Contrasted pollen capture mechanisms in Phyllocladaceae and certain Podocarpaceae (Coniferales)

Comparative study shows that Phyllocladus and representative Podocarpaceae differ in the mechanism by which pollen is introduced into the pollen chamber and onto the apex of the nucellus ("pollen capture"). Both types involve a pollination drop, but only in Podocarpaceae is it consistently inverted and in contact with adjacent surfaces. Phyllocladus has functionally nonsaccate pollen (although a vestigial saccus has been claimed); its pollen is wettable and sinks in water. Podocarpaceae (except Saxegorhaea) have saccate pollen, which is nonwettable and floats on water. In Phyllocladus the pollination drop receives the pollen directly and presence of pollen stimulates complete drop withdrawal, which may be a metabolic process. Once pollinated, an ovule does not resecrete a pollination drop. In Podocarpaceae the drop usually receives the pollen indirectly via pollen scavenging and saccate pollen is preferentially captured. The retraction of the drop appears to be the result of evaporation and is presumably nonmetabolic. Drop secretion can be repeated in the presence of pollen. A major consequence of these contrasted mechanisms is that in Phyllocladus the entire contents of the pollination drop are ingested, whereas in Podocarpaceae only that part of the drop that includes saccate pollen is ingested. Because of differences in repeatability of the secretion process, Podocarpaceae are likely to capture more pollen. In neither mechanism does the process favor 'own" pollen. but in Podocarpaceae all but saccate pollen is excluded. We thus have further evidence for differences in pollen capture mechanisms in conifers with a pollination drop, and differences in the behavior of the pollination drop itself.     

Osmunda (Osmundaceae) from the Triassic of Antarctica: an example of evolutionary stasis

Compressed specimens of the fern Osmunda are described from the Triassic of the Allan Hills, Antarctica. The specimens consist of a once pinnate, deeply pinnatifid fertile frond as well as several sterile specimens. Six pinnae are present on the partial fertile rachis, with two sterile pinnae above four fertile pinnae. Both sterile and fertile specimens are virtually identical to the modern species Osmunda claytoniana. Entire fronds are fragmentary; the longest is 21 cm in length. Sterile pinnae are alternate and deeply pinnatifid, with slightly toothed pinnules and dichotomous venation. Fertile pinnae are 1-1.3 cm long, once pinnate, and lack vegetative lamina. Sporangia are clustered, each 300-375 um in diameter, and possess a transverse annulus 6-8 cells long; dehiscence is by a vertical slit. Fronds arise from a rhizome 4 cm long by 1 cm wide; two croziers are present on the rhizome. Two frond segments up to 6 cm long and three deeply pinnatifid pinnae are present on the uppermost part of one rachis. Pinnules are ~4 mm long and 2-3 mm wide. The presence of this Osmunda species in the Triassic demonstrates stasis of frond morphology, both fertile and vegetative, for the genus.     

New perspectives on the Mesozoic seed fern order Corystospermales based on attached organs from the Triassic of Antarctica

A new Triassic corystosperm is described from the Shackleton Glacier region of Antarctica. The compression fossils include cupulate organs (Umkomasia uniramia) and leaves (Dicroidium odontopteroides) attached to short shoot-bearing branches. The cupulate organs occur in groups near the apices of the short shoots, and each consists of a single axis with a pair of bracts and a subapical whorl of five to eight ovoid cupules. This unique architecture indicates that the cupules are individual megasporophylls rather than leaflets of a compound megasporophyll. A branch bearing an attached D. odontopteroides leaf provides the first unequivocal evidence that Umkomasia cupulate organs and Dicroidium leaves were produced by the same plants. Although this had previously been assumed based on organ associations, the new specimens are important in demonstrating that a single species of corystosperm produced the unique cupulate organs described here and the geographically and stratigraphically widespread and common D. odontopteroides leaf. Therefore, biostratigraphic, paleoecological, and phylogenetic studies that treat Dicroidium leaf morphospecies as proxies for biological species of entire plants should be reconsidered. Phylogenetic analysis suggests that the corystosperm cupule is an unlikely homologue for the angiosperm carpel or outer integument.     

Developmental shoot morphology in Phyllocladus (Podocarpaceae)

TOMLINSON, P. B., TAKASO, T. & RATTENBURY, J. A., 1989. Developmental shoot morphology in Phyllocladus (Podocarpaceae). Shoot architecture in the adult phase of Phyllocladus is established by a succession of units of extension that develop a system of permanent axes supporting photosynthetic units (phylloclades) each of which represents a branch complex with three branch orders. Seedlings have needle foliage leaves comparable to those of other conifers, but in adult plants all leaves are ephemeral, non-photosynthetic scales that for the most part subtend no axillary buds. Once rhythmic growth (usually seasonal) is established in the adult phase, each increment produces a whorl of phylloclades so that a regular tiered arrangement develops, with the tiers progressively reduced on outer units. In the resting terminal bud of permanent axes only scale primordia are present; with bud burst and beginning of extension of the unit the phylloclades are produced by syllepsis and complete the initiation and expansion of all axis orders in the short flushing cycle. Segments retain strict distichy throughout, but in a dorsiventral and not a lateral plane. Phylloclades may be either determinate, when the apex of the first-order axis develops as a terminal flattened segment, or indeterminate, when the apex retains radial symmetry and forms a resting bud that can continue axis extension as a permanent shoot in subsequent years. A phylloclade consequently only produces flattened lateral segments in its season of initiation. Reiteration from reserve buds is not possible, because none are produced in the adult phase, but is possible from detached meristems formed in the axils of needle leaves on juvenile shoots. Reiteration in the adult phase is thus possible only by axis dedifferentiation, that is, change from plagiotropy to orthoptropy. The distinctive massive vascular connection of the phylloclade is made possible by syllepsis. In this way the normal structural constraints of elaborate appendage development in conifers is fully overcome.     

A taxodiaceous seed cone from the Triassic of Antarctica

A silicified seed cone is described from the lower Middle Triassic of A silicified seed cone is described from the lower Middle Triassic of Antarctica. The cone measures up to 3.4 cm long and 1.4 cm wide, and consists of helically arranged cone scales attached to a eustelic axis. Bract and ovuliferous scale are approximately of equal length and fused at the base. The bract is entire and vascularized by a single trace. The ovuliferous scale contains five distal lobes, each vascularized by a terete strand that divides to form a smaller trace to each of the five inverted ovules. Ovules are small and flattened with the three-parted integument attenuated into oppositely positioned lateral wings. The Triassic specimens are compared with both extant and fossil conifer seed cones and believed to have their closest affinities within the Taxodiaceae.     

Studies on Pollen Morphology of Podocarpaceae from China

The pollen morphology and ultrastructure of exine of Podocarpaceae in China were examined with light microscope, scanning electron microscope and transmission electron microscope. The pollen grains of Podocarpus have rather large and prominent sacs on both sides of body, and are 53.9-74.8 μm long in total, with their bodies 29.6-45.2 μm long and 19.1-31.3 μm wide. The sacs are smooth on outer surface with perforation, but reticulate inside. On distal view, they are obviously of radial muri from its base. The body is oblate or spheroidal, laddershaped on distal face. The exine of the capis tuberculate, but more distinctly on the margin than in the centre. The pollen grains of Dacrydium are of small and indistinct sacs around body, which are composed of many small bladders. The body is subcircular in outline. Both body and sacs are irrugulate tuberculate under SEM. Examination of thin sections of Podocarpus macrophyllus var. maki with TEM reveals that the exine includes ectexine and endexine. It is interesting to note that foot layer of ectexine possesses lamellar stru cture, but endexine is homogeneous in structure and lighter in colour. This character is specific in the gymnosperms. Based on informations of fossil pollen grains, Podocarpaceae is rather primitive and of ancient origin.     

Microsporogenesis and microgametogenesis of the Argentinian species of Podocarpus (Podocarpaceae)

The development of the microsporangium and male gametophyte of three species of Podocarpus was studied with light microscopy (LM) and the morphology of pollen with scanning and transmission electron microscopy (SEM and TEM). During early stages, the male cone is covered with coriaceous scales. The archesporid cells go through a dormant period. Later the pollen mother cells differentiate and undergo meiosis. Callose is detected around the tetrad and between each monad. The microspore nucleus divides several times to give rise to a multicellular gametophyte, which includes the tube cell, the stalk and body cells, and four prothallial cells. The exine of the pollen grain is rugulate in the corpus and quite smooth in the sacci. The ultrastructure of the pollen wall consists of the alveolate sexine, the laminate nexine I and the amorphous nexine II. The intine is very thin. Comparison of the mature grain of some fossils with living members of the Podocarpaceae reveals great similarity.     

Rainforest conifers of Eocene Patagonia: Attached cones and foliage of the extant Southeast Asian and Australasian genus Dacrycarpus (Podocarpaceae)

? Premise of the study: Eocene caldera-lake beds at Laguna del Hunco (LH, ca. 52.2 Ma) and Río Pichileufú (RP, ca. 47.7 Ma) in Argentine Patagonia provide copious information about the biological history of Gondwana. Several plant genera from these sites are known as fossils from southern Australia and New Zealand and survive only in Australasian rainforests. The potential presence of Dacrycarpus (Podocarpaceae) holds considerable interest due to its extensive foliage-fossil record in Gondwana, its remarkably broad modern distribution in Southeast Asian and Australasian rainforests, its high physiological moisture requirements, and its bird-dispersed seeds. However, the unique seed cones that firmly diagnose Dacrycarpus were not previously known from the fossil record. ? Methods: I describe and interpret fertile (LH) and vegetative (LH and RP) material of Dacrycarpus and present a nomenclatural revision for fossil Dacrycarpus from South America. ? Key results: Dacrycarpus puertae sp. nov. is the first fossil occurrence of the unusual seed cones that typify living Dacrycarpus, attached to characteristic foliage, and of attached Dacrycarpus pollen cones and foliage. Dacrycarpus puertae is indistinguishable from living D. imbricatus (montane, Burma to Fiji). Dacrycarpus chilensis (Engelhardt) comb. nov. is proposed for Eocene vegetative material from Chile. ? Conclusions: Modern-aspect Dacrycarpus was present in Eocene Patagonia, demonstrating an astonishingly wide-ranging paleogeographic history and implying a long evolutionary association with bird dispersers. Dacrycarpus puertae provides the first significant Asian link for Eocene Patagonian floras, strengthens the biogeographic connections from Patagonia to Australasia across Antarctica during the warm Eocene, and indicates high-rainfall paleoenvironments.     

Evolution and biogeography of the austral genus Phyllocladus (Podocarpaceae)

Aim To infer evolutionary relationships within the genus Phyllocladus and among its close relatives by phylogenetic analysis of DNA sequences. Interpret the inferred relationships in association with the fossil record to examine the origin and diversification of the genus. Location Australasia. Methods Phylogenetic analyses of rbcL, matK and internal transcribed spacer (ITS) sequences representing all of the extant species of Phyllocladus and a selection of outgroups from Podocarpaceae and Araucariaceae. Results The rbcL and matK sequences exhibit little variation within Phyllocladus, but ally its members to Podocarpaceae although its immediate sister remains unclear. The ITS sequences resolve all five species of Phyllocladus and two intraspecific ecotypes of P. alpinus. Main conclusions Phyllocladus forms a distinct lineage that diverged early in the evolutionary history of Podocarpaceae. The fossil record indicates that the genus was more widely distributed and morphologically diverse during the early Tertiary than at present. Although of Mesozoic origin, the level of sequence variation within Phyllocladus suggests that the extant species radiated during the late Tertiary c. 6.3 ± 0.9 Ma. New Zealand is the present centre of species diversity.     

Fossilized spermatozoa preserved in a 50-Myr-old annelid cocoon from Antarctica

The origin and evolution of clitellate annelids—earthworms, leeches and their relatives—is poorly understood, partly because body fossils of these delicate organisms are exceedingly rare. The distinctive egg cases (cocoons) of Clitellata, however, are relatively common in the fossil record, although their potential for phylogenetic studies has remained largely unexplored. Here, we report the remarkable discovery of fossilized spermatozoa preserved within the secreted wall layers of a 50-Myr-old clitellate cocoon from Antarctica, representing the oldest fossil animal sperm yet known. Sperm characters are highly informative for the classification of extant Annelida. The Antarctic fossil spermatozoa have several features that point to affinities with the peculiar, leech-like ‘crayfish worms'' (Branchiobdellida). We anticipate that systematic surveys of cocoon fossils coupled with advances in non-destructive analytical methods may open a new window into the evolution of minute, soft-bodied life forms that are otherwise only rarely observed in the fossil record.     

Structurally preserved sphenophytes from the Triassic of Antarctica: reproductive remains of Spaciinodum

Permineralized cones found organically attached to Spaciinodum collinsonii stems are described from the early Middle Triassic silicified flora from the Fremouw Formation of Antarctica, and the species diagnosis is emended to include the reproductive specimens. The apical cones are organized into internodal and leaf-bearing nodal regions. Nodal septations span the central pith and cortex, and thin fimbrils subdivide the internodal areas into smaller chambers. The vascular system consists of 31-33 continuous bundles that do not alternate in position between successive nodes and internodes. Simple sporangia are associated with the cortical chambers and occur in one whorl on the axis. Spores are small, lack elaters, and have no discernible ultrastructure preserved, and they are interpreted to be immature. The Antarctic cones are different in structure from typical cones of modern and fossil members of Equisetales; however, they share similarities with some morphologically aberrant cones of extant Equisetum and several Late Paleozoic and Mesozoic compression-impression fossils. Spaciinodum is now the most complete anatomically described Mesozoic sphenophyte.     

Remarkably preserved annelid worms from the La Meseta Formation (Eocene), Seymour Island, Antarctica

Abstract: Worm tubes, which exhibit the replaced tube-lining membrane, have been collected from the lowermost Lower Eocene Acantilados Allomember of the La Meseta Formation, Seymour Island, Antarctica. The discovery represents the oldest examples of preservation of the tube-lining membrane of tube-dwelling Polychaeta in the fossil record. A new genus and species, Caprascolex antarcticus , are described. The specimens are preserved as thin coatings of amorphous iron oxide on the inner surface of the moulds interpreted to be the replaced tube-lining membrane. Examination of the rarely described tube-lining mucosoid membrane in extant polychaetes shows that the fossils are nearly identical in morphology and scale to extant forms. These fossils record in remarkable detail the morphology of the tube-lining membrane, which appears to be composed of growth bands formed as the worm constructed the tube. The tube-lining is believed to have been originally preserved as pyrite, with subsequent oxidation to iron oxide. The tube-lining membrane of worm tubes possibly is known from only one other fossil occurrence.     

Northern Hemisphere megafossil of Dacrycarpus (Podocarpaceae) from the Miocene of South China and its evolutionary and paleoecological implications

The modern genus Dacrycarpus (Endl.) de Laub. of the family Podocarpaceae, containing nine species, is mainly distributed in tropical mountain rainforests of the southwestern Pacific region, ranging from New Zealand to low‐latitude Asia. This genus has abundant fossil records in both hemispheres, but all the known megafossils were limited to Australasia and South America. Here we report on Dacrycarpus guipingensis sp. nov. from the Miocene Erzitang Formation of Guangxi, South China. This is the first megafossil of Dacrycarpus in the Northern Hemisphere. The new species is represented by mummified dimorphic foliage, ovuliferous shoots, and a male cone with in situ pollen. It resembles the extant Dacrycarpus imbricatus (Blume) de Laub., which is common in rainforests from southern China and northern Myanmar to Fiji. This paper presents the first data on the anatomical structure of seed cone and exine ultrastructure of Dacrycarpus in situ pollen grains from a fossil material by using computed tomography scanning and ultrathin sectioning. For comparative purpose, data on the pollen morphology and ultrastructure were obtained for modern D. imbricatus for the first time. The D. guipingensis fossils strongly suggest the Miocene arrival of Dacrycarpus in Asia from the Southern Hemisphere. Based on the modern ecological niche and related fossil elements, this fossil locality was probably covered by conifer–broad‐leaved mountain rainforests during the Miocene.     

Phylogeny of Podocarpaceae: comparison of evidence from morphology and 18S rDNA

Maximum parsimony analyses of the genera of Podocarpaceae were conducted using sequence data from 18S ribosomal DNA. Trees from sequence, morphological, and combined data differ in taxon arrangement, but are similar in that Podocarpus sensu lato and Dacrydium s.l. are unnatural, while Podocarpaceae (including Phyllocladus) are monophyletic. The clade Microcachrys + Microstrobos is recognized in all analyses, but its placement differs, i.e., nested among other scale-leaved taxa in the morphological analysis, but associated with Nageia and other tropical genera in the sequence analyses. Trees from combined data reflect this ambiguity. Podocarpus sensu stricto is paraphyletic according to most trees. Inferences of plesiomorphic character states within the family are largely consistent between analyses and support the view that prototypical podocarps had bifacial leaves, cones with several fertile cone scales, and large epimatia (cone scales) that covered the inverted ovules.     

Evolutionary relationships of the New Caledonian heterotrophic conifer, Parasitaxus usta (Podocarpaceae), inferred from chloroplast trnL-F intron/spacer and nuclear rDNA ITS2 sequences

 The phylogenetic position of Parasitaxus (Podocarpaceae) has been inferred from a cladistic analysis of molecular characters from chloroplast and nuclear genomes including all genera of Podocarpaceae. In all 24 most parsimonious trees, based on combined datasets, Phyllocladus resided outside Podocarpaceae s. str. while Lepidothamnus was basal to the latter. Most other genera were arranged in two major clades. The evidence confirms previous studies, which have suggested a relationship between Lagarostrobos, Manoao and Parasitaxus. Parasitaxus is not directly related to its host Falcatifolium taxoides. Instead it appears to be most closely related to Manoao and Lagarostrobos. No other members of this group now occur on New Caledonia. However, if the evolution of Parasitaxus were autochthonous, a free-living member of this group must once have occurred there. An accelerated evolutionary rate of the chloroplast sequence analysed was suggested, indicating that the plant behaves like a holoparasite. Received January 4, 2002; accepted April 3, 2002 Published online: September 13, 2002     

A unique mode of parasitism in the conifer coral tree Parasitaxus ustus (Podocarpaceae)

Almost all parasitic plants, including more than 3000 species, are angiosperms. The only suggested gymnosperm exception is the New Caledonian conifer, Parasitaxus ustus , which forms a bizarre graft-like attachment to the roots of another conifer Falcatifolium taxoides . Yet, the degree of resource dependence of Parasitaxus on Falcatifolium has remained speculative. Here we show that Parasitaxus is definitively parasitic, but it displays a physiological habit unlike any known angiosperm parasite. Despite possessing chloroplasts, it was found that the burgundy red shoots of Parasitaxus lack significant photosynthetic electron transport. However unlike non-photosynthetic angiosperm parasites (holoparasites), tissues of Parasitaxus are considerably enriched in ¹³carbon relative to its host. In line with anatomical observations of fungal hyphae embedded in the parasite/host union, stable carbon isotopic measurements indicate that carbon transport from the host to Parasitaxus most likely involves a fungal partner. Therefore, Parasitaxus parallels fungus-feeding angiosperms (mycoheterotrophs) that steal carbon from soil mycorrhizal fungi. Yet with its tree-like habit, association with fungi residing within the host union, high stomatal conductance, and low water potential, it is demonstrated that Parasitaxus functions unlike any known angiosperm mycoheterotroph or holoparasite. Parasitaxus appears to present a unique physiological chimera of mistletoe-like water relations and fungal-mediated carbon trafficking from the host.     

The Ichnofossils of the Triassic Hope Bay Formation,Trinity Peninsula Group,Antarctic Peninsula

The Triassic Hope Bay Formation (Trinity Peninsula Group) includes a diverse ichnocoenosis in the Puerto Moro succession (Hope Bay, Antarctic Peninsula). The Hope Bay Formation is a thick turbidite succession with a minimum vertical exposure of 533 meters along the Hope Bay coast. The rocks are locally affected by contact metamorphism related to later arc magmatism. The ichnofossils are found mainly in thick- and thin-bedded sandstone-mudstone facies composed of a monotonous repetition of sandstone-mudstone cycles. The sandstones are usually medium grained, massive or parallel laminated; the mudstones are massive and rarely laminated. In the fine-grained rocks, mainly the mudstones, there are distinct densities of bioturbation, and at least six patterns were observed. The following ichnogenera were recognized: Arenicolites Salter 1857, Lophoctenium Richter 1850, Taenidium Heer 1877, Palaeophycus Hall 1847, Phycosiphon von Fischer-Ooster 1858 and Rhizocorallium Zenker 1836. All appear to be feeding-traces. The trace fossil assemblages occur mainly in black mudstones rich in organic material that suggest a low oxygen environment. The stratigraphic interval in which they occur is interpreted as progradational supra-fan lobes with channel fill and levee deposits. The thin-bedded turbidite and mudstone lithofacies, where the ichnofossils are abundant, is interpreted as a distal fan turbidite or levee deposit related to a long-term channel fill. This study is the first significant report of trace fossils in the Hope Bay Formation.