| HOME | HELP | FEEDBACK | SUBSCRIPTIONS | ARCHIVE | SEARCH | TABLE OF CONTENTS |
|
|
|
|||||||
|
|||||||||
Paleobotany |
School of Plant Science, University of Tasmania, Private Bag 55, Hobart 7001, Tasmania, Australia
Received for publication August 1, 2002. Accepted for publication November 21, 2002.
 |
ABSTRACT |
|---|
 TOP ABSTRACT INTRODUCTIONMATERIALS AND METHODSRESULTSSYSTEMATIC DESCRIPTIONDISCUSSIONLITERATURE CITED |
|---|
Key Words: Antarctica • Australia • Caryophyllaceae • Centrospermae • Eocene • fossil • Paleogene • phylogeny • pollen
|
INTRODUCTION |
|---|
|
TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSSYSTEMATIC DESCRIPTIONDISCUSSIONLITERATURE CITED |
|---|
). Only three of the genera are large shrubs or small trees, Sanctambrosia, endemic to San Ambrosio Island, Chile, and the Hawaiian endemics, Alsinidendron and Schiedea (Bittrich, 1993
). The family is included in the order Caryophyllales, which Cronquist (1981)
defined to contain about 600 genera and 10 000 species. This is a monophyletic group (Cuenod et al., 2002
). Cactaceae, Aizoaceae, Amaranthaceae, and Portulacaceae are other large families in the same order. Savolainen et al. (2000)
defined Caryophyllales more broadly but this work will use Cronquist's (1981)
narrow definition.
The fossil record of the family is sparse, with published pre-Pleistocene records consisting almost exclusively of pollen. Most Caryophyllaceae have more or less spherical, periporate, spinulose, microechinate or microreticulate annular perforate or punctate pollen with simple apertures (Nowicke, 1994
; Punt and Höen, 1995
). Fossil pollen matching this description are included in Periporopollenites Pflug & Thompson in Thompson & Pflug 1953, Malvacipollis and Chenopodipollis Krutzsch 1966. Periporopollenites includes pollen previously assigned to Periporites Van der Hammen 1956, Caryophyllidites Couper 1960, Parsonsidites Couper 1960, Liquidambarpollenites Ratz 1937 ex Potonié 1960, Polyporina Naumova 1939 ex Potonié 1960, Caryophyllaceaepites Biswas 1962, and Chenopodipollis. Malvacipollis and Chenopodipollis are still in use, but now only encompasses pollen of Euphorbiaceae and Amaranthaceae (defined to include Chenopodiaceae), respectively (Truswell et al., 1985
).
Periporopollenites polyoratus (Couper 1960) Stover and Partridge 1973
has the longest geological range of fossil pollen types assigned to Caryophyllaceae. Its earliest record is Late Campanian from the Gippsland and Otway Basins in southeast Australia (Stover and Partridge, 1973
) and South Island, New Zealand (Raine, 1984
). Its last recorded appearance is Oligocene. Younger records of Periporopollenites that may be derived from members of Caryophyllaceae are morphologically distinct species. They include Oligocene specimens of P. pallidus and P. vesicus (Nott and Owen, 1992
), Early Miocene specimens of P. vesicus and two undescribed Periporopollenites species (Owen, 1988
), and an undescribed Early-?Middle Miocene grain (Tulip et al., 1982
).
Caryophyllaceae-type pollen first appears in Europe in the Miocene (Muller, 1981
). This late appearance is surprising considering the much higher diversity of the family in the Northern Hemisphere (Bittrich, 1993
) and the earlier Southern Hemisphere records. Eocene seeds from England, Hantsia pulchra and H. glabra (Chandler, 1961
, 1963
), have been attributed to the family. These are the only macrofossil records of the family, but their taxonomic status is uncertain (Chandler, 1961
).
Morphological convergence means that the pollen records of Caryophyllaceae need to be interpreted carefully. All the features of typical Caryophyllaceae pollen occur in other families of the order, notably Amaranthaceae, Rivinaceae, and Nyctaginaceae. Periporate pollen also occurs in unrelated families such as Papaveraceae, Convolvulaceae, Euphorbiaceae, and Hamamelidaceae.
This work describes a fossil inflorescence from Middle-Late Eocene sediments at Locharbour, northwestern Tasmania, Australia, and the relationships of the in situ pollen with dispersed fossil pollen types. Recent progress in understanding angiosperm evolution has been made by determining the relationships of extinct species to living taxa without forcing fossils into extant genera or families (Dilcher, 2000
). This work therefore uses a phylogenetic analysis to determine the relationships of the fossil to extant groups. Formal phylogenetic analysis was necessary because of extensive morphological homoplasy within the order.
|
MATERIALS AND METHODS |
|---|
|
TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSSYSTEMATIC DESCRIPTIONDISCUSSIONLITERATURE CITED |
|---|
). The associated palynoflora falls in the Lower Nothofagidites asperus zone of Stover and Partridge (1973)
, implying a Middle or possibly Late Eocene age.
The living relatives of fossil species at Locharbour and coeval sites suggest that floristically the vegetation at the time of deposition was diverse, cool climate rainforest dominated by genera with extant species in temperate and tropic-montane rainforests (Carpenter et al., 1994
; Macphail et al., 1994
). The Locharbour fossil flora includes the conifers Phyllocladus aberensis (Hill, 1989
), Willungia oppositifolia (Hill and Pole, 1992
), Acmopyle tasmanica, Dacrycarpus mucronatus (Hill and Carpenter, 1991
), and Araucaria (Carpenter et al., 1994
), and the angiosperms Eucryphia aberensis (Barnes and Jordan, 2000
) and Banksieaephyllum attenuatum (Hill and Christophel, 1988
).
Specimen preparation
One partial inflorescence was found by manually splitting a block of sediment. Brushing the block with a fine brush under running water exposed this fossil progressively. Some individual anthers were squashed on glass slides and mounted in phenol glycerine jelly for light microscopy. Others were soaked in 50% hydrofluoric acid for approximately 1 h to remove residual sediments, then placed on double-sided conductive tape on aluminum stubs. These were sputter coated with gold to approximately 40 nm thick and observed under high vacuum with an ESEM 2020 Environmental Scanning Electron Microscope (ElectroScan, Wilmington, Massachusetts, USA).
Data for cladistic analyses
This work seeks the most parsimonious phylogeny of extant species and the fossil (e.g., Jordan and Hill, 2000
). Molecular and morphological data were used for extant species and only morphological data for the fossil. Eighty extant genera with published rbcL or matK nucleotide sequences were used (Appendix 1, archived at the Botanical Society of America website [http://ajbsupp.botany.org/V90/]). They were 75 genera of Caryophyllales and genera from five families chosen as outgroups based on Savolainen et al. (2000)
and Cuenod et al. (2002)
. All recognized families of the order were represented.
In total 41 rbcL and 69 matK sequences were used, with both rbcL and matK sequences for 30 genera. For 76 genera only one species was used. In 14 genera the rbcL and matK sequences were from different species, and the sequences were combined to minimize the sparsity of the data matrices. This approach could cause misinterpretation if the relevant pairs of species were not monophyletic with regard to the other species analyzed. However, the sparse sampling of taxa makes this unlikely. Scoring of morphological characters allowed for polymorphism in these genera.
After alignment the rbcL sequences were trimmed to 1389 sites. Two sequences for Halophytum ameghinoi were combined to produce a usable length of 1187 sites. The matK alignments followed Cuenod et al. (2002)
. Ten informative presence/absence characters from rbcL sequences and five from matK sequences were created for indels. The missing nucleotides were treated as missing values (Simmons and Ochoterena, 2000
).
Most morphological/anatomical characters (Table 1) were derived from Rodman's (1994)
characters, but were scored following an exemplar approach (Prendini, 2001
) rather than inferring basal states for families. Preference was given to characters that could be scored unambiguously on the fossil. The data included 23 characters. Twenty morphological characters were compiled from Ross-Craig (1951)
, Roles (1957)
, Davis (1967)
, Dyer (1975)
, Nowicke and Skvarla (1979)
, Manly and Hutchins (1980)
, Cronquist (1981)
, Correa (1984)
, Eliasson (1993)
, Kubitzki et al. (1993)
, Nowicke (1994
, 1996)
, Hofmann (1994)
, and Punt and Höen (1995)
. Of these, 19 could be scored on the fossil. One character was biochemical (pigment type; scored from Clements et al. [1994
] and Cuenod et al. [2002]
). Two characters described sieve element plastids (Behnke, 1994
).
|
) was used to seek most-parsimonious trees. All characters were analyzed both for extant species only and for extant species and the fossil (Appendix 2, archived at the Botanical Society of America website [http://ajbsupp.botany.org/V90/]). These analyses used random addition sequences replicated 100 times, with tree bisection-reconnection, saving multiple trees. Branch support for the analysis of extant species was assessed by bootstrap with 1000 replicate heuristic searches, each using simple addition sequence, saving a single tree at each step and, because of memory limitations, restricting each replicate to 108 rearrangements.
The power of phylogenetic placements using the limited data available on the fossil was tested following Jordan and Hill (2000)
. For each extant species, a matrix (Appendix 3, archived at the Botanical Society of America website [http://ajbsupp.botany.org/V90/]) was created. The relevant species was coded only for characters that could be scored on the fossil (thus simulating a fossil). All characters were coded for the other 79 extant species. The most-parsimonious phylogenetic placement of each extant species using this restricted data was then compared with its placement based on the complete data.
|
RESULTS |
|---|
|
TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSSYSTEMATIC DESCRIPTIONDISCUSSIONLITERATURE CITED |
|---|
|
). The placement within Caryophyllaceae is better supported: the shortest trees placing the fossil outside the family were three steps longer. These placed the fossil in a number of basal positions with the Caryophyllales.
|
|
SYSTEMATIC DESCRIPTION |
|---|
|
TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSSYSTEMATIC DESCRIPTIONDISCUSSIONLITERATURE CITED |
|---|
Order
Caryophyllales.
Family
Caryophyllaceae.
Genus
Caryophylloflora G. J. Jord. & Macphail genus nov.
Etymology
In reference to the association with Caryophyllaceae.
Diagnosis
Inflorescence dichasial, bracts fused around each node. Flowers with several bracts appressed to the calyx. Calyx a five-lobed tube. Stamens free, five opposite and five alternate to the sepals. Ovary syncarpous, superior. Pollen a sphaeroidal, periporate monad with nonannulate, circular pores; exine perforate, tectate with clearly visible columellae.
Species
Caryophylloflora paleogenica G. J. Jord. & Macphail sp. nov. (Figs. 3–22).
|
Locality
Locharbour mine, northeastern Tasmania, Australia 40°56'16'' S, 148°0'34'' E.
Age
Middle to Late Eocene.
Etymology
In reference to the Paleogene age of the fossil.
Diagnosis
Inflorescence an open dichasium with at least three flowers. Flowers hermaphrodite or unisexual. Pedicels 5–10 mm long, subtended by acute bracts, pairs of bracts fused around the peduncle. Floral bracts acute, imbricate, about as long as the calyx tube. Calyx tube approximately 3 mm long, lobes triangular, obtuse, approximately two-thirds as long as the tube. Stamens slightly exserted, filaments tapering slightly towards the apex. Anthers approximately 1 mm long, dorsifixed, tetrasporangiate, opening latrorsely by longitudinal slits. Ovary opening by five valves. Pollen 24–(30)–32 µm in diameter; amb circular; pores approximately 16, 3–3.5 µm in diameter, with or without a pore membrane; exine 2–2.5 µm thick, sexine approximately 1.5 µm thick, microreticulate-microechinate, thinner around pores, nexine <<1 µm thick, protruding through or invaginated within pores.
|
DISCUSSION |
|---|
|
TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSSYSTEMATIC DESCRIPTIONDISCUSSIONLITERATURE CITED |
|---|
. The four major clades, Caryophyllales, higher Caryophyllales, Caryophyllaceae, and higher Caryophyllaceae (Fig. 1), were also strongly supported by earlier analyses (Downie and Palmer, 1994
; Manhart and Rettig, 1994
; Savolainen et al., 2000
; Cuenod et al., 2002
; Smissen et al., 2002
). In the most parsimonious trees (Fig. 1), all morphological characters are homoplasious, including some traditionally considered to be of taxonomic significance. Pollen ornamentation, aperture number and shape, stamen number and arrangement, perianth merosity and fusion all appear to have converged or reverted in several clades. For example, tricolpate and periporate pollen occur in both Caryophyllaceae and higher Caryophyllales. The strong support for these clades implies convergence in aperture shape and number.
Phylogenetic placement of the fossil
The analyses of the power to correctly classify fossils suggest that, given the characters available on the fossil, the approach used here would place only members of higher Caryophyllaceae within that clade. Combined with the additional numbers of steps required to place the fossil outside the clade, this makes the placement of C. paleogenica as a member of higher Caryophyllaceae (Fig. 2) convincing, in spite of extensive homoplasy. These interpretations assume that the fossil is a member of the order. However, the combination of periporate pollen, cymose inflorescence, fused calyx, and pentamerous flowers, with five stamens opposite the sepals and five alternate, makes placement in other groups unlikely.
The pollen grains of C. paleogenica (Figs. 9–22) are consistent with dispersed grains of Periporopollenites polyoratus from Australia and East Antarctica (Figs. 23–33). The dispersed grains are larger (39 µm vs. 30 µm), but this is probably due to oxidization from weathering and palynological preparation. The grains of C. paleogenica differ from other Periporopollenites species described in Australia. They have larger and more pores than P. demarcatus Stover & Partridge 1973
(Figs. 34–41) and P. hexaporus Macphail & Hill 1994
(Figs. 43–45). They are smaller than P. vesicus Stover & Partridge 1973
, which does not have distinctly defined columellae. Periporopollenites pallidus Truswell & Owen 1988 has more pores (20–30) and a thinner exine (1–1.3 µm) that does not decrease in thickness around the pores.
|
|
; Nowicke and Skvarla, 1979
; Moar, 1993
; Nowicke, 1994
, 1996
; Punt and Höen, 1995
). Most extant Caryophyllaceae (e.g., Figs. 46–53) have (1) spinulose, not echinate pollen, (2) conspicuous spinulose opercula over the apertures, and (3) conspicuous perforations (Nowicke, 1994
; Punt and Höen, 1995
). Many have much larger perforations through the sexine (Nowicke, 1994
; Punt and Höen, 1995
). Some species, such as Melandrium apricum and Silene aperta, are finely reticulate (Nowicke, 1994
; Punt and Höen, 1995
), but this is due to large perforations unlike the fine ones of the fossil (Fig. 10). Illecebrum verticillatum, like the fossil, has echinate grains with fine perforations and little elaboration of the exine over the apertures, but has only 6–9 pores (Punt and Höen, 1995
). The pollen is also distinct from other periporate Caryophyllales (e.g., Figs. 54–57). In particular, Amaranthaceae (including Chenopodiaceae) have annulate pores.
Caryophylloflora paleogenica is clearly distinct in floral form from any of the eight genera and 40 or so species of Caryophyllaceae native to Australia (Hnatiuk, 1990
). Gypsophila lacks bracts appressed to the calyx. Arenaria, Stellaria, Spergularia, Colobanthus, Polycarpea, and Drymaria have free sepals. In Scleranthus the stamens arise from the top of a hypanthium. Caryophylloflora paleogenica is therefore best considered as a member of an extinct genus. To establish the relationships of C. paleogenica within Caryophyllaceae more precisely would require a detailed phylogeny. This may be indecisive considering the morphological convergence in the order and lack of resolution among extant species (Fig. 1; Smissen et al., 2002
).
Evolutionary and biogeographic implications
Caryophylloflora paleogenica implies that Caryophyllaceae were in Australia by the Late Eocene. It also confirms that at least some fossil pollen assigned to Periporopollenites polyoratus was produced by Caryophyllaceae. The placement of the fossil among or as sister to the higher Caryophyllaceae suggests that considerable divergence had occurred within the family by the Late Eocene. Furthermore, the distinct nature of C. paleogenica and the absence of P. polyoratus after the Oligocene implies that Paleogene Caryophyllaceae in Australia may not have been closely related to extant Australian members of the family.
Most extant Australian genera of Caryophyllaceae are best interpreted as having arrived during the Neogene or Quaternary. Arenaria, Gypsophila, Stellaria, and Spergularia are cosmopolitan or primarily Eurasian, and the Australian species are identical to or closely related to temperate Eurasian species. Fossil pollen closely resembling that produced by Stellaria (Polyporina granulatus Martin 1973) and Silene (Polyporina reticulata Martin 1973) first appear in southern Australia during the Late Miocene-Early Pliocene (Macphail, 1997
). This suggests relatively recent dispersal. Polycarpea and the Australian species of Drymaria, D. cordata, are more or less pantropical (Bittrich, 1993
). Transoceanic dispersal appears to have been significant in both Colobanthus and Scleranthus. The former occurs on isolated islands of relatively recent origin (e.g., Macquarie Island), and species of the latter (S. biflorus and S. brockei) occur in both Tasmania and New Zealand, a distribution unlikely to have arisen through vicariance (see Jordan, 2001
). Thus, although C. paleogenica demonstrates that the family grew in the Southern Hemisphere by the Middle-Late Eocene, modern Australian Caryophyllaceae appear to be independent, recent arrivals.
|
|
FOOTNOTES |
|---|
2 Author for reprint requests (greg.jordan{at}utas.edu.au
)
3 Current address: Department of Archaeology and Natural History, Research School of Pacific and Asian Studies, Australian National University, Canberra 0200, A. C. T. Australia
|
LITERATURE CITED |
|---|
|
TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSSYSTEMATIC DESCRIPTIONDISCUSSIONLITERATURE CITED |
|---|
Barnes R. W. G. J. Jordan 2000 Eucryphia (Cunoniaceae) reproductive and leaf macrofossils from Australian Cainozoic sediments. Australian Systematic Botany 13: 373-394[CrossRef][ISI]
Behnke H.-D. 1994 Sieve-element plastids: their significance for the evolution and systematics of the order. In H.-D. Behnke and J. T. Mabry [eds.], Caryophyllales: evolution and systematics, 87–122. Springer Verlag, Berlin, Germany
Bittrich V. 1993 Caryophyllaceae. In K. Kubitzki, J. G. Rohwer, and V. Bittrich [eds.], The families and genera of vascular plants, vol. II, Flowering plants, dicotyledons, Magnoliid, Hamamelid and Caryophyllid Families, 206–236. Springer-Verlag, Berlin, Germany
Carpenter R. J. R. S. Hill G. J. Jordan 1994 Cenozoic vegetation in Tasmania. In R. S. Hill [ed.], History of the Australian vegetation: Cretaceous to Recent, 276–298. Cambridge University Press, Cambridge, UK
Chandler E. M. 1961 Flora of the Lower Headon Beds of Hampshire and the Isle of Wight. Bulletin of the British Museum of Natural History: Geology 5: 91-158
Chandler E. M. 1963 The lower Tertiary floras of South England. III. Flora of the Bournemouth Beds; the Boscombe, and the Highcliff Sands. The British Museum, London, UK
Clements J. S. J. T. Mabry H. Wyler A. S. Drieding 1994 Chemical review and evolutionary significance of betalaines. In H.-D. Behnke and J. T. Mabry, [eds.], Caryophyllales: evolution and systematics, 247–262. Springer Verlag, Berlin, Germany
Correa M. N. 1984 Flora Patagonica. Instituto Nacional de Tecnologia Agropecuaria, Buenos Aires, Argentina
Cronquist A. 1981 An integrated system of classification of flowering plants. Columbia University Press, New York, New York, USA
Cuenod P. V. Savolainen L. W. Chatrou M. Powell R. E. J. Grayer M. W. Chase 2002 Molecular phylogenetics of Caryophyllales based on nuclear 18s rDNA and plastid rbcL, atpB, and matK DNA sequences. American Journal of Botany 89: 132-144
Davis P. H. 1967 Flora of Turkey and the East Aegean Islands, vol. 2. Edinburgh University Press, Edinburgh, UK
Dilcher D. L. 2000 Toward a new synthesis: major evolutionary trends in the angiosperm fossil record. Proceedings of the Natonal Academy of Sciences 97: 7030-7036[CrossRef]
Downie S. R. D. Palmer 1994 A chloroplast DNA phylogeny of the Caryophyllales based on structural and inverted repeat restriction site analysis. Systematic Botany 19: 236-252[CrossRef][ISI]
Dyer R. A. 1975 The genera of southern African flowering plants, vol. 1, Dicotyledons. Department of Agriculture Technical Services, Pretoria, South Africa
Eliasson U. 1993 35A, Phytolaccaceae, Flora of Ecuador, 46. University of Goteborg Press, Goteborg, Sweden
Heusser C. J. 1971 Pollen and spores of Chile. Modern types of the Pteridophyta, Gymnospermae, and Angiospermae. University of Arizona Press, Tucson, Arizona, USA
Hill R. S. 1989 New species of Phyllocladus (Podocarpaceae) macrofossils from south eastern Australia. Alcheringa 13: 193-208
Hill R. S. R. J. Carpenter 1991 Evolution of Acmopyle and Dacrycarpus (Podocarpaceae) foliage as inferred from macrofossils in south-eastern Australia. Australian Systematic Botany 4: 449-479[CrossRef]
Hill R. S. D. C. Christophel 1988 Tertiary leaves of the tribe Banksieae (Proteaceae) from southeastern Australia. Botanical Journal of the Linnean Society 97: 205-227
Hill R. S. M. S. Pole 1992 Leaf and shoot morphology of extant Afrocarpus, Nageia and Retrophyllum (Podocarpaceae) species, and species with similar leaf arrangement from Tertiary sediments in Australasia. Australian Systematic Botany 5: 337-358
Hnatiuk R. J. 1990 Census of Australian vascular plants. Australian Government Publishing Service, Canberra, Australia
Hofmann U. 1994 Flower morphology and ontogeny. In H.-D. Behnke and J. T. Mabry [eds.], Caryophyllales: evolution and systematics, 123–166. Springer Verlag, Berlin, Germany
Jordan G. J. 2001 An investigation of long-distance dispersal based on species native to both Tasmania and New Zealand. Australian Journal of Botany 49: 333-340[CrossRef][ISI]
Jordan G. J. R. S. Hill 2000 The phylogenetic affinities of Nothofagus leaf fossils based on combined molecular and morphological data. International Journal of Plant Science 160: 1177-1188
Källersjö M. J. S. Farris A. G. Kluge C. Bult 1992 Skewness and permutation. Cladistics 8: 275-287
Kubitzki K. J. G. Rohwer V. Bittrich 1993 The families and genera of vascular plants, vol. II, Flowering plants, dicotyledons, Magnoliid, Hamamelid and Caryophyllid Families. Springer-Verlag, Berlin, Germany
Macphail M. K. 1997 Late Neogene climates in Australia: fossil pollen- and spore-based estimates in retrospect and prospect. Australian Journal of Botany 45: 425-464[CrossRef]
Macphail M. K. N. Alley E. M. Truswell I. R. Sluiter 1994 Early Tertiary vegetation: evidence from pollen and spores. In R. S. Hill [ed.], History of the Australian vegetation: Cretaceous to Recent, 189–261. Cambridge University Press, Cambridge, UK
Manhart J. R. J. H. Rettig 1994 Gene sequence data. In H.-D. Behnke and J. T. Mabry [eds.], Caryophyllales: evolution and systematics, 235–246. Springer Verlag, Berlin, Germany
Manly W. C. C. R. Hutchins 1980 A flora of New Mexico. J. Cramer, Vaduz, Germany
Moar N. T. 1993 Pollen grains of New Zealand dicotyledonous plants. Manaaki Whenua Press, Lincoln, New Zealand
Muller J. 1981 Fossil pollen records of extant angiosperms. Botanical Review 47: 1-142
Nott J. F. J. A. K. Owen 1992 An Oligocene palynoflora from the middle Shoalhaven catchment N.S.W. and the Tertiary evolution of flora and climate in the southeast Australian highlands. Palaeogeography, Palaeoclimatology, Palaeoecology 95: 135-151[CrossRef]
Nowicke J. W. 1994 Pollen morphology and exine ultrastructure. In H.-D. Behnke and J. T. Mabry [eds.], Caryophyllales: evolution and systematics, 168–221. Springer Verlag, Berlin, Germany
Nowicke J. W. 1996 Pollen morphology, exine structure and the relationships of Basellaceae and Didieraceae to Portulacaceae. Systematic Botany 21: 187-208[CrossRef][ISI]
Nowicke J. W. J. J. Skvarla 1979 Pollen morphology: the potential influence in higher order systematics. Annals of the Missouri Botanic Gardens 66: 633-700[CrossRef][ISI]
Owen J. A. K. 1988 Miocene palynomorph assemblages from Kiandra, New South Wales. Alcheringa 12: 269-297[ISI]
Prendini L. 2001 Species or supraspecific taxa as terminals in cladistic analysis? Groundplans versus exemplars revisited. Systematic Biology 50: 290-300[CrossRef][ISI][Medline]
Punt W. P. P. Höen 1995 Caryophyllaceae. Review of Palaeobotany and Palynology 88: 82-272
Raine J. I. 1984 Outline of a palynological zonation of Cretaceous to Paleogene terrestrial sediments in West Coast region, South Island, New Zealand. New Zealand Geological Survey Report 109: 1-82
Rodman J. E. 1994 Cladistic and phenetic studies. In H.-D. Behnke and J. T. Mabry [eds.], Caryophyllales: evolution and systematics, 279–301. Springer Verlag, Berlin, Germany
Roles S. J. 1957 Flora of the British Isles, illustrations, part 1. Cambridge University Press, Cambridge, UK
Ross-Craig S. 1951 Drawings of the British Flora, vol. 5, Caryophyllaceae. G. Bell and Sons, London, Great Britain
Savolainen V. et al 2000 Phylogeny of the eudicots: a nearly complete familial analysis based on rbcL gene sequences. Kew Bulletin 555: 257-309
Simmons M. P. H. Ochoterena 2000 Gaps as characters in sequence-based phylogenetic analyses. Systematic Biology 49: 369-381[CrossRef][ISI][Medline]
Smissen R. D. J. C. Clement P. J. Garnock-Jones G. K. Chambers 2002 Subfamilial relationships within Caryophyllaceae as inferred from 5' ndhF sequences. American Journal of Botany 89: 1336-1341
Stover L. E. A. D. Partridge 1973 Tertiary and Late Cretaceous spores and pollen from the Gippsland Basin, south-eastern Australia. Proceedings of the Royal Society of Victoria 85: 237-286
Swofford D. L. 2000 PAUP*: phylogenetic analysis using parsimony (*and other methods), version 4. Sinauer, Sunderland, Massachusetts, USA
Truswell E. M. I. R. Sluiter W. K. Harris 1985 Palynology of the Oligocene-Miocene sequence in the Oakvale-1 corehole, western Murray Basin, South Australia. BMR Journal of Australian Geology and Geophysics 9: 267-295
Tulip J. R. G. Taylor E. M. Truswell 1982 Palynology of Tertiary Lake Bunyan, Cooma, New South Wales. BMR Journal of Australian Geology & Geophysics 7: 255-268[ISI]
This article has been cited by other articles:
|
|
 |
|
  S. R. S. Cevallos-Ferriz, E. Estrada-Ruiz, and B. R. Perez-Hernandez Phytolaccaceae infructescence from Cerro del Pueblo Formation, Upper Cretaceous (late Campanian), Coahuila, Mexico Am. J. Botany, January 1, 2008; 95(1): 77 - 83. [Abstract] [Full Text] [PDF]
|
|
| ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
| HOME | HELP | FEEDBACK | SUBSCRIPTIONS | ARCHIVE | SEARCH | TABLE OF CONTENTS |
