The Morphology of Pollen Tetrads and Viscin Threads in Some Tertiary,Rhododendron-Like Ericaceae

The morphology of pollen tetrads and viscin threads is described in fossil Ericaceae pollen from various Eocene/Oligocene/Miocene localities in Europe (Germany, Austria), North America (eastern U.S.A.), and Asia (eastern China). The typical characters of the tetrad configuration, the exine ornamentation and sculpturing, and especially the viscin thread morphology are extremely similar to or even indistinguishable from that in extant members of Rhododendron. All these pollen morphological features strongly suggest that all the investigated material can be assigned to a modern taxon of the Ericaceae: either to Rhododendroideae or even to Rhododendron itself.     

Exine architecture and function in some lythraceae and sonneratiaceae

The pollen morphology of selected species of Crenea, Lagerstroemia, Lafoensia and Diplusodon (Lythraceae) and of Sonneratia (Sonneratiaceae) is compared from a functional point of view. From a basically tricolporate type, multiplication of colpi, reduction of colpi, development of meridional ridges, development of polar caps, differentiation of pores and reduction of pores have led to partly diverging, partly converging structural types which are interpreted as adaptation to controlled harmomegathy. In some species of Lagerstroemia with dimorphic stamens, the exine of the pollen produced by the feeding anthers shows a specialized structure, which enables the pollen grain to retain moisture and become sticky. For Sonneratia the fossil record allows a check on the postulated evolutionary trends.     

Fossil pollen grains of Asteraceae from the Miocene of Patagonia: Nassauviinae affinity

Fossil pollen grains with morphological features unique in the subtribe Nassauviinae (tribe Mutisieae, Asteraceae) occur in Miocene marine deposits of eastern Patagonia, southern South America. A new morphogenus and two morphospecies are proposed to assemble fossil pollen grains characterized by having a complex bilayered exine structure with delicate columellae, separated by an internal tectum. Subprolate specimens with Trixis exine type (ectosexine thinner than endosexine, straight internal tectum) are referred to Huanilipollis cabrerae. This species is similar to pollen of recent Holocheilus, Jungia, and Proustia. Suboblate specimens with Oxyphyllum exine type (ectosexine and endosexine equally thick, zigzag internal tectum) are referred to Huanilipollis criscii. This species is similar to pollen of recent Triptilion. The spore/pollen sequences in which Nassauviinae pollen types occur suggest a wide range of vegetation types varying from forest dominated during the Early Miocene (Chenque Formation) to virtually xerophytic ones during the Late Miocene (Puerto Madryn Formation). The subtribe Nassauviinae comprises 25 genera and ca. 320 species of vines, shrubs and low trees endemic to America with a wide range of ecological preferences; the nearest living relatives of the fossil types being mostly confined to humid landscapes. The unusual occurrence of these groups during the arid characterized Late Miocene time could be attributed to the complex interplay of the mountain uplift and global circulation patterns. These forcing factors would have created a mosaic of different habitats with both patches of forest and dry-adapted species developing in relatively small regions. This is the first fossil record of Nassauviinae and confirms that this subtribe of Asteraceae was already differentiated in the Miocene.     

PALYNOLOGY AND SYSTEMATICS OF THE LYTHRACEAE. II. GENERA HAITIA THROUGH PEPLIS

Pollen morphology for 12 genera of the Lythraceae is described, using light microscopy (LM), scanning (SEM) and transmission electron microscopy (TEM). The genera surveyed are Haitia, Heimia, Hionanthera, Lafoensia, Lagerstroemia, Lawsonia, Lythrum, Nesaea, Orias (= Lagerstroemia), Pehria, Pemphis, and Peplis. Among results of taxonomic significance are 1) the similarity of pollen between Haitia and Ginoria, suggesting that future classifications reflect a close relationship between these genera, 2) a comparable similarity in pollen between Pehria and Adenaria, 3) the distinction of Hionanthera pollen from that of Ammannia, 4) an exine similar in certain ultrastructural characteristics between Orias and some species of Lagerstroemia, although in general aspect they appear quite different, 5) that within the Didiplis-Lythrum-Peplis complex, there are three palynologically distinct groups, with Didiplis standing apart from the somewhat more similar, but nevertheless, distinct Lythrum and Peplis, and 6) that considerable differences exist between pollen of Pemphis acidula and Pemphis madagascariensis, supporting removal of the latter species to a new genus. This pollen survey is part of a multi-disciplinary effort to clarify generic limits in the Lythraceae, and evaluate relationships within the family and among families of the Myrtales.     

Diverse fossil Onagraceae pollen from a Miocene palynoflora of north-east China: early steps in resolving the phytogeographic history of the family

The origin and evolution of angiosperms can be unravelled by using fossil records to determine first occurrences and phytogeographic histories of plant families and genera. Many angiosperm families, for example the Onagraceae, have a poor macrofossil record, but are more common in palynological records. Modern Onagraceae produce pollen clearly distinct from that of other angiosperms. Combined morphological features obtained by use of light and scanning electron microscopy have enabled assignment of fossil Onagraceae pollen to extant genera, and therefore tracing of the origin and past distributions of extant Onagraceae lineages. We studied a Miocene palynoflora from the Daotaiqiao Formation of north-east China. Using the single-grain technique, we examined individual Onagraceae pollen/tetrads using both light and scanning electron microscopy. Fossil Onagraceae pollen is more frequent than macrofossil remains, but is still rare, and usually represented by a single taxon in palynological samples. Remarkably, samples from the Miocene of north-east China contain five different species: two of Circaea, one of Epilobium, and two of Ludwigia. Such a large number of Onagraceae taxa from a single palynoflora is unknown elsewhere. Whereas Ludwigia pollen is known from Cenozoic sediments of the northern hemisphere, the Circaea pollen is the first fossil pollen assignable to this extant genus. This is also the first fossil record of Epilobium from China. Although the young geological age of this sample does not enable consideration of time of origin for the genera encountered, the co-occurrence of Circaea, Epilobium, and Ludwigia in the mid to late-Miocene of East Asia sheds some light on their phytogeographic histories.     

Vegetation, climate and palaeoaltitude reconstructions of the Eastern Alps during the Miocene based on pollen records from Austria, Central Europe

Aim To reconstruct the flora, vegetation, climate and palaeoaltitude during the Miocene (23.03–5.33 Ma) in Central Europe. Location Six outcrop sections located in different basins of the Central Paratethys in Austria. Methods Pollen analysis was used for the reconstruction of the vegetation and climate. The altitude of the Eastern Alps that are adjacent to the Alpine Foreland and Vienna basins has been estimated using a new quantification method based on pollen data. This method uses biogeographical and climatological criteria such as the composition of the modern vegetation belts in the European mountains and Miocene annual temperature estimates obtained from fossil pollen data. Results Pollen changes from Early to Late Miocene have been observed. The vegetation during the Burdigalian and Langhian (20.43–13.65 Ma) was dominated by thermophilous elements such as evergreen trees, typical of a present‐day evergreen rain forest at low altitudes (i.e. south‐eastern China). During the Serravallian and Tortonian (13.65–7.25 Ma) several thermophilous elements strongly decreased, and some disappeared from the Central European region. This kind of vegetation was progressively substituted by one enriched in deciduous and mesothermic plants. Middle‐altitude (Cathaya, Cedrus and Tsuga) and high‐altitude (Abies and Picea) conifers increased considerably during the Langhian and later on during the Serravallian and Tortonian. Main conclusions Pollen changes are related to climatic changes and to the uplift of the Alpine massifs. The vegetation during the Burdigalian and Langhian reflects the Miocene climatic optimum. The decrease in thermophilous plants during the Serravallian and Tortonian can be interpreted as a climatic cooling and can be correlated with global and regional climatic changes. This study shows that the palaeoaltitude of the eastern part of the Eastern Alps during the Burdigalian was not high enough for Abies and Picea to form a forest. Therefore, we inferred that the summits of most of the mountains would have been less than 1800 m. The substantial increase of middle‐ and high‐altitude conifers in the pollen spectra suggests that the uplift rate increased during the Langhian in this region. Based on higher palaeoaltitude estimations for the pollen floras from the studied sections of Austria, we infer that the uplift of the easternmost part of the Alpine chain continued during the Serravallian and Tortonian.     

<Emphasis Type="Italic">Altingioxylon hainanensis</Emphasis> sp. nov.: earliest fossil wood record of the family Altingiaceae in Eastern Asia and its implications for historical biogeography

A new species, Altingioxylon hainanensis, is described from the Eocene Changchang Formation of the Changchang Basin on Hainan Island, South China. It is the first record of a fossil wood assigned to Altingiaceae found in China, and the most ancient evidence of wood for this family in eastern Asia. The new species is similar to A. rhodoleioides, known since the Miocene in India and Java Island, and to Altingia hisauchii from the Miocene to Pliocene of Japan. The close resemblance between these species and Liquidambar sp., known from the Middle Miocene of western North America, provides additional evidence for the migration of their ancestors from Asia to North America across the Bering land bridge during the Miocene. Distinctions in ray sizes between the eastern Asian specimens and their contemporaries from Europe to Kazakhstan is suggested as a result of the divergence between the large eastern Asian clade and the North American–west Asian clade within Altingiaceae during the Eocene–Oligocene. The presence of crystals in ray cells may be considered an ancestral condition that persists in the eastern Asian lineages up to the extant Altingia and Semiliquidambar, but which was lost in other Altingiaceae in the course of evolution.     

New fossil leaves and fruits of Lauraceae from the Middle Miocene of Fujian,southeastern China differentiated using a cluster analysis

The fossil record of Lauraceae can be traced back to the Early Cretaceous of eastern Asia based on fossil flowers. Here, we refer a number of new occurrences of leaf and fruit fossils of Lauraceae from the Middle Miocene of Zhangpu, Fujian, China, to seven species. These data provide evidence supporting the fact that a diverse subtropical, or tropical, Lauraceae-dominated evergreen forest surrounded this region 15 million years ago (Mya). The Lauraceae fossils presented in this paper provide evidence for the evolution of this group as well as new materials that enable the study of the Fujian Province Neogene flora. The fossils described in this paper fill in the gaps in studies about Lauraceae pollen in the Middle Miocene from Fotan, Fujian, China. In addition, these fossils also enrich the Middle Miocene fossil records of Lauraceae in eastern Asia, especially improving the study of the macrostructures and reproductive organs of fossil Lauraceae from southern China. The similarity between fossil and modern fruits shows that during the Middle Miocene the fruit morphological of Lauraceae have changed very little. We also identify families where the fossils we report belong to their closest relatives and can be used to reconstruct the paleoenvironment of Fujian in the Middle Miocene.     

Warm and humid Trans-Himalaya during the late Miocene: plant fossil evidence

Neogene fossil records from the Indus Basin sedimentary rocks (IBSR), deposited in the Indus Tsangpo Suture Zone (ITSZ), are very rare, but are important to understand the history of plant diversity and paleoclimate in the Himalaya. We report fossil wood ascribed to Ebenoxylon siwalicus Prakash from late Miocene sediments of the Karit Formation belonging to ITSZ. The anatomical details of the fossil wood, such as small to medium-sized vessels occluded with tyloses, scanty paratracheal to diffuse-in-aggregate axial parenchyma, 1–3 seriate homo to heterocellular rays, bordered intervessel pits with lenticular apertures and simple perforations, suggest its close affinity with Diospyros Linnaeus of the family Ebenaceae. Further anatomical details suggest a close resemblance with extant D. ehretioides Don and D. macrophylla Blume. The present fossil, along with previously known fossil records of Lagerstroemia (Lythraceae) and palms, indicate that the Trans-Himalaya was warm and humid during the late Miocene, quite different from the modern cool and dry climate in the study area.     

A Review of the Fossil Record of the Genus Itea (Iteaceae, Saxifragales) with Comments on its Historical Biogeography

Itea is a genus of about 20 species of trees and shrubs that are today native to southeastern North America, eastern Asia, and eastern Africa. In this paper, I review the fossil record of Itea, which is based on four types of fossils: diporate, psilate pollen attributed to Itea or the dispersed pollen genus Iteapollis; carpofossils representing fruits and seeds attributed to Itea europaea; flowers preserved in amber and assigned to Adenanthemum iteoides; and leaf impressions attributed to Itea. The distributions of these fossils indicate that Itea was present in western North America from the early Eocene to Miocene, in eastern North America beginning no later than the early Miocene, and in western Eurasia from the late Eocene to Pliocene. Only one datapoint is known from eastern Asia; it is early Miocene in age. Based on the fossil record, it can be inferred that Itea crossed between continents over both the Bering Land Bridge and North American Land Bridge, and that it reached Africa from Europe via Anatolia. Thus, it is predicted that the sole extant North American species, I. virginica, may be most closely related to the sole extant African species, I. rhamnoides. The potential application of Itea fossils to calibrating phylogenetic trees generated from molecular sequence data is also discussed.     

基于荧光原位杂交技术的紫薇属植物核型分析

以紫薇(Lagerstroemia indica)、尾叶紫薇(L.caudata)、屋久岛紫薇(L.fauriei)和福建紫薇(L.limii)4种紫薇属植物为材料,利用染色体荧光原位杂交技术(FISH)获得了4种紫薇属植物的有丝分裂中期染色体FISH图及核型参数,分析了45SrDNA在紫薇属植物染色体上的数量和分布特点。结果表明,4种紫薇属植物染色体上均具有1对45SrDNA杂交位点,位于较长染色体短臂的近端部,紫薇、尾叶紫薇、屋久岛紫薇和福建紫薇的核型公式分别为2n=48=2M+24m+22sm、2n=48=30m+18sm、2n=48=2M+20m+26sm和2n=48=2M+32m+14sm,均为2A型。该研究首次获得了紫薇属植物45SrDNA荧光原位杂交核型,为紫薇属植物亲缘关系研究和细胞生物学研究提供了分子细胞学依据。     

Fossil Records in the Lythraceae

The Lythraceae (Myrtales) are a family of 28 genera and ca. 600 species constituting with the Combretaceae and sister family Onagraceae a major lineage of the Myrtales and including the former Sonneratiaceae, Duabangaceae, Punicaceae, and Trapaceae. The fossil record of the family is extensive and significant new discoveries have been added to the record in recent years. This review provides a vetted summary of fossils attributed to the Lythraceae, their geographic distributions, and their stratigraphic ranges. It anticipates the use of the information to generate robustly dated molecular phylogenies to accurately reconstruct the evolutionary and biogeographic history of the family. Fossils of 44 genera or form genera have been attributed to the Lythraceae; 24 are accepted here as lythracean. Fourteen of the 28 modern genera have fossil representatives: Adenaria, Crenea, Cuphea, Decodon, Duabanga, Lafoensia, Lagerstroemia, Lawsonia, Lythrum, Pemphis, Punica, Sonneratia, Trapa, and Woodfordia. Ten extinct genera are recognized. The most common kinds of fossil remains are seeds and pollen. The only fossil flower confidently accepted in the family is the extinct genus Sahnianthus from the Early Paleocene of India. The oldest confirmed evidence of the Lythraceae is pollen of Lythrum/Peplis from the Late Cretaceous (early Campanian, 82?81 Ma) of Wyoming. Seeds of Decodon from the late Campanian (73.5 Ma) of northern Mexico are next oldest. Sonneratia, Lagerstroemia, and extinct Sahnianthus first appear in the Paleocene of the Indian subcontinent; extinct Hemitrapa fruits first occur in the Paleocene of northwestern North America. Diversification of the Lythraceae occurred primarily during two major periods of global temperature change, during the Paleocene-Eocene Thermal Maximum and from the middle Miocene forward when temperatures decreased markedly and seasonality and dry-adapted vegetation types became more prominent. Fossils of the Lythraceae from South America and Africa are limited in number. The few dates available for South American genera are comparatively young and diversification of the largest genus, Cuphea (ca. 240 species), was mainly a Quaternary event. A phylogeny of the family is briefly explored and examples of specialized characters occurring in the oldest known genera are noted. The fossil record of the Lythraceae is presently too fragmentary to confidently reconstruct the early history of the family. The record indicates, however, that the family was well-diversified and widely dispersed globally over a wide latitudinal range by the end of the Paleocene.     

Paleobotanical remains from the Paleocene–lower Eocene Vagadkhol Formation, western India and their paleoclimatic and phytogeographic implications

A small assemblage of macro- and micro floral remains comprising fossil leaf impressions, silicified wood, spores, and pollen grains is reported from the Paleocene–lower Eocene Vagadkhol Formation (=Olpad Formation) exposed around Vagadkhol village in the Bharuch District of Gujarat, western India. The fossil leaves are represented by five genera and six species, namely, Polyalthia palaeosimiarum (Annonaceae), Acronychia siwalica (Rutaceae), Terminalia palaeocatapa and T. panandhroensis (Combretaceae), Lagerstroemia patelii (Lythraceae), and a new species, Gardenia vagadkholia (Rubiaceae). The lone fossil wood has been attributed to a new species, Schleicheroxylon bharuchense (Sapindaceae). The palynological assemblage, consisting of pollen grains and spores, comprises eleven taxa with more or less equal representation of pteridophytes, gymnosperms, and angiosperms. Angiospermous pollen grains include a new species Palmidites magnus. Spores are mostly pteridophytic but some fungal spores were also recovered. All the fossil species have been identified in the extant genera. The present day distribution of modern taxa comparable to the fossil assemblage recorded from the Vagadkhol area mostly indicate terrestrial lowland environment. Low frequency of pollen of two highland temperate taxa (Pinaceae) in the assemblage suggests that they may have been transported from a distant source. The wood and leaf taxa in the fossil assemblage are suggestive of tropical moist or wet forest with some deciduousness during the Paleocene–early Eocene. The presence of many fungal taxa further suggests the prevalence of enough humidity at the time of sedimentation.     

A new species of Macaranga from the middle Miocene of Fujian,China and its significance

A well-preserved Macaranga leaf fossil from the middle Miocene Fotan Group of Zhangpu County, Fujian, South-eastern China is examined and described as a new species, Macaranga zhangpuensis Z.X. Wang et B.N. Sun sp. nov. The present fossil species represents the highest latitudinal distribution of a reliable Macaranga fossil in the world, and we present the first Macaranga fossil described with detailed cuticular characteristics from China. Based on the global palaeogeographic distribution of Macaranga, we infer that the genus probably originated during the Oligocene in Africa and spread from Africa to India and then to South Fujian, China, further into the Mariana Islands and finally into the Philippine Islands and Malaysia, leading to its present distribution. In addition, the new leaf material described herein is the first Macaranga fossil record with three leaf tips globally. This finding demonstrates that Macaranga with three leaf tips already existed during the Miocene in Fujian, South China, and it provides new information for understanding climatic changes between the Miocene and the present -day.     

Leaves and pollen of bamboos from the Polish Neogene

Recently discovered fossil bamboo leaves and pollen are described from the Neogene of Poland. Morphological analysis of fossil leaves from the Late Miocene deposits of the Bełchatów Lignite Mine, Central Poland have allowed them to be determined as “Bambusa” lugdunensis Saporta. It is the first record of this species in the Cenozoic of Poland. The fossils have been compared with other records of bamboos from the Cenozoic of Europe. Pollen grains from the Middle Miocene (Badenian) deposits from the Legnica brown coal deposit (Lower Silesia) provide additional evidence of bamboos. These are assigned to Graminidites bambusoides Stuchlik, and are similar to those of recent Arundinaria. We infer that plant (plants?) that produced the pollen of G. bambusoides could have grown in swamp forests and/or reed marshes. This report provides a general survey of the occurrence of bamboos in the European Neogene, from which the leaves and pollen documented here present important data on the distribution of this group of monocots in Poland and eastern Europe.     

The Phytogeography of the extinct angiosperm Nordenskioeldia (Trochodendraceae) and its response to climate changes

Elucidating the spatio-temporal distributions of terrestrial plants is a key for interpreting the origin of distribution patterns and the tempo of intercontinental disjunction. Nordenskioeldia was distributed in eastern Asia and North America from the Late Cretaceous to the Miocene. Its fossil record provides important information on former patterns of disjunction and dispersal in the Northern Hemisphere. New specimens from the Paleocene of China allow us to further extend the history of the group and provide the impetus to review its distribution in space and time. The comparative morphological survey on fossil Nordenskioeldia found in the Paleocene sediments in both eastern Asia and North America confirms that they belonged to the same morpho-species, which indicates a close floristic continuity between both continents due to land connection available during that time. The spatio-temporal distributions of Nordenskioeldia indicate that the taxon probably expanded eastward from eastern Asia into North America by the end of Early/Middle Maastrichtian, subsequently colonized Greenland, northeastern North America and Spitsbergen in the Early Paleocene, and finally became extinct in the Miocene. The fluctuations in its northern limits took place in response to climate changes: warming from the Paleocene to the Eocene, cooling during the Eocene–Oligocene and amelioration during the Late Oligocene–Mid-Miocene.     

FUCHSIA POLLEN FROM THE EARLY MIOCENE OF NEW ZEALAND

Fossil pollen of Diporites aspis Pocknall & Mildenhall, from the early Miocene of New Zealand, was examined by combined light, scanning, and transmission electron microscopy, in order to determine its relationship to extant Fuchsia pollen. Based on overall morphology, exine structure, exine surface sculpture, viscin thread morphology, and apertural features, the fossil pollen definitely can be considered to represent Fuchsia. Parallel studies of extant Fuchsia pollen indicate that a more precise identification of the fossils is not possible. With the examination of additional fossil material, it should be possible to learn a great deal about the timing of origin and migration of the genus Fuchsia, but not of its constituent parts.     

Rudabánya: A late miocene subtropical swamp deposit with evidence of the origin of the African apes and humans

Rudabánya, a rich late Miocene fossil site in northern central Hungary, has yielded an abundant record of fossil primates, including the primitive catarrhine Anapithecus and the early great ape Dryopithecus. While the affinities of Anapithecus are not clear, Dryopithecus is clearly a great ape sharing numerous characteristics of its dental, cranial and postcranial anatomy with living great apes. Like all Miocene hominids (great apes and humans), Dryopithecus is more primitive in a number of ways than any living hominid, which is probably related to the passage of time since the divergence of the various lineages of living hominids, allowing for similar refinements in morphology and adaptation to take place independently. On the other hand, Dryopithecus (and Ouranopithecus) share derived characters with hominines (African apes and humans), and Sivapithecus (and Ankarapithecus) share derived characters with orangutans, thus dating the split between pongines and hominines to a time before the evolution of these fossil great apes. Pongines and hominines follow similar fates in the late Miocene, the pongines moving south into Southeast Asia from southern or eastern Asia and the hominines moving south into East Africa from the Mediterranean region, between 6 to 9 Ma.     

The diversity and paleoenvironmental significance of Calophyllum (Clusiaceae) from the Miocene of southeastern China

Abstract

Three species within the genus Calophyllum collected from middle Miocene Fotan Group sediments in Zhangpu County, Fujian, southeastern China are described in this paper. These fossils include Calophyllum zhangpuensis sp. nov., Calophyllum striatum, and Calophyllum suraikholaensis. The new fossil species C. zhangpuensis sp. nov. is oval, possesses entire leaves with closely spaced parallel secondary veins and has a round, or slightly retuse, apex. These specimens represent the first known fossil records of this relative wide leaf-type form of Calophyllum from China and have a length:width (L:W) ratio less than 3:1. In combination with the known modern geographic distribution and habitats of this wide leaf-type Calophyllum and other plants, data suggest that the middle Miocene Fotan flora is indicative of a warm climate. Thus, based on available fossil data, we speculate that this genus probably originated in India during the Paleocene before spreading from India to Bangladesh and into China, Sumatra, Malaysia, Indonesia, and Java during the Neogene, leading to its modern distribution. At least, the 3 fossil species in this region can explain floristic exchange between India, Fujian, and South China, which is consistent with previous studies; the occurrence of these 3 species indicates that Calophyllum began to diversity in China no later than the Miocene.     

Pollen morphology and fossil record of the feathery mistletoe family Misodendraceae

Misodendraceae is a small family of mistletoes in the order Santalales. Its distribution is restricted to the southern South American temperate forests. The family comprises the sole genus Misodendrum with eight species of hemiparasitic shrubs, mainly parasitising the southern beech Nothofagus. This contribution presents palynological evidence from seven species, using light microscopy and scanning electron microscopy. Pollen grains are consistently small, periporate and echinate, although differences in the length of echini and number and size of pores were noted. Pollen features can be used to distinguish groups of species and, in some cases, individual species. Cluster analysis of pollen characters differentiates two main groups: one includes M. brachystachyum, M. oblongifolium and M. quadriflorum; and the other includes M. gayanum, M. linearifolium, M. punctulatum and M. angulatum. Palynological results are compared with previous systematic studies of the family. The South American fossil pollen record is summarised and characters of the fossil pollen are analysed using UPGMA to test the relationships between extant and fossil species. Miocene pollen resulted similar to species of subgenus Angelopogon while Eocene pollen is disimilar to extant species of Misodendraceae.