the related species, P. solve; longest axis36 mm. hallii and P. nivalis, are easily recognized by NAME: Trithuria inconspicua. strongly folded (rugulate) exine over the prox- COMMENT: These fragile grains are easily dis- imal cap. Grains of Prumnopitys ferruginea and torted and to date have not been recorded in a P. taxifolia are often confused; the larger pollen analysis. SUMMARY: OX, W, P1, D1. grains, clearly projecting saccae, with conspic- uous folds at the point of insertion, and thick proximal cap often smooth in the centre, 2. MAGNOLIIDAE s.s. usually separate the former from the latter. Pollen of Lepidothamnus and Manoao are Lauraceae similar, but the fine, regular reticulum of GENERA: Beilschmiedia (2), Cassytha (1), Litsea Manoao contrasts with the more open, broken (1). reticulum of both Lepidothamnus intermedium POLLEN: Grains echinate, delicate, often col- and L. laxifolium. lapsed; intectate; minor differences not easily NAMES: Dacrycarpus dacrydioides; Dacrydium resolved under bright field light microscopy. cupressinum; Halocarpus; Lepidothamnus; Man- NAME: Beilschmiedia type. oao colensoi; Phyllocladus; Podocarpus; Prum- COMMENTS: Grains recorded only a very few nopitys ferruginea; Prumnopitys taxifolia. times, and then usually as single examples; COMMENTS: Sometimes the pollen of Lepi- possibly one of the most under-represented dothamnus, Manoao, Podocarpus and Prumnop- pollen types in the flora. See discussion in itys are variously distorted and so may defy Macphail (1980). SUMMARY (general): O1, I, P1, D1. ready identification. In such cases, refer to as bisaccate-Podocarpaceae. Podocarp pollen types are always recorded when source plants Monimiaceae are present in the surrounding vegetation. GENERA: Hedycarya (1), Laurelia (1). Lepidothamnus is only rarely recorded, perhaps POLLEN: Grains of Hedycarya are inaperturate because it often resembles Prumnopitys taxifolia tetragonal tetrads; those of Laurelia are dicol- when not well displayed. It is not a major pate, finely reticulate monads, often splitting component of present-day vegetation. into halves. SUMMARY: Dacrycarpus dacrydioides O4, NAMES: Hedycarya arborea; Laurelia novae- W, P2, D3; Dacrydium cupressinum O4, zelandiae. W, P3, D3; Halocarpus O4, W, P2, D2; SUMMARY: Hedycarya arborea O1, I, P1, D2; Lepidothamnus O1, W, P2, D1; Manoao Laurelia novae-zelandiae O1, I, P1, D1. colensoi O1 (4), W, P2, D2; Phyllocladus O4, W, P3, D3; Podocarpus O3, W, P3, D3; Prumnopitys ferruginea O3, W, P3, D3; Piperaceae Prumnopitys taxifolia O4, W, P3, D3. GENERA: Macropiper (2), Peperomia (2). 206 NT Moar et al.

POLLEN: Grains small. Macropiper monocol- POLLEN: Grains monosulcate; sulcus long, nar- pate; colpus circular, its membrane bearing row, but wider at ends; exine up to 2 mm thick coarse exine fragments; exine faintly verrucate. on proximal surface; exine mostly smooth or Peperomia acolpate; exine coarsely verrucate. may be slightly pitted or vaguely rugulate. NAMES: Macropiper; Peperomia. NAME: Rhopalostylis. COMMENTS: Macropiper mostly rarely recorded, COMMENTS: Unlikely to be confused with other but occasionally frequent as on Tawhiti Rahi monosulcate pollen types. R. cheesemanii ( R. Island, Poor Knights Islands (Wilmshurst & baueri) occurs in the Kermadec group (Prebble Moar unpubl. data); Peperomia not recorded in & Dowe 2008), whereas R. sapida is the species pollen analyses. of the two main islands. In the South Island it SUMMARY: Macropiper, Peperomia O1, W, P1, occurs south to Banks Peninsula and Hokitika D1. in Canterbury and Westland, respectively. It is also found on Chatham Island. In the North Island it is recorded from sites in northern Winteraceae North Island, Great Barrier Island (Horrocks GENERA: Pseudowintera (4). et al. 2002), and Tawhiti Rahi of the Poor POLLEN: Grains tetrahedral tetrads; exine Knights Islands (Wilmshurst & Moar unpub- boldly reticulate; each tetrad member mono- lished data). Most, if not all, records from the porate. main islands refer to R. sapida. NAME: Pseudowintera. SUMMARY: O1 (4), I (W), P3, D1. COMMENT: The most widely distributed species is P. colorata, and in most instances grains recorded in Holocene samples are probably Asphodelaceae (Xanthorrhoeaceae) derived from plants of this taxon. GENUS: Bulbinella (6). SUMMARY: O1, I, P1, D1. POLLEN: Grains monosulcate; sulcus very long; exine rugulate, especially so in B. rossii; rugulae often absent on distal surface especially along 3. CHLORANTHALES sulcus margin. NAMES: Bulbinella rossii; Bulbinella. Chloranthaceae COMMENTS: The only grain easily distinguished GENUS: Ascarina (1). is that of B. rossii because of its much bolder POLLEN: Grains bilaterally symmetrical, mono- rugulae. This grain is frequently recorded from colpate; exine intectate, clavate; clavae ar- Auckland and Campbell islands (McGlone ranged in reticuloid pattern. 2002). Pollen of the other taxa, restricted to NAME: Ascarina lucida. the main islands, is much less boldly rugulate COMMENT: The characteristic clavae may be and only occasionally recorded. Although there partly or completely stripped from the exine; in are subtle differences in the nature and dis- subfossil preparations there may be a gradation tribution of the rugulae of these grains, identi- from complete grains through to those from fication is difficult. Rugulae, which are least which the clavae are completely absent. well developed on the grains of B. modesta, are SUMMARY: O3, W, P3, D2. sometimes absent, which may provide a means of further separation. However, a decision needs to be made taking into account the 4. MONOCOTYLEDONS location of the site and the number of grains recorded during an analysis. Arecaceae SUMMARY: Bulbinella rossii O3, I, P2, D1; GENUS: Rhopalostylis (2). Bulbinella O1, I, P2, D1. Standardizing names in New Zealand palynology 207

Asteliaceae elliptic to oval in polar view, longest axis up GENERA: Astelia (13), Collospermum (2). to 34 mm. POLLEN: Grains of Astelia and Collospermum NAME: Iphigenia novae-zelandiae. cannot be confused. Although both Astelia and COMMENT: Not recorded in pollen analyses. Collospermum grains are monosulcate, and in SUMMARY: OX, I?, P1, D1. both the sulci are long, Astelia grains are elliptic in shape and bear spinules up to 2 mm long, whereas Collospermum grains are generally Cyperaceae spheroidal, and the sulcus is often very broad. GENERA: Baumea (7), Carex (80), Carpha (1), Unlike C. microspermum, the exine of C. Cyperus (2), Desmoschoenus (1), Eleocharis (5), hastatum is perforate and bears scattered very Fimbristylis (1), Gahnia (6), Isolepis (14), Lepi- short, broad-based spinules. dosperma (3), Machaerina (1), Morelotia (1), NAMES: Astelia; Collospermum hastatum; Col- Oreobolus (3), Schoenus (9), Tetraria (1), Un- lospermum microsporum. cinia (32). COMMENTS: Both genera are recorded in pollen POLLEN: Two main types may be distinguished, analyses, but never in great numbers. However, spheroidal and variously cup-shaped or pear- in the subantarctic Auckland and Campbell shaped. Spheroidal grains occur in Baumea, Islands, Astelia pollen, either A. linearis or A. Fimbristylis, Lepidosperma, Machaerina, Tetra- subulata, may be an important element in ria and Oreobolus. Of these, Fimbristylis pollen pollen diagrams. Collospermum pollen, prob- is recognized by its coarsely verrucate exine and ably C. hastatum, is recorded occasionally from poorly defined apertures; the exine of Baumea, North Island sites. Lepidosperma, Machaerina, Tetraria and Oreo- SUMMARY: Astelia O1 (2), W (V), P2, D1; bolus is smooth, and of these, the apertures of Collospermum hastatum, C. microsporum O1, Oreobolus are difficult to resolve, in contrast to W (V), P2, D1. those of the other four genera. The grains of all other genera are broadly rectangular, and usually either cup-shaped or pear-shaped. The most easily recognized are Eleocharis and Centrolepidaceae Schoenus; the former by its very large elongate GENERA Centrolepis Gaimardia : (4), (1). apertures, relative to grain size, and the latter POLLEN: Grains ulcerate; the aperture rims by its narrow wedge shape. Pollen of E. formed by the irregular and abrupt ending of sphacelata is distinguished from other species the exine; the exine is perforate (scrobiculate) of that genus by its larger size, recorded as and the scrobiculi are often joined by shallow being between 35 and 46 mm long, in contrast to canals. the length of the other species (24Á31 mm). Of NAMES: Centrolepis type (mainland sites); Cen- the other taxa, Isolepis and Uncinia grains are trolepis for subantarctic sites. mostly cup-shaped, and with 9 circular or COMMENT: Gaimardia is not recorded from the elliptical lacunae. These characters separate subantarctic islands. them from the remaining genera, which possess SUMMARY: O1 (3), W, P1, D1. short lacunae, never more than half the length of the grain (Carex), by elongate and narrow lacunae (Cyperus ustulatus), or by diffuse, often Colchicaceae very narrow elongate lacunae (Carpha alpina). GENUS: Iphigenia (1). For further detail consult Moar & Wilmshurst POLLEN: Grains monosulcate; sulcus very long, (2003). sometimes wider at end; exine very thin,1 mm, NAMES: Baumea type (Baumea, Lepidosperma, minutely reticulate, heterobrochate; shape Machaerina and Tetraria); Carex; Carpha 208 NT Moar et al. alpina; Cyperus ustulatus; Eleocharis sphacelata; COMMENT: Rarely recorded in pollen analyses Eleocharis; Fimbristylis squarrosa; Gahnia type (Mildenhall 1980b). (Gahnia and Morelotia); Isolepis type (Isolepis SUMMARY: O1, W (I), P1, D1. and Uncinia); Schoenus type (Bolboschoenus, Desmoschoenus, Schoenoplectus and Schoenus); Oreobolus. Juncaceae COMMENTS: These notes, especially regarding GENERA: Juncus (16), Luzula (11), Marsippos- large genera such as Carex, are based on a permum (1), Rostkovia (1). limited number of taxa and should be seen as a POLLEN: Grains tetrahedral tetrads; each mem- guide only to identifying the pollen of this ber of the tetrad bears a large pore occupying difficult group. Pollen of the Baumea type, most of the distal surface; exine very thin, Isolepis type, Eleocharis and Schoenus type are delicate; faintly verrucate. the most easily recognized; if there is any NAME: Juncaceae. doubt, refer to Cyperaceae. COMMENTS: Acetolysis usually destroys the SUMMARY (general): O4, W, P2Á3, D1. internal walls of the tetrad leading to its collapse and destruction. Rarely recorded in pollen analyses (Harris & Mildenhall 1980). Hemerocallidaceae (Xanthorrhoeaceae) SUMMARY: O1, W, P2, D1. GENERA: Dianella (3), Herpolirion (1), Phor- mium (2). POLLEN: Grains trichotomosulcate; exine reti- Juncaginaceae culate (Phormium), or microreticulate GENUS: Triglochin (2). (Dianella, Herpolirion); shape triangular or POLLEN: Grains inaperturate; finely reticulate, sub-triangular; Phormium tenax grains, 35Á37 reticulum not easily resolved; shape 9 spher- mm, are the largest; P. cookianum,27Á32 mm, oidal; size20 mm. slightly smaller; Dianella,19Á27 mm, and Her- NAME: Triglochin. polirion,22Á31 mm, are considerably smaller COMMENTS: The poorly resolved reticulum and may be difficult to separate. separates Triglochin from Potamogeton in NAMES: Herpolirion type (Dianella and Herpo- which the reticulum is clearly resolved. Not lirion); Phormium (P. tenax and P. cookianum). recorded in pollen analyses. Dianella pollen has been occasionally recorded SUMMARY: OX, W, P1, D1. (e.g. Mildenhall 1980b). COMMENTS: Of these genera, Phormium is the most likely to be recorded. Phormium pollen is Laxmanniaceae (Asparagaceae) extremely under-represented in relation to its GENERA: Arthropodium (3), Cordyline (5). prominence in many wetland areas. POLLEN: Grains monosulcate; sulcus very long. SUMMARY: Herpolirion type O1, I, P1, D1; Exine of Arthropodium1 mm thick, reticulate, Phormium O1, V (I), P2, D1. heterobrochate; simplibaculate; longest axis45 mm. Exine of Cordyline varies in thickness from B1 mm(C. australis)to2mm Iridaceae (C. pumilio); surface undulating; surface of C. GENUS: Libertia (8). indivisa sparingly perforate, perforations small, POLLEN: Grains monosulcate; exine microreti- visible only by lux obscuritas analysis (LO, see: culate; size variable between 27 and 40 mm. Moore et al. 1991). NAME: Libertia NAMES: Arthropodium; Cordyline. Standardizing names in New Zealand palynology 209

COMMENTS: Pollen of these two genera cannot Ehrhartoideae, Pooideae, Arundinoideae, Pa- be confused. The latter is more likely to be nicoideae and Chloridoidaea (Edgar & Connor recorded, and then only occasionally. 2000); pollen of Chloridoidaea have not been SUMMARY: Arthropodium O1, I, P2, D1; Cordy- examined. Genera examined are: Microlaena line O1, I, P2, D1. (Ehrhartoideae); Achnatherum, Anemanthele, Austrostipa, Deschampsia, Deyeuxia, Elymus, Festuca, Hierochloe, Lachnagrostis, Poa, Trise- Luzuriagaceae (Alstroemeriaceae) tum (Pooideae); Chionochloa, Cortaderia, Ryti- GENUS: Luzuriaga (1). dosperma (Arundinoideae); Isachne, Spinifex POLLEN: Grains monosulcate; sulcus often cir- (Panicoideae). Some genera, e.g. Anemanthele, cular, occupying much of distal pole, margins are represented in New Zealand by only one irregular; exine surface areolate; shape spher- species, others by numerous species, e.g. Chio- oidal; size32 mm. nochloa and Rytidosperma. NAME: Luzuriaga. POLLEN: Grains monoporate; pores with thick- COMMENT: Rarely recorded in pollen analyses ened annulus; exine1 mm thick, tectate, bacu- (Mildenhall 1980a). late; bacula often difficult to resolve; at 1000 SUMMARY: O1, I, P1, D1. the tectum may carry separate spinules (e.g. Chionochloa), groups of spinules on ‘islands’ separated by shallow channels (e.g. Poa), or Orchidaceae there may be an intermediate state in which COMMENTS: Detailed comment is reserved until some spinules are in groups, whereas ongoing revisions are completed (Brian Molloy others appear to be single (e.g. Cortaderia). personal communication). Pollen may occur as The apparent ‘islands’ are interpreted to be the tetrads, monads, or as pollinia. All grains result of an undulating exine surface; shape examined are reticulate; the exine of different spheroidal; size based on the limited number of taxa varies in thickness, and there is consider- genera examined varies between 23 and 49 mm. able variation in size of individual grains. NAMES: Chionochloa type (all species of Chiono- Pollen can be identified to family level without chloa examined, Lachnagrostis filiformis); Poa difficulty, but they are rarely seen in the fossil type (species examined Deyeuxia aucklandica, record. Hierochloe fusca, H. brunonis, Poa breviglumis, NAME: Orchidaceae. P. cita, P. colensoi, Rytidosperma caespitosum, R. SUMMARY: O1, I, P2, D1. unarede, Spinifex sericeus, Trisetum spicatum); Cortaderia type (Cortaderia spp., Hierochloe redolens, Rytidosperma clavatum and R. macula- Pandanaceae tum). Whenever in doubt, refer to Poaceae. GENUS: Freycinetia (1). COMMENTS: A close examination of the exine POLLEN: Grains monoaperturate; aperture 9 surface, especially by phase contrast, offers circular, grain pouch-shaped. some hope for a better understanding of the NAME: Freycinetia banksii. pollen grains, but studies by electron micro- COMMENT: Sometimes recorded in high num- scope, scanning electron microscope in parti- bers, especially in Westland. cular, are needed to make any progress. SUMMARY: O1, I (V), P3, D2. Chionochloa species, and others, have clearly separated spinules although the degree of separation may depend upon the vagaries of Poaceae preparation. Taxa in which ‘islands’ are GENERA: There are 34 genera native to New most consistently seen occur in the genus Poa, Zealand distributed between five subfamilies, although there is considerable variation 210 NT Moar et al. between species. Pollen of 10 specimens named between Empodisma and Leptocarpus, but eco- Poa colensoi collected by B. Molloy from logical considerations will often offer the best different parts of Otago and Canterbury, and clue as to identification. one from Mount Ruapehu, were examined. The SUMMARY (general): O1, W, P2, D1. exine ‘islands’ varied considerably; some were long and narrow, some were 9 circular and some, especially on pollen from the Mount Smilacaceae Ruapehu specimen, were broad and 9 rectan- GENUS: Ripogonum (1). gular in shape. Long and narrow ‘islands’ have POLLEN: Grains monosulcate; sulcus extending been described as ruguloid. Intermediate ar- along length of distal surface; exine microreti- rangements in which the spinules are aggre- culate; muri very narrow, bearing very short gated into groups are found in the Cortaderia spinules. type. NAME: Ripogonum scandens. SUMMARY (general): O4, W, P2, D3. COMMENTS: The spinules are not easily resolved under bright field light microscopy or even with phase contrast and require scanning electron Potamogetonaceae microscopy study for confirmation. The pollen has been recorded as frequent in some Taranaki GENUS: Potamogeton (4). sites. POLLEN: Grains inaperturate; exine very thin, SUMMARY: O1, I (W), P1, D1. semitectate, baculate, reticulate; reticulum het- erobrochate, delicate. NAME: Potamogeton. Typhaceae COMMENT: Pollen grains of Potamogeton and GENERA: Sparganium (1), Typha (1). Triglochin (Juncaginaceae) are similar but the POLLEN: Grains monoporate, pore circular; easily resolved, delicate reticulum in Potamo- exine slightly raised around pore to form a geton separates it from Triglochin. distinct ring; exine reticulate, muri of variable SUMMARY: O1, W (A), P2, D1. thickness and mostly duplibaculate. NAMES: Sparganium subglobosum; Typha orien- talis. Restionaceae COMMENT: Typha and Sparganium may be GENERA: Empodisma (1), Leptocarpus (1), Spor- distinguished by differences in exine structure adanthus (2). and by differences in the pore margins. POLLEN: Grains monoporate; pores large, often SUMMARY: Sparganium subglobosum O1, W, P1, 9 circular; exine perforate (scrobiculate), un- D1; Typha orientalis O1 (3), W, P3, D1. dulate and marked by grooves or channels that emphasize the undulate nature of the exine, especially in Sporadanthus. Xeronemataceae NAMES: Empodisma type (Empodisma and Lep- GENUS: Xeronema (1). tocarpus); Sporadanthus. POLLEN: Grains monosulcate; exine microreti- COMMENTS: Of these three genera, Spora- culate, muri coarse, duplibaculate. danthus is most easily recognized by its clearly NAME: Xeronema callistemon. undulate tectum, by the narrow clear grooves COMMENTS: Occasionally recorded on Poor or channels separating the undulations, and by Knights group (Wilmshurst & Moar unpub- the numerous evenly distributed perforations lished data); pollen of Xeronema and Arthro- (scrobiculi). There are subtle differences podium (Laxmanniaceae) are similar although Standardizing names in New Zealand palynology 211 the duplibaculate muri of the former separate it NAME: Alternanthera. from the latter. COMMENTS: These grains cannot be confused SUMMARY: O1, V, P1 (2), D1. with any other New Zealand pollen type; they are rarely recorded in a pollen analysis (Mil- denhall 1980b). The two species in New Zeal- 5. EUDICOTS and are A. nahui and A. denticulata. Of these, the former is endemic, but the status of the Argophyllaceae latter is uncertain and may be naturalized. GENUS: Corokia (3). SUMMARY: O1, W, P2, D1. POLLEN: Grains tricolporate, endoapertures H- shaped; exine faintly rugulate. NAME: Corokia. Apiaceae COMMENT: Not noted in pollen analyses. GENERA: Aciphylla (40), Actinotus (1), Aniso- SUMMARY: OX, I, P1, D1. tome (16), Apium (1), Azorella (1), Centella (1), Chaerophyllum (4), Daucus (1), Eryngium (1), Gingidia (7), Hydrocotyle (10), Lignocarpa (2), Aizoaceae Lilaeopsis (2), Scandia (2), Schizeilema (11), GENERA: Disphyma (2), Tetragonia (2). Stilbocarpa (3). POLLEN: Grains tricolpate (Disphyma australe, POLLEN: Grains easily recognized at the family Tetragonia tetragonioides) or mostly tetracol- level; identification sometimes possible at the pate (D. papillatum, T. trigyna); tectum min- generic level, and occasionally at the specific utely spinulose (D. australe, Tetragonia spp.), level. reticulate (D. papillatum), perforate (D. aus- Aciphylla, Anisotome and Scandia are similar, trale); ectoapertures have raised margins pro- but Aciphylla pollen is distinguished by its ducing a pouting effect in polar view, obviously prolate shape and pointed poles, in sometimes syncolpate (D. australe). contrast to the sub-prolate shape and rounded NAMES: Disphyma australe type; D. papillatum poles of Anisotome and Scandia. type. Actinotus, Apium, Daucus and Lilaeopsis are COMMENT: Only rarely noted in pollen analyses. easily confused, although the long ectoaper- SUMMARY (general): O1, I, P1, D1. tures of Actinotus and Apium ( two-thirds length of grain) contrast with those of Daucus and Lilaeopsis (two-thirds length of grain or Alseuosmiaceae less). GENUS: Alseuosmia (7). Centella and Eryngium are similar, although POLLEN: Grains variable, mainly tricolporate, some grains of Centella may be tetra- or angulaperturate; tectum verrucate or rugulo- hexacolporate. verrucate. Chaerophyllum pollen may be recognized by the NAME: Alseuosmia. crimped appearance of the exine when seen in COMMENT: Rarely recorded (Mildenhall 1980b). optical, equatorial view. SUMMARY: O1, I, P1, D1. Gingidia and Lignocarpa pollen grains are characterized by markedly protruding aper- tures and are difficult to separate. Amaranthaceae Hydrocotyle grains have obviously ‘offset’ po- GENUS: Alternanthera (2). lar exine, a characteristic shared with Azorella POLLEN: Grains with12 deeply recessed aper- and with some Schizeilema. Small rhomboidal tures, separated by narrow, finely spinulose grains with pointed or rounded poles identify muri. Azorella selago and H. moschata. Pollen of 212 NT Moar et al. other taxa may vary in size, aperture number POLLEN: Typically the pollen is reticulate, sub- and surface pattern, but separation is difficult, prolate, and the endoapertures are lalongate, especially as few grains are recorded in any relatively short and rectangular in shape. Pollen pollen count. However, the larger grains of H. of Meryta sinclairii has the boldest reticulum novae-zeelandiae may sometimes be recognized. with lumina up to 5 mm across and is distinct Schizeilema is identified by its strongly ‘bowed’ from other New Zealand Araliaceae. The shape in equatorial view; polar exine in S. reticulum of Raukaua type (Mitchell et al. hydrocotyloides is strongly ‘offset’, and tetra- 1997) is very small, often B 1 mm across, and colporate grains occur in S. allanii and S. sometimes difficult to resolve. Pollen of Pseu- colensoi. dopanax chathamicus, P. crassifolius, P. disco- Stilbocarpa grains are psilate, prolate and have lor, P. ferox and P. linearis are close in general narrow lalongate endoapertures fused to form appearance to those of Raukaua, but the an equatorial ring. reticulum is easily resolved. Pollen of the NAMES: Aciphylla; Anisotome type (Anisotome, remaining Pseudopanax species are bolder, Scandia); Centella type (Centella, Eryngium); and P. colensoi is recognized either by a Chaerophyllum; Gingidia type (Gingidia, Ligno- markedly rugulate mesocolpium or by irregu- carpa); Hydrocotyle type (Hydrocotyle moscha- larly spaced verrucae and spines. The grains of ta, Azorella selago); Hydrocotyle pterocarpa the remaining species of Pseudopanax, includ- type (Hydrocotyle dissecta, H. heteromeria, H. ing P. arboreus, come between the two extremes novae-zeelandiae, H. pterocarpa, H. sulcata); noted above, while Schefflera is generally Lilaeopsis type (Lilaeopsis novae-zelandiae, L. smaller, prolateÁspheroidal, and the rectangu- ruthiana, Actinotus novae-zealandiae, Apium lar endoapertures are sometimes H-shaped. prostratum, Daucus glochidiatus); Schizeilema, NAMES: Meryta sinclairii; Pseudopanax colensoi; Stilbocarpa. Pseudopanax arboreus type (P. arboreus, P. COMMENTS: Aciphylla, Hydrocotyle type and kermadecensis, P. laetus); Pseudopanax crassi- Lilaeopsis type pollen are most frequently folius type (P. chathamicus, P. crassifolius, P. encountered. Although Azorella selago pollen discolor, P. ferox, P. linearis); Raukaua (R. is close to that of Hydrocotyle moschata type, anomalus, R. edgerleyi, R. simplex); Schefflera most grains from Holocene sites are likely to be digitata. Refer to Araliaceae if uncertain. derived from Hydrocotyle moschata. COMMENTS: Pollen of the Pseudopanax arboreus SUMMARY (general): O1, I, P2, D1. type are most frequently observed. P. colensoi is sometimes seen in higher altitude sites from the South Island. Apocynaceae SUMMARY (general): O1 (2), I, P2, D2. GENUS: Parsonsia (3). POLLEN: Grains heteropolar, diporateÁtetrapo- Asteraceae rate, but mostly triporate. GENERA: There are 32 genera distributed among NAME: Parsonsia. five tribes, the Anthemideae, Astereae, Gna- COMMENT: Occasionally recorded; may be con- phalieae, Lactuceae and Senecioneae. fused with Geniostoma (Loganiaceae). POLLEN: Grains of Anthemideae and Lactuceae SUMMARY: O1, I, P2, D1. are easily recognized, but those of the Astereae, Gnaphalieae and Senecioneae are often difficult to separate. Araliaceae Pollen of the Anthemideae are characterized by GENERA: Meryta (1), Pseudopanax (14), Rau- a two-layered exine comprising an inner layer kaua (3), Schefflera (1). of coarse bacula supporting a diffuse tectum Standardizing names in New Zealand palynology 213 carrying short, fine bacula, which in turn other Lactuceae); Olearia colensoi type (O. support a perforate, echinate tectum. Haastia colensoi); Celmisia type (all Celmisia spp., grains (tribe Senecioneae) also have a two- Damnamenia vernicosa, Pachystegia spp., Pleur- layered exine, but the inner layer is amorphous ophyllum speciosum); Raoulia type (all Raoulia and has no obvious baculi; these have not been spp., Ozothamnus); Senecio type (all Senecio- recorded. neae); Asteraceae for all others or whenever Lactuceae grains are fenestrate and character- recognition is uncertain. ized by large depressed areas (lacuneae), sepa- COMMENTS: Grains of the Anthemideae and rated by high, narrow ridges of echinate exine. Lactuceae are generally distinguished easily The complexity of the apertures, involving an enough, but those referred to the other three ectoaperture, a mesoaperture, and an endoa- tribes are more difficult and see notes above. perture, is not often clear. Of the sixgenera Because Olearia lyallii is considered to have (Embergeria, Kirkianella, Microseris, Picris, been introduced to the Auckland Islands early Sonchus, Taraxacum) Embergeria is easily re- in the 19th century, pollen of the O. colensoi cognized by its long spines, up to 6 mm long, type recorded from the Auckland Islands and by much reduced lacunae and exine ridges; before the European era may be referred to as it has never been recorded. Pleurophyllum. Embergeria has not been re- Pollen of some in the Astereae, including corded. Olearia colensoi, O. lyallii, Pleurophyllum crini- SUMMARY (general): O3, I, P3, D2. ferum and P. hookeri, are characterized by short, close-set spines, and together comprise a well-defined group. Others, including Pleur- Avicenniaceae (Acanthaceae) ophyllum speciosum, Damnamenia vernicosa, GENUS: Avicennia (1). most Celmisia spp. and Pachystegia spp., POLLEN: Grains tricolporate; endoapertures characterized by acute, generally crowded small, slightly lolongate with sinuous margins, spines 4Á5 mm long, cannot easily be distin- often indistinct; exine up to 3 mm thick, guished from each other. reticulate, muri generally wide. Pollen of Raoulia and Ozothamnus (Gnapha- NAME: Avicennia marina. lieae) are difficult to separate. Their ectoaper- COMMENTS: Unlikely to be confused with pollen tures are rarely more than half the length of the of other taxa and occasionally recorded. Mil- grain, which distinguishes them from pollen of denhall and Brown (1987) have recorded it in other tribes, and their spines are usually no mid-Holocene sites south of its present limit in more than 2Á3 mm long. the North Island. Senecioneae pollen, like the Lactuceae, possess SUMMARY: O1, I, P1, D1. compound apertures (Moar 1993); spines tend to be blunt, in contrast to the acutely tipped spines of other groups, and in many a hole is Balanophoraceae visible (e.g. Brachyglottis repanda); ectoaper- GENUS: Dactylanthus (1). tures are mostly more than two-thirds the POLLEN: Grains characterized by 9Á12 irregu- length of the grain, and the lalongate endoa- larly distributed circular pores, each with an pertures are always very long, up to 14 mmor obvious annulus. longer. NAME: Dactylanthus taylorii. All other taxa are so similar that they cannot COMMENTS: Pollen of this bat-pollinated plant easily be distinguished and may be recorded as is occasionally recorded in late Quaternary Asteraceae. sediments (Macphail & Mildenhall 1980). For NAMES: Leptinella type (Leptinella, Cotula); a rare, low-growing plant, it is surprising that it Embergeria grandifolia; Kirkianella type (all is recorded at all. Wilmshurst (1995) suggests 214 NT Moar et al. that the pollen may be distributed in bat Brassicaceae droppings. GENERA: Cardamine (7), Lepidium (9), Nototh- SUMMARY: O1, V, P1, D1. laspi (2), Pachycladon (9), Rorippa (3). POLLEN: Grains tricolpate; reticulate; bacula and lumina vary, up to 3 mm long and 3Á4 Bignoniaceae mm across (Cardamine and Notothlaspi), bacula GENUS: Tecomanthe (1). 2 mm long and lumina 2 mm across (Lepidium, POLLEN: Grains tricolpate; ectoaperture split in Pachycladon, Rorippa), or bacula and lumina two or more places; exine boldly reticulate, not more than1 mm(Pachycladon cheesema- reticulum heterobrochate; muri relatively nii); easily recognized at the family level. coarse, duplibaculate. NAME: Cardamine type (Cardamine, Notothlas- pi); Lepidium type (Lepidium, Pachycladon NAME: Tecomanthe speciosa. Rorippa); Pachycladon cheesemanii. COMMENTS: Restricted to the Three Kings COMMENTS: Brassicaceae pollen is not often Islands. The splits across the ectoaperture are recorded; the three pollen types are not easily similar to those of Myoporum, but the bold distinguished. If in doubt, refer to Brassicaceae. reticulum makes confusion unlikely. Not re- SUMMARY (general): O1, I, P1, D1. corded in a pollen analysis. SUMMARY: OX, I, P1, D1. Callitrichaceae (Plantaginaceae) GENUS: Callitriche (4). Boraginaceae POLLEN: Grains small, thin walled; reticuloid or GENERA: Myosotidium (1), Myosotis (34). retipilate; vaguely colpoid. POLLEN: Grains of both genera are heterocol- NAME: Callitriche. Myosotidium hortensium pate. pollen is very COMMENTS: Occasionally recorded; exine struc- small, mostly B 11 mm long, in contrast to the ture and the amphibious or terrestrial habit of larger, but still small grains of Myosotis which most taxa suggest that pollination may be may be up to 22 mm long. Myosotis grains are anemophilous (see Martinsson 1993) or geito- divided into three groups: (a) Simple colpi are nogamous (Osborn & Philbrick 1994). Mole- parasyncolpate forming a characteristic polar cular studies support inclusion in the cap, e.g. M. uniflora. (b) Grains are flattened at Plantaginaceae, but here we follow Heywood the poles and constricted at the equator, e.g. M. et al. (2007) who on the basis of morphological angustata. (c) Grains are rounded at the poles characteristics prefer retention of the family and not constricted at the equator, e.g. M. Callitrichaceae. Pollen cannot be confused with australis. Plantago or members of the Scrophulariaceae. NAMES: Myosotidium hortensium; Myosotis uni- SUMMARY: O1, A?, P1, D1. flora type; Myosotis angustata type; Myosotis australis type. COMMENTS: Of the Myosotis pollen grains those Campanulaceae of M. uniflora type are the most easily recog- GENERA: Colensoa (1), Lobelia (11), Wahlenber- nized. If there is any doubt, refer to Myosotis. gia (10). Myosotidium has been recorded from the Cha- POLLEN: Grains usually triporate (Wahlenber- tham Islands by Mildenhall (1994). gia), or tricolporate (Colensoa, Lobelia). Exine SUMMARY: Myosotis types O1, I, P1, D1; bearing very short, evenly distributed spinules Myosotidium hortensium O1, I, P1, D1. (Wahlenbergia); mostly finely reticulate in Standardizing names in New Zealand palynology 215

Colensoa and Lobelia, but rugulo-striate in L. may have as many as 90. These numbers are fatiscens. Endoapertures lalongate in Lobelia, not helpful in pollen analysis because there is but H-shaped in Colensoa. considerable variation in both pore number and NAMES: Colensoa physaloides; Lobelia; Wahlen- size in any taxon, a difficulty compounded by bergia. the very few grains usually recorded in any COMMENTS: Pollen grains of C. physaloides, sample. restricted to the northern North Island, are NAME: Chenopodiaceae. not recorded; pollen of Lobelia is rarely re- COMMENTS: Sarcocornia pollen is the most corded; and the pollen of W. gracilis is tripor- distinctive in terms of pore number and size. ateÁhexaporate, but mostly tetraporate. The If site characteristics are appropriate, and the occasional grains noted during pollen analysis number of grains numerous enough, Sarcocor- are mostly triporate. nia quinqueflora may be used. Some pollen may SUMMARY (general): O1, I, P1, D1. be derived by long-distance transport from Australia, e.g. McGlone & Meurk (2000). SUMMARY: O1, W, P1, D1. Caryophyllaceae GENERA: Colobanthus (16), Scleranthus (3), Spergularia (1), Stellaria (6). Convolvulaceae POLLEN: Grains periporate except Spergularia, GENERA: Calystegia (3), Convolvulus (3), Di- which are tricolpate; exine spinulose, except in chondra (2), Ipomoea (2). Scleranthus and Spergularia; perforate in all POLLEN: Grains either tricolpate (Convolvulus, taxa; number of pores in Stellaria B20, in Dichondra) or periporate (Calystegia, Ipomoea). Colobanthus and Scleranthus 30. Convolvulus is the larger of the tricolpate grains NAMES: Colobanthus; Scleranthus; Spergularia (polar axis 55Á62 mm), and unlike the very thin, media; Stellaria. irregularly spinulose exine of Dichondra, the COMMENTS: Colobanthus and Scleranthus are exine of Convolvulus is up to 5 mm thick, close, but the absence of spinules in Scleranthus perforate and coarsely baculate. The periporate separates the two genera. Colobanthus is the grains (Calystegia, Ipomoea) are spheroidal, most frequently recorded of the two genera. very large, up to 91 mm for Ipomoea, with a SUMMARY (general): O1, I, P1, D1. thick (5 mm) perforate exine. Calystegia grains are spinulose, those of Ipomoea bear spines up Celastraceae to 8 mm long with very short spinules scattered in between. GENUS: Stackhousia (1). NAMES: Convolvulus; Dichondra; Calystegia; POLLEN: Grains reticulate; bacula short, often Ipomoea. projecting from the floor of the lumina singly, COMMENT: Only Dichondra and Calystegia in pairs, or in groups of three. recorded and then only occasionally. NAME: Stackhousia minima. SUMMARY (general): O1, I, P1, D1. SUMMARY: O1, I, P1, D1.

Chenopodiaceae (Amaranthaceae) Coriariaceae GENERA: Atriplex (4), Chenopodium (3), Einadia GENUS: Coriaria (8). (2), Sarcocornia (1), Suaeda (1). POLLEN: Grains small, usually tricolporate; POLLEN: Grains periporate, often with70 ectoapertures short, slit-like; endoapertures pores, but number varies considerably, e.g. short, about as broad as long. Einadia has60, Suaeda80, and Sarcocornia NAME: Coriaria. 216 NT Moar et al.

COMMENTS: Tetracolporate grains may occur. Donatiaceae (Stylidiaceae) The generally smaller grains, shorter apertures, GENUS: Donatia (1). and the absence of endocracks easily separate POLLEN: Grains of Donatia novae-zelandiae are Coriaria from Coprosma. mostly tricolporate in contrast to the colpate SUMMARY: O1, W, P2, D2. grains (tri-, tetra- or pentacolpate) of Forstera, Oreostylidium and Phyllachne sometimes asso- ciated with it, but here placed in Stylidiaceae. In Corynocarpaceae Donatia the often obscure endoapertures are H- GENUS: Corynocarpus (1). shaped. POLLEN: Grains dicolporate; heteropolar; ellip- NAME: Donatia novae-zelandiae. tical in polar view. SUMMARY: O1, I, P1, D1. NAME: Corynocarpus laevigatus. COMMENT: Rarely recorded. SUMMARY: O1, I, P1, D1. Droseraceae GENUS: Drosera (6). POLLEN: Grains spinulose tetrads; in Drosera Crassulaceae arcturi the tetrad is a compact, 8Á12-porate unit; in all other species, the units of the tetrad GENUS: Crassula (12). are distinguished by stalked proximal surfaces, POLLEN: Grains small; ectoapertures very long, sometimes fusing to form a polar cap; endoa- essentially the germinating apparatus. pertures H-shaped. NAMES: Drosera arcturi; Drosera spatulata type (all other species of Drosera). NAME: Crassula. COMMENTS: Drosera arcturi pollen is easily COMMENT: Rarely recorded. distinguished from other Drosera pollen. Spine SUMMARY: O1, I, P1, D1. length varies in pollen of the other five species, but is not reliable enough as a character to permit separation. Only occasionally recorded. Cucurbitaceae SUMMARY (general): O1, I, P1, D1. GENUS: Sicyos (1). POLLEN: Grains 7Á9 colpate; exine minutely reticulate; tectum spinulose. Elaeocarpaceae NAME: Sicyos australis. GENERA: Aristotelia (2), Elaeocarpus (2). COMMENT: Not recorded in pollen analyses. POLLEN: Grains tricolporate, small; exine SUMMARY: OX, I, P1, D1. smooth in Elaeocarpus, faintly and finely re- ticulate in Aristotelia. NAMES: Aristotelia; Elaeocarpus. Cunoniaceae COMMENTS: Elaeocarpus grains are smaller than GENERA: Ackama (2), Weinmannia (2). those of Aristotelia. Aristotelia grains recorded POLLEN: Grains small, reticulate; in Ackama the in glacial sites are probably from A. fruticosa. reticulum can only be resolved under oil SUMMARY: Aristotelia O1, I, P1, D2; Elaeocar- immersion; in Weinmannia the reticulum is pus O2, I, P1, D2. easily resolved at 400 magnification. NAMES: Ackama; Weinmannia. COMMENT: Pollen recorded in South Island sites Elatinaceae may be treated as Weinmannia racemosa. GENUS: Elatine (1). SUMMARY: Ackama O1, I, P2, D1; Weinmannia POLLEN: Grains are of two forms; in the first O4, I (W), P3, D2. the tectum has clustered perforations in Standardizing names in New Zealand palynology 217 mesocolpia; in the second the exine is finely and Pollen of Androstoma empetrifolia and Cy- faintly reticulate. athodes pumila generally occur as dyads be- NAME: Elatine gratioloides. cause two members of the tetrad are aborted; COMMENT: Not recorded in pollen analyses. the aborted cells are usually small, and scrobi- SUMMARY: OX, I, P1, D1. culi are scattered over the exine. In Lepteco- phylla juniperina and L. robusta the aborted grains are not much smaller than the fertile Ericaceae grain and the exine is smooth. GENERA: Acrothamnus (2), Androstoma (2), NAMES: Archeria; Cyathodes (Acrothamnus co- Archeria (2), Cyathodes (2), Dracophyllum ( lensoi, A. suaveolens, Androstoma empetrifolia, 35), Epacris (2), Gaultheria (12), Leptecophylla Cyathodes dealbata, C. pumila); Dracophyllum (2), Leucopogon (5), Pentachondra (1), Sprenge- or Dracophyllum and Oreothamnus; Epacris lia (1). alpina; Epacris pauciflora; Leucopogon colensoi; POLLEN: Grains tetrahedral tetrads. In most Leucopogon fasciculatus; Leucopogon fraseri; taxa the four members of the tetrad are fully Pentachondra pumila; Sprengelia incarnata. developed (T-type grains). Of these Archeria COMMENT: Mostly occasional, but some, e.g. and Dracophyllum subgenus Oreothamnus Dracophyllum, may be frequent as in the Auck- grains have very short ectoapertures (5Á10 mm land, Campbell and Chatham islands (Milden- long), in contrast to the long or very long hall 1994; McGlone et al. 2000, 2007). ectoapertures (14Á32 mm) of the others. SUMMARY (general): O1, I, P1, D1. The exine of these T-type grains is smooth (Archeria, Acrothamnus colensoi, Pentachondra pumila), verrucateÁechinate (Epacris alpina), Euphorbiaceae finely reticulate (E. pauciflora) or faintly rugu- GENERA: Euphorbia (1), Homalanthus (1), Por- late (Gaultheria); Dracophyllum grains have an anthera (2). intectate area in the mesocolpia. Sprengelia POLLEN: Grains medium (Homalanthus, Por- incarnata is recognized by its thick perforate anthera) to large (Euphorbia); ectoapertures exine often broken into coarse exinous patches. very long, fosse-aperturate in Homalanthus; Pollen of the two subgenera of Dracophyllum, endoapertures obscure (Euphorbia), large rec- Dracophyllum and Oreothamnus, can be distin- tangular (Poranthera), or long, narrow, taper- guished; the former by longer ectoapertures (20 ing (Homalanthus). mm) and straighter sides, e.g. D. menziesii, the NAMES: Homalanthus polyandrous; Euphorbia latter by shorter ectoapertures and obviously glauca; Poranthera. lobed appearance, e.g. D. muscoides. Penta- SUMMARY (general): OX, I, P1, D1. chondra pumila can be distinguished from Acrothamnus colensoi by shorter, broader ec- toapertures and by more numerous and more Fabaceae easily resolved endocracks; the ectoapertures of GENERA: Canavalia (1), Carmichaelia (22), Leucopogon colensoi may be bifurcate. Clianthus (2), Montigena (1), Sophora (8). The tetrads of the remaining taxa are aborted in POLLEN: Grains mostly prolateÁsubprolate; tri- varying degrees. The three aborted components colporate, endoapertures lalongate; exine var- of Leucopogon fraseri are so reduced that the iously reticulate. Exceptions are Canavalia pollen may be mistaken for monads (S-type rosea and Sophora. Canavalia rosea grains are grains); they are heavily patterned with endo- anisopolar, with smooth and very thick exine cracks. By contrast, the three aborted grains of often broken into exinous islands at one pole; L. fasciculatus, although small, are usually bacula short. Endoapertures in Sophora are obvious as separate entities. lolongate. Montigena grains are similar to those 218 NT Moar et al. of Sophora, but endoapertures tend to be often over-represented, and some are found in lalongate. Most Carmichaelia grains are finely almost all sites, including the subantarctic reticulate and cannot be distinguished although islands; Nothofagus menziesii grains are gener- there are differences in size. However, C. ally under-represented, but they may be well carmichaeliae, C. crassicaulis, C. glabrescens, represented if source plants are locally domi- C. muritai, C. stevensonii, and C. torulosa are nant. generally more boldly reticulate, especially C. SUMMARY: Fuscospora O4, W, P3, D3; Notho- stevensonii. Clianthus grains are generally larger fagus menziesii O4, W, P3, D2. than those of the other taxa. NAMES: Canavalia rosea; Carmichaelia (Carmi- chaelia), all species; Clianthus (Clianthus max- Gentianaceae imus, C. puniceus); Sophora type (Sophora, Montigena). GENERA: Gentianella (30), Sebaea (1). POLLEN: Grains easily recognized by their bold COMMENTS: The only recorded pollen types are those of Carmichaelia and Sophora. The more reticulate or striateÁreticulate exine. Gentianella boldly reticulate grains of Carmichaelia have grains are larger (polar axis of G. bellidifolia not been distinguished. 58Á71 mm) than those of Sebaea (polar axis SUMMARY: Carmichaelia O1, I, P2, D1; Sophora 25Á29 mm). type O1, V, P2, D1. NAMES: Gentianella; Sebaea ovata. COMMENTS: There is considerable variation in surface pattern (Moar 1993; Glenny 2004) and Fagaceae for this reason most records are best referred to GENUS: Nothofagus (5). Gentianella; in subantarctic records those POLLEN: Grains easily recognized. Nothofagus from the Campbell Islands may be referred to menziesii pollen with7 short apertures is G. antarctica, from the Antipodes Islands to stephanocolpate, angulaperturate, apertures G. antipoda, and from the Auckland Islands not always easily resolved; the exine is very reference to G. cerina and G. concinna thin, and bears very short, more or less evenly may be made. Sebaea ovata grains are rarely distributed spinules; shape is peroblate. Grains recorded. of all other species are more robust, and SUMMARY (general): O1, I, P1, D1. smaller; apertures (5Á8) are clearly defined, short, and thickened around the margins; exine up to 1.5 mm thick, bearing spinules; shape Geraniaceae oblate, more or less circular in polar view. An GENERA: Geranium (8), Pelargonium (1). analysis of aperture distribution (Harris 1956a, POLLEN: Grains tricolporate; ectoapertures in b; Newnham 1992; Hanks & Fairbrothers 1976) Geranium are very short (approximatley one- shows 80% of all grains have either 6 or 7 apertures. N. fusca has approximately equal quarter length of the grain), contrasting with frequencies of both 6 or 7, whereas in the other the very long ectoapertures of Pelargonium; taxa,61Á66% have 7 apertures. This may be exine in both very thick, semitectate; Geranium useful to confirm the presence of N. fusca. exine bears blunt bold processes; Pelargonium NAMES: Fuscospora (subgenus includes N. fusca, exine bears fine spinules. N. solandri, N. cliffortioides, N. truncata); NAMES: Geranium; Pelargonium inodorum. Nothofagus menziesii. COMMENT: Geranium is the only pollen so far COMMENTS: Grains of the Fuscospora group recorded, and then only sparingly. (note: often mistakenly spelt Fuscaspora) are SUMMARY: Geranium O1, I, P1, D1. Standardizing names in New Zealand palynology 219

Gesneriaceae all species of Gonocarpus and in Haloragis GENUS: Rhabdothamnus (1). erecta. (b) Grains tricolpate to pentacolpate; POLLEN: Grains tricolporate, angulaperturate; exine up to 5 mm thick; tectum surface rough- ectoapertures very long, endoapertures lolon- ened. Sometimes occurs in Gonocarpus aggre- gate; exine finely reticulate; grains small. gatus, Haloragis erecta and Myriophyllum NAME: Rhabdothamnus solandri. robustum. (c) Grains triporate to hexaporate, COMMENT: Not recorded in pollen analyses. pores generally protruding; exine 1Á2 mm thick, SUMMARY: OX, V, P1, D1. surface smooth or verrucate. Characteristic of all species of Myriophyllum. Myriophyllum grains are either triporate (M. Goodeniaceae propinquum, M. votschii) or mainly tetraporate GENERA: Scaevola (1), Selliera (2). to hexaporate (M. pedunculatum, M. triphyl- POLLEN: Recognized by their coarse bacula lum). Two pollen types are produced by M. which are often bifurcate and longer at the robustum. The first is similar to those of M. polar area. Scaevola is larger and bolder than pedunculatum and the second similar to type (b) Selliera. noted above. NAMES: Scaevola gracilis; Selliera radicans. NAMES: Gonocarpus type (all Gonocarpus spp. COMMENT: Selliera is the only pollen type and Haloragis erecta); Haloragis type (pollen of recorded, and then only occasionally. Gonocarpus aggregatus, H. erecta and Myrio- SUMMARY: Selliera O1, I, P1, D1. phyllum robustum as described under type (b)); Myriophyllum type (all species of Myriophyl- lum). Griseliniaceae COMMENTS: Good indicators of wet habitats. It GENUS: Griselinia (2). has been suggested that the thick-walled grains POLLEN: Grains characterized by small lalon- noted under type (b) above may be unviable gate, slit-like endoapertures and faintly and (Moar 1993). irregularly striate exine. SUMMARY (general): O1, W, P2, D1. NAME: Griselinia. SUMMARY: O2, I, P1, D1. Hypericaceae Gunneraceae GENUS: Hypericum (4). POLLEN: Grains of medium size; ectoapertures GENUS: Gunnera (5). bordered by smooth margins; exine finely POLLEN: Grains tricolpate, markedly fossaper- reticulate. turate, finely reticulate. NAME: Hypericum. NAME: Gunnera. SUMMARY: O1, I, P2, D1. COMMENT: Often recorded in late-glacial sedi- ments. SUMMARY: O1, I, P2, D1. Ixerbaceae GENUS: Ixerba (1). Haloragaceae POLLEN: Grains tetraporate or pentaporate, GENERA: Gonocarpus (4), Haloragis (1), Myr- mostly the latter; angulaperturate; exine iophyllum (4). smooth, thinner along endoapertures to form POLLEN: Three pollen types are recognized: (a) a visible border. Grains tricolpate to pentacolpate; exine up to 2 NAME: Ixerba brexioides. mm thick; tectum generally smooth. Found in SUMMARY: O1, I, P1, D1. 220 NT Moar et al.

Lamiaceae optical section. Mitrasacme tricolporate; exine GENERA: Mentha (1), Scutellaria (1). finely reticulate. Grains of both taxa small. POLLEN: Mentha grains usually hexacolpate; NAMES: Geniostoma rupestre; Mitrasacme. apertures very long, sometimes syncolpate; COMMENTS: Geniostoma and Parsonsia (Apoc- exine finely reticulate. Scutellaria pollen tricol- ynaceae) grains are similar and may be con- pate; aperture membrane often ruptured in two fused. The smaller isopolar grains of or more places; exine reticulate. Geniostoma with their thick exine,1 mm, NAMES: Mentha cunninghamii; Scutellaria separate them from those of Parsonsia. Genios- novae-zelandiae. toma rupestre rarely recorded. Mitrasacme not COMMENTS: Mentha cunninghamii is occasion- recorded in pollen analyses. ally recorded; Scutellaria has not been recorded SUMMARY (general): O1, I, P1, D1. in pollen analyses. SUMMARY: Mentha cunninghamii O1, I, P1, D1. Loranthaceae GENERA: Alepis (1), Ileostylis (1) Peraxilla (2), Lentibulariaceae Trilepidea (1), Tupeia (1). GENUS: Utricularia (3). POLLEN: The small oblate or suboblate pollen of POLLEN: Grains tricolporate (Utricularia dichot- Ileostylis and Tupeia cannot be confused with oma) or pentacolporate (U. lateriflora); ectoa- the medium to large peroblate grains of Alepis, pertures in tricolporate grains very long, Peraxilla and Trilepidea. shorter in pentacolporate grains; endoapertures The spiny exine of Tupeia separates it from small and slit-like. Ileostylis of which the exine surface is faintly NAMES: Utricularia lateriflora; Utricularia di- rugulate or smooth. chotoma. The peroblate grains of Alepis, Peraxilla, and COMMENTS: Rarely, if ever recorded. Pollen of Trilepidea are variously striate, markedly trian- U. australis was not available for description. gular with often concave sides, and with long, SUMMARY (general): O1, I, P1, D1. usually syncolpate ectoapertures. NAMES: Ileostylis micranthus; Peraxilla type (Peraxilla spp., Alepis, Trilepidea); Tupeia ant- Linaceae arctica. GENUS: Linum (1). COMMENTS: Alepis, Peraxilla and Trilepidea POLLEN: Grains large (70Á75 mm); pericolpate pollen may be distinguished on the basis of (up to 12 apertures), very thick exine; baculi the exine pattern, but variation is considerable very short, of two sizes. often making this separation difficult. Peraxilla NAME: Linum monogynum. colensoi is probably most commonly encoun- COMMENT: Rarely recorded. tered in South Island material; Trilepidea SUMMARY: O1, W, P2, D1. adamsii, now extinct, was restricted to northern districts of the North Island. Peraxilla type and Tupeia antarctica are the most commonly Loganiaceae recorded pollen, but not in great numbers. GENERA: Geniostoma (1), Mitrasacme (2). SUMMARY: Ileostylis micranthus O1, I, P1, D1; POLLEN: Grains of Geniostoma triporate; ex- Peraxilla type O2, V, P2, D1; Tupeia antarctica ine2 mm around pore; surface smooth in O1, I, P1, D1. Standardizing names in New Zealand palynology 221

Malvaceae Moraceae GENERA: Hibiscus (2), Hoheria (7), Plagianthus GENUS: Streblus (3). (2). POLLEN: Grains diporate; exine very thin, POLLEN: Grains medium (Hoheria, Plagianthus) slightly thickened at pore margins. to very large (Hibiscus); tricolporateÁ NAME: Streblus. pentacolporate (Hoheria, mainly tetracolporate; SUMMARY: O1, W, P2, D2. agianthus, mainly pentacolporate) or periporate (Hibiscus); spinulose, spines35 mm long in Hibiscus,upto5mm long in Hoheria, and rarely Myoporaceae (Scrophulariaceae) 2 mm long in Plagianthus. GENUS: Myoporum (1). NAMES: Hibiscus; Hoheria; Plagianthus. POLLEN: Grains tricolpate; colpi characterized COMMENT: Hoheria angustifolia sometimes has by up to two lalongate fissures; exine reticulate. narrow, needle-shaped spines, e.g. from Peel NAME: Myoporum. Forest, Canterbury (Moar 1993), and such COMMENTS: Myoporum laetum grains are larger grains could be referred to H. angustifolia in (polar axis 35Á40 mm) than those of M. debile any discussion. Although spine length may vary (polar axis 26Á30 mm), which apparently is now Hoheria considerably, it is possible to separate extinct (Webb et al. 1988); apertures are similar Plagianthus and grains. Spines and ectexine are to those of Tecomanthe speciosa (Bignonia- sometimes stripped from the grains; these are ceae). Grains of Myoporum are occasionally Hoheria Plagianthus referable to or on the basis recorded. of aperture number. Pollen of the northern SUMMARY: O1, I, P2, D1. Hibiscus cannot be confused with other pollen types. SUMMARY: Hibiscus OX, I, P2, D1; Hoheria O2, I, P2, D2; Plagianthus O2, IW, P2, D2. Myrsinaceae (Primulaceae) GENERA: Elingamita (1), Myrsine (11). POLLEN: Grains of Myrsine have short aper- Meliaceae tures, usually four, and a smooth exine. Elin- GENUS: Dysoxylum (1). gamita johnsonii pollen is tricolporate and POLLEN: Grains isopolar or subisopolar, usually faintly reticulate. tetracolporate; ectoapertures very short. NAMES: Elingamita johnsonii; Myrsine. NAME: Dysoxylum spectabile. COMMENTS: Myrsine pollen is often well repre- COMMENT: Rarely recorded. Pollen dispersal sented in late-glacial sediments and in mainland may be restricted because the inflorescence is sites may include either the prostrate M. cauliflorous. nummularia or the shrub M. divaricata;in SUMMARY: O1, I, P1, D1. post-glacial sites any of the mainland tree or shrub taxa may be represented. In the suban- tarctic Auckland and Campbell islands the Menyanthaceae source taxon is M. divaricata; in Chatham GENUS: Liparophyllum (1). Island sites the source taxa may be either M. POLLEN: Grains peroblate, triangular in polar chathamica or M. coxii. Although there are view, tectum generally boldly rugulate or rugu- subtle differences, pollen of the different taxa lo-striate; ectoapertures parasyncolpate. cannot easily be distinguished. Pollen of Elin- NAME: Liparophyllum gunnii. gamita johnsonii has not been recorded in COMMENT: The bold exine pattern clearly pollen analyses. separates these grains from those of Myrtaceae. SUMMARY: Elingamita johnsonii OX, P1, D1; SUMMARY: O1, I, P1, D1. Myrsine O4, I/W?, P3, D3. 222 NT Moar et al.

Myrtaceae Metrosideros robusta type (M. excelsa and M. GENERA: Kunzea (2), Leptospermum (1), Lopho- robusta) O3, I (V), P3, D2; Neomyrtus type myrtus (2), Metrosideros (12), Neomyrtus (1), (Lophomyrtus, Neomyrtus) O1, I, P3, D1; Syzygium (1). Syzygium O1, I, P2 (3), D1. POLLEN: Grains peroblate, tricolporate, usually parasyncolpate except for Lophomyrtus and Neomyrtus; angulaperturate; angles more or Nyctaginaceae less rounded except in the acutely angled GENUS: Pisonia (1). Syzygium; exine surface usually smooth or POLLEN: Grains pericolpate, 12Á15 aperturate; faintly marked, but clearly verrucate in Lopho- tectum often minutely spinulose, perforate. myrtus and Neomyrtus; Leptospermum and NAME: Pisonia brunoniana. Kunzea have the smallest grains at15Á20 mm COMMENTS: Grains not recorded in pollen in equatorial plane, all others24 mmor analyses. Superficially similar to Montia but greater. Metrosideros excelsa and M. robusta distinguished by much shorter and broader grains tend to be larger and more robust than apertures. those of other Metrosideros species. SUMMARY: OX, I?, P1, D1. NAMES: Leptospermum type (Kunzea, Leptos- permum); Metrosideros, Metrosideros robusta type; Neomyrtus type (Lophomyrtus, Neomyr- Oleaceae tus); Syzygium maire. GENUS: Nestegis (4). COMMENTS: Because Kunzea was formerly in- POLLEN: Grains tricolpate; apertures short, cluded in Leptospermum, and the pollen grains recessed, exine reticulate. of the two genera are indistinguishable, the NAME: Nestegis. latter name is chosen for the pollen type. COMMENTS: Grains regularly recorded in North Lophomyrtus pollen is sometimes more boldly Island samples; in the South Island mostly in verrucate than that of Neomyrtus, but because interglacial samples. The grains have been the distinction is often not clear, the pollen type confused with those of Brassicaceae and Calli- is named after the more widely distributed triche. Neomyrtus. Of the Metrosideros species repre- SUMMARY: O2, I, P2, D2. sented in New Zealand, M. excelsa, M. robusta, and M. umbellata are most likely to be encoun- tered during a pollen analysis. Occasionally a Onagraceae larger and more robust grain is identified as M. GENERA: Epilobium (30), Fuchsia (3). robusta type, especially in North Island sites. POLLEN: Grains with protruding compound When southern sites are being studied, as in apertures; thick spongy tectum; viscin threads Westland, the probability that the pollen is often present. Epilobium pollen usually shed as derived from M. umbellata, especially if it is loosely aggregated tetrads; individual grains abundant, should be noted in the text. In tricolporate; Fuchsia always monads, dicolpo- material from the Auckland Islands grains rate. could be identified as M. umbellata, although NAMES: Epilobium; Fuchsia. if grains are infrequent the possibility of long- COMMENT: The differences between pollen of distance transport, including Eucalyptus from species in both genera are too slight to allow Australia, must be considered. greater precision of identification. SUMMARY: Leptospermum type (Kunzea, Lep- SUMMARY: Epilobium O1, I, P1, D1; Fuchsia tospermum) O4, I, P3, D1; Metrosideros or O1, V, P1, D1. Standardizing names in New Zealand palynology 223

Oxalidaceae POLLEN: Grains periporate, pores up to seven, GENUS: Oxalis (4). small; tectum verrucate; shape spheroidal. POLLEN: Grains tricolpate; finely reticulate; NAME: Plantago. tectum in O. magellanica bears unevenly spaced COMMENT: Mostly recorded in glacial or late- spinules. glacial sediments. NAME: Oxalis. SUMMARY: O1, W, P2, D2. COMMENT: Not recorded in pollen analyses. SUMMARY: OX, I, P1, D1. Polygonaceae GENERA: Muehlenbeckia (5), Persicaria (1), Passifloraceae Rumex (2). GENUS: Passiflora (1). POLLEN: Grains of the three genera are easily POLLEN: Grains, large, hexacolpate, colpi in distinguished. Muehlenbeckia and Rumex three pairs; exine finely reticulate. grains are tricolporate, but the thicker exine NAME: Passiflora tetrandra. of Muehlenbeckia (2 mm) and its striateÁ SUMMARY: O1, V, P2, D1. reticulate pattern separates it from the thinner, faintly reticulate grains of Rumex. Persicaria Pennantiaceae grains are periporate, the exine is thick and the GENUS: Pennantia (2). pores deeply recessed. POLLEN: Grains tricolporate, often fossapertu- NAMES: Muehlenbeckia; Persicaria decipiens; rate in polar view; endoapertures short, slit- Rumex. like, exine surface generally smooth, but some- SUMMARY: Muehlenbeckia O1, I, P3, D1; Persi- times roughened. caria decipiens O1, I, P1, D1; Rumex O1, W, NAME: Pennantia. P2, D1. COMMENTS: May be confused with Griselinia but distinguished by the smooth or roughened exine surface in contrast to the finely striate Portulacaceae surface of Griselinia; rarely recorded. GENERA: Hectorella (1); Montia (8). SUMMARY: O1, I, P2, D1. POLLEN: Hectorella grains tricolpate, colpi broad, very long; tectum bearing short scat- tered spinules and perforate. Montia grains Pittosporaceae pericolpate, colpi broad, 12Á30 in number; GENUS: Pittosporum (22). tectum bearing widely spaced minute spinules, POLLEN: Grains tricolporate; ectoapertures very and perforate. long; exine smooth; tectum with scattered NAMES: Hectorella caespitosa; Montia. perforations. Exine reticulate in P. dallii; un- COMMENT: Pollen grains of all species of Montia even and undulate in P. pimeleoides. are similar and variable colpi numbers make NAME: Pittosporum. separation difficult; usually recorded in sub- COMMENTS: Not recorded in great numbers; neither P. dallii nor P. pimeleoides recorded in alpine or late-glacial sediments. There are pollen analyses. similarities with Pisonia pollen, but confusion SUMMARY: O1, I, P1, D1. is unlikely because of the marked differences in distribution and plant communities. Hectorella pollen not recorded in pollen analyses. Plantaginaceae SUMMARY: Hectorella caespitosa OX, I, P1, D1; GENUS: Plantago (9). Montia O1, I, P2, D1. 224 NT Moar et al.

Primulaceae Ranunculus pollen see Moar (1993). The tricol- GENUS: Samolus (1). pate grains of Ceratocephalus, Myosurus and POLLEN: Grains small, finely reticulate, lumina Psychrophila bear very short, more or less larger in mesocolpia. evenly distributed spinules, and cannot easily NAME: Samolus repens. be distinguished. COMMENT: Rarely recorded. NAMES: Psychrophila type (Ceratocephalus, SUMMARY: O1, I, P1, D1. Myosurus, Psychrophila); Ranunculus type (An- one, Clematis, e.g. R. grahamii, R. biternatus); Ranunculus carsei type (e.g. R. carsei, R. Proteaceae cheesemanii); Ranunculus nivicola type (e.g. R. GENERA: Knightia (1), Toronia (1). nivicola, R. insiginis). POLLEN: Grains triporate, angulaperturate; COMMENTS: Variation within and between the Knightia finely reticulate, Toronia exine perfo- genera make precise identification difficult and rate, with scattered gemmae. using the name Ranunculaceae would be ac- NAMES: Knightia excelsa; Toronia toru. ceptable for most situations. Pollen is mostly COMMENT: Knightia is the most frequently recorded in late-glacial sediment, but is never recorded, although never in great numbers. recorded in great numbers. SUMMARY: Knightia excelsa O1, V, P2, D1; SUMMARY (general): O1, I, P2, D1. Toronia toru O1, V, P1, D1.

Rhamnaceae Quintiniaceae (Paracryphiaceae) GENERA: Discaria (1), Pomaderris (7). GENUS: Quintinia (3). POLLEN: Grains of Discaria are generally tri- POLLEN: Grains small, tetracolporate or penta- colporate, angulaperturate; ectoapertures very colporate, mostly the latter, ectoapertures very long; endoapertures small, more or less circu- long, endoapertures small, elliptical; exine very lar; exine smooth or faintly verrucate or thin, smooth. scabrate. Pomaderris kumeraho grains are simi- NAME: Quintinia. lar, but ectoapertures are obviously costate and COMMENT: Generally only occasionally re- endoapertures are H-shaped. The pollen grains corded, but sometimes abundant. of other Pomaderris species are characterized SUMMARY: O1, I, P1, D2. by as many as eight, often distorted, ectoaper- tures. NAMES: Discaria toumatou; Pomaderris. Ranunculaceae COMMENTS: Although it may be difficult to GENERA: Anemone (1), Ceratocephalus (1), separate Pomaderris kumeraho from Discaria Clematis (10), Myosurus (1), Psychrophila (2), toumatou when viewed in polar view, in practice Ranunculus (45). this is not a major difficulty because Pomader- POLLEN: Grains of Ceratocephalus, Clematis, ris is mostly restricted to northern areas of the Myosurus, Psychrophila and many species of North Island. Pomaderris phylicifolia extends Ranunculus, e.g. R. grahami, are tricolpate; southwards into Canterbury, but only in scat- other species of Ranunculus are pericolpate, tered localities. Pomaderris pollen from e.g. R. nivicola. Anemone grains are tricolpate Australia may occur in surface samples, espe- or hexacolpate. Exine characters are variously cially on the western slopes of the Southern spinulose, perforate and, in the case of Ranun- Alps. Both are generally only rarely recorded, culus the surface is either slightly undulate, e.g. but Discaria is sometimes frequent, e.g. Cowan R. nivicola, or obviously undulate, e.g. R. & McGlone (1991). carsei. For discussion on the differences in SUMMARY (general): O1, I, P2, D1. Standardizing names in New Zealand palynology 225

Rosaceae short and narrow; endoapertures are broad GENERA: Acaena (18), Geum (7), Potentilla (1), and often long; endocracks are common, and Rubus (5). in Nertera often numerous and anastomosing; POLLEN: Grains mostly tricolporate, but Geum in Nertera the exine is often gemmate. pollen is tetracolporate or pentacolporate ex- NAMES: Coprosma; Galium; Nertera. cept for the tricolporate G. uniflorum. Acaena COMMENTS: Sometimes the characteristic gem- and Potentilla have a well-defined operculum mae of Nertera pollen are lost; identification over each ectoaperture, but are easily distin- may be possible if endocracks are clearly guished by differences in exine pattern Á visible. Tetracolporate grains, not often re- rugulate in Acaena, striate in Potentilla. Rubus corded, may reflect the presence of Coprosma pollen is striate, but apertures lack opercula, as perpusilla, especially in material from the sub- do Geum pollen, which are often sub-rectangu- antarctic islands. lar or square in polar view and possess rugulate SUMMARY: Coprosma O3, W, P3, D3; Galium or spinulose exine. O1, I, P1, D1; Nertera O1, W, P3, D2. COMMENT: Grains most frequently recorded are those of Acaena and Rubus. NAMES: Acaena; Geum; Potentilla anserinoides; Rutaceae Rubus. GENERA: Melicope (2), Phebalium (1). SUMMARY: Acaena O1, WÁI?, P2, D1; Geum POLLEN: Grains tricolporate, reticulate; Pheba- O1, I, P1, D1; Potentilla O1, I, P2, D1; Rubus lium is considerably smaller than Melicope, has O1, I, P3, D1. a finer reticulum and sometimes an H-shaped endoaperture is obvious. NAMES: Phebalium nudum; Melicope. Rousseaceae SUMMARY (general): O1, I, P2, D1. GENUS: Carpodetus (1). POLLEN: Grains tetrahedral tetrads; ectoaper- tures and endoapertures of about equal length; Santalaceae exine often faintly rugulose. Key character GENERA: Exocarpus (1), Mida (1). separating Carpodetus from other tetrad pollen POLLEN: Grains of Exocarpus are characterized is the presence of ectoapertures on the internal by short ectoapertures and by the fusion of walls. endoapertures to form a continuous equatorial NAME: Carpodetus serratus. ring. In contrast, Mida pollen is heteropolar, COMMENT: Often recorded in low numbers. and ectoapertures are parasyncolpate at one SUMMARY: O1, I, P1, D1. pole, bifurcate at the other. NAMES: Exocarpus bidwillii; Mida salicifolia. COMMENTS: Although the fused endoapertures Rubiaceae of Exocarpus are similar to those of Stilbocarpa GENERA: Coprosma (60), Galium (3), Nertera (Araliaceae) the smaller grains of Exocarpus (8). (polar axis 21Á23 mm), and their shorter POLLEN: Grains of Galium are small, pentacol- ectoapertures, separate them from those of pate or hexacolpate, and minutely spinulose. Stilbocarpa (polar axis 31Á36 mm). Pollen not Coprosma and Nertera grains are usually tri- recorded in pollen analyses. colporate, but some, especially Coprosma per- SUMMARY: Exocarpus bidwillii OX, I, P1, D1; pusilla, are tetracolporate; ectoapertures are Mida salicifolia OX, I, P1, D1. 226 NT Moar et al.

Sapindaceae varying degrees, a characteristic that separates GENERA: Alectryon (1), Dodonaea (1). them from other genera in the family. These POLLEN: Grains of Alectryon are characterized five genera are considered part of Veronica s.l. by a finely striateÁreticulate exine, in contrast by other authors. to the scabrate exine of Dodonaea, which Euphrasia grains are larger, at the upper end of sometimes exhibits parallel folds in the meso- the medium size range, mostly tricolpate; colpia, and slightly protruding lalongate en- colpus membrane often ruptured; tectum min- doapertures. utely reticulate. Mimulus and Limosella grains NAMES: Alectryon excelsus; Dodonaea viscosa. are small to medium in size, semitectate and SUMMARY: Alectryon excelsus O1, I, P2, D1; reticulate. Dodonaea viscosa O1, W, P3, D2. Grains of Mazus and Ourisia are close, but the former are larger, up to 42 mm, in contrast to the 25Á30 mm for Ourisia. Both are clearly Sapotaceae reticulate with Mazus generally having the GENUS: Pouteria (1). coarser reticulum. POLLEN: Grains mainly rectangular; tetracolpo- NAME: Euphrasia; Glossostigma; Gratiola type rate; flattened at pole and nearly square in (Gratiola, Jovellana); Hebe type (Chionohebe, polar view. Hebe, Heliohebe, Leonohebe, Parahebe); Limo- NAME: Pouteria costata. sella type (Limosella, Mimulus); Ourisia type SUMMARY: OX P2, P1, D1. (Mazus, Ourisia). COMMENTS: Of these pollen grains those of Euphrasia and the Hebe type are the most Scrophulariaceae frequently recorded; the others rarely so or GENERA: All genera here are included in Scro- not at all. Euphrasia is occasionally recorded in phulariaceae, but alternative families as indi- high numbers, e.g. Dickson (1972); those of the cated by recent phylogenetic studies are also Hebe type always in such low numbers that shown in parentheses. Chionohebe (5), Gratiola more precise identification would be difficult. (3), Hebe (c. 80), Heliohebe (6), Leonohebe (5), SUMMARY: Euphrasia O1, I, P2, D1; Glossos- Ourisia (13), Parahebe (17), (Plantaginaceae); tigma, Gratiola type; Limosella type O1, I, P1, Euphrasia (15) (Orobanchaceae); Jovellana (2) D1; Hebe type O2, I, P3, D2; Ourisia type OX, (Calceolariaceae); Glossostigma (2), Mazus (2), I, P2, D1. Mimulus (1) (Phrymaceae); Limosella (2) (Scro- phulariaceae). POLLEN: Grains mostly variously reticulate and Solanaceae colpate. Exceptions are Glossostigma, Gratiola GENUS: Solanum (3). and Jovellana. Glossostigma grains are minutely POLLEN: Grains tricolporate (Solanum ameri- spinulose and perforate, mostly tetracolpate, canum, S. aviculare) or mostly tetracolporate and in the case of G. elatinoides the polar exine (S. laciniatum); vestibulate. is markedly thickened. Gratiola and Jovellana NAME: Solanum. pollen is colporate; Gratiola nana is usually COMMENT: Grains not recorded in pollen ana- tricolporate, but may be tetracolporate, in lyses. contrast to G. sexdentata which is usually SUMMARY: OX, I, P2, D1. tetracolporate, but may be tricolporate. Jovel- lana is mostly tricolporate. Pollen grains of Hebe, Heliohebe, Chionohebe, Stylidiaceae Leonohebe, and Parahebe are very similar Á all GENERA: Forstera (5), Oreostylidium (1), Phyl- are finely reticulate or reticulateÁstriate in lachne (3). Standardizing names in New Zealand palynology 227

POLLEN: Pollen of Forstera, Oreostylidium and POLLEN: Grains mostly triporate, but Elatoste- Phyllachne are spinulose; in all taxa the spinules ma grains are diporate, and Parietaria grains are usually very short and best resolved by LO are mostly tetraporate. analysis (Moore et al. 1991); grains may be NAME: Urtica type (all taxa). tricolpate as in Forstera sedifolia, tricolpa- COMMENT: Urtica grains are most likely to be teÁpentacolpate as in Phyllachne species, or recorded, but only rarely. tetracolpateÁpentacolpate as in Oreostylidium SUMMARY: Urtica O1, W, P2, D1. subulatum, which is distinguished by a perforate exine. NAMES: Phyllachne type (Phyllachne, Forstera); Verbenaceae Oreostylidium subulatum. GENERA: Teucridium (1), Vitex (1). SUMMARY (general): O1, I, P1, D1. POLLEN: Grains tricolpate; Teucridium parvifo- lium characterized by a finely verrucate tectum and well-developed operculum, which may be Tetrachondraceae destroyed by processing; Vitex by a smooth, GENUS: Tetrachondra (1). perforate, sometimes faintly striate tectum. POLLEN: Grains tricolporate, small; tectum NAMES: Teucridium parvifolium; Vitex lucens. smooth, but rugulate in mesocolpium, perfo- COMMENTS: Teucridium not recorded; cannot be rate elsewhere. confused with Rosaceae pollen. NAME: Tetrachondra hamiltonii. SUMMARY: Teucridium parvifolium OX, W, P1, SUMMARY: OX, W?, P1, D1. D1; Vitex lucens O1, W, P1, D1.

Thymelaeaceae Violaceae GENERA: Kelleria (8), Pimelea (29). GENERA: Melicytus (10), Viola (3). POLLEN: Grains display croton pattern. POLLEN: Grains generally similar except for the NAME: Pimelea type. larger size of Viola, polar axis up to 30 mm long, COMMENTS: There is some variation between in contrast to the more usual 21 mm for taxa; numbers of grains recorded are few and Melicytus. In both genera, the tectum appears usually only from sites reflecting cool condi- smooth, or nearly so in Bright Field (BF), but tions. may be finely reticulate (Melicytus ramiflorus), SUMMARY: O1, I, P2, D1. or rugulate; perforate (M. crassifolius) under scanning electron microscopy. NAMES: Melicytus; Viola. Tiliaceae (Malvaceae) COMMENT: Melicytus is more frequently re- GENUS: Entelea (1). corded than Viola, but neither is common. POLLEN: Surface exine either with supratectal SUMMARY (general): O1, I, P1, D1. reticulum or rugulate; tectum perforate. NAME: Entelea arborescens. SUMMARY: O1, I, P1, D1. Viscaceae (Santalaceae) GENUS: Korthalsella (3). POLLEN: Grains small, exine in mesocolpium Urticaceae much thicker than polar exine. GENERA: Australina (1), Boehmeria (1), Elatos- NAME: Korthalsella. tema (1), Parietaria (1), Urtica (6). SUMMARY: O1, I, P1, D1. 228 NT Moar et al.

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