Reproductive structures of Rhamnaceae from the Cerro del Pueblo (Late Cretaceous, Coahuila) and Coatzingo (Oligocene, Puebla) Formations, Mexico

Recently discovered fossil flowers from the Cretaceous Cerro del Pueblo and flowers and fruits from the Oligocene Coatzingo Formations are assigned to the Rhamnaceae. The Cretaceous flower, Coahuilanthus belindae Calvillo-Canadell and Cevallos-Ferriz, gen. et sp. nov., is actinomorphic with fused perianth parts forming a slightly campanulate to cupulate floral cup, with sepals slightly keeled and spatulate clawed petals. The Oligocene fossils include Nahinda axamilpensis Calvillo-Canadell and Cevallos-Ferriz, gen. et sp. nov. (characterized by its campanulate bisexual flower with stamens opposite, adnate to and enfolded by petals; and with the ovary ripening into a drupe), and a winged fruit assigned to Ventilago engoto Calvillo-Canadell and Cevallos-Ferriz, sp. nov. The flowers and drupe features indicate closer affinity to Zizipheae and/or Rhamneae, while the single samaroid fruit suggests the presence of Ventilagineae. However, the unique character combination in the fossil flowers precludes placing them in extant genera. Nevertheless, the history of the family is long and can be traced back to the Campanian. A detailed phylogenetic revision of the group that uses morphological characters from both extant and fossil plants is needed to better understand the significance of these records as well as other important fossils of the family.     

Upper Cretaceous aquatic plants from Northern Mexico

Two aquatic plant genera assignable to Decodon (Lythraceae) and Ceratophyllum (Ceratophyllaceae) are described based on reproductive structures collected from the Cerro del Pueblo Formation (late Campanian [73.5 ma]), Coahuila, Northeast Mexico. Decodon is represented by three small seeds with a pyramidal shape, rounded borders, and a concave ventral surface with a rectangular valve towards the center of the seed ventral surface. The Ceratophyllum spiny fruit has an ellipsoidal central body and two proximal long spines flanking a short pedicel opposite the stylar projection. These new reports confirm the presence of both genera in the Upper Cretaceous sediments of Northeastern Mexico, and add to our recognition of diversity within the widely distributed freshwater communities along the margin of the epicontinental sea.     

Identification of asymmetrically winged samaras from the Western Hemisphere

A key is presented for use in identifying asymmetrically winged fruits (samaras) with either proximal or distal locules. It aids identification based on dispersed fuit morphology and can be used to identify undetermined extant herbarium specimens or fossil fruits to the correct extant family and genus. The 39 genera from 11 families (Aceraceae, Anacardiaceae, Fabaceae, Malpighiaceae, Phytolaccaceae, Polygalaceae, Polygonaceae, Rutaceae, Sapindaceae, Trigoniaceae, Ulmaceae) are distinguished on the basis of wing venation, size of fruit, presence and position of attachment surface, presence and type of subsidiary wings on the ovary wall. ornamentation, size and shape of the ovary, locule position, shape of locule cross section, style position and ornamentation, distinction between ovary wall and wing, and angle of attachment between individual samaras. The developmental origins of some of these features are discussed.     

Miocene winged fruits of Loxopterygium (Anacardiaceae) from the Ecuadorian Andes

A new species of asymmetrically winged fruit is described from Miocene sediments of Andean Ecuador. The new fruit is readily placed in the genus Loxopterygium of the Anacardiaceae based on the size, position of the stigma, wing venation, and serration of the wing tip. The new fossil species is very similar to extant species of Loxopterygium now distributed in dry habitats of coastal Ecuador and Peru, as well as dry interior forests of Bolivia and northern Argentina. We use the fossil to calibrate a molecular-based phylogeny of some members of the Anacardiaceae, showing that dry forest habitats may have been present in South America for more than 10 million years.     

Late Miocene Palaeocarya (Engelhardieae: Juglandaceae) from Southwest China and its biogeographic implications

Fossil fruits of Palaeocarya (Juglandaceae) are described from late Miocene sediments of southeastern Yunnan, China. The fruits present a tri-lobed wing consisting of an intact oblong-ovate middle lobe and two lateral lobes. The lobes are apically obovate, and have pinnate venation. The middle lobe is thicker at the base and gradually tapers to the apex. The nutlet, located at the base of the winged fruit, is round and hispid, and is subdivided by a septum into two compartments. Based on extensive morphological comparisons to previously documented fossil fruits, we found that the fossil fruits align most closely with members of the genusPalaeocarya, but have a unique combination of characters. Thus, we describe the fossils as a new species, Palaeocarya hispida sp. nov. This species represents an important range expansion for low-latitude occurrences of Palaeocarya in the late Miocene and therefore substantially improves our understanding of the biogeographic history of the genus. We propose that the wide distribution of Palaeocarya and relatively narrow distributions of close relatives, Engelhardia, Alfaropsis, and Oreomunnea, might be associated with a stepwise cooling and a major ice sheet expansion in the Antarctic and Arctic from the late middle Miocene to early Pliocene. In particular, the climatic oscillations during the Quaternary, such as the last glacial maximum, might have led to a decrease in the geographic distribution of Engelhardieae.     

Fruits of Tetrapterys (Malpighiaceae) from the Oligocene of Hungary and Slovenia

Propeller-like winged fruits of Tetrapterys harpyiarum Unger from the Oligocene of Sotzka, Budapest, Eger-Vécsey valley, and a new occurrence at Eger-Kiseged, were reinvestigated and compared in detail with extant species of Tetrapterys (Malpighiaceae) and with other dicotyledonous genera with four winged fruits. T. harpyiarum fruits are bilaterally symmetrical, consisting of a globose nut surrounded by four elongate wings with parallel venation. Tetrapterys is now distributed only in tropical America and this implies that there was an opportunity for Tetrapterys to spread between the Partethys region and the New World during the Tertiary.     

A fossil fruit wing of Dipterocarpus from the middle Miocene of Fujian, China and its palaeoclimatic significance

Dipterocarpus zhengae sp. nov. is described from the middle Miocene Fotan Group of Zhangpu county, Fujian Province, Southeast China on the basis of a fruit wing. Three kinds of venation of the calyx longer lobes (enlarged sepals) occur in the winged fruits of extant Dipterocarpus. The fossil species is referred to the kind having three primary veins and within that category is most similar to extant Dipterocarpus gracilis Blume in the size of the longer lobe as well as its venation. The occurrence of the fruit wing of Dipterocarpus, together with palynological evidence, indicates unequivocally that a tropical climate and tropical rain forest occurred in Zhangpu during the middle Miocene, warmer and more humid than at the present day. The palaeobiogeography of Dipterocarpus is reviewed.     

Reconstructing the ancestral angiosperm flower and its initial specializations

Increasingly robust understanding of angiosperm phylogeny allows more secure reconstruction of the flower in the most recent common ancestor of extant angiosperms and its early evolution. The surprising emergence of several extant and fossil taxa with simple flowers near the base of the angiosperms-Chloranthaceae, Ceratophyllum, Hydatellaceae, and the Early Cretaceous fossil Archaefructus (the last three are water plants)-has brought a new twist to this problem. We evaluate early floral evolution in angiosperms by parsimony optimization of morphological characters on phylogenetic trees derived from morphological and molecular data. Our analyses imply that Ceratophyllum may be related to Chloranthaceae, and Archaefructus to either Hydatellaceae or Ceratophyllum. Inferred ancestral features include more than two whorls (or series) of tepals and stamens, stamens with protruding adaxial or lateral pollen sacs, several free, ascidiate carpels closed by secretion, extended stigma, extragynoecial compitum, and one or several ventral pendent ovule(s). The ancestral state in other characters is equivocal: e.g., bisexual vs. unisexual flowers, whorled vs. spiral floral phyllotaxis, presence vs. absence of tepal differentiation, anatropous vs. orthotropous ovules. Our results indicate that the simple flowers of the newly recognized basal groups are reduced rather than primitively simple.     

Molecular data reveal convergence in fruit characters used in the classification of Thlaspi s. I. (Brassicaceae)

Phylogenetic relationships of 18 Thlaspi s.l. species were inferred from nuclear ribosomal internal transcribed spacer (ITS) sequence data. These species represent all sections of the basic classification system of Schulz primarily based on fruit characters. The molecular phylogeny supported six clades that are largely congruent with species groups recognized by Meyer on the basis of differences in seed coat anatomy, i.e. Thlaspi s. s., Thlaspkeras, Moccaea {Raparia included), Microthhspi, Vania and Neurotropy. Some of these lineages include species which are morphologically diverse in fruit shape (e.g. Thlaspi s. s.: T. arvense - fruits broadly winged, T. ceratocarpum - fruits with prominent horns at apex, T. alliaceum - fruits very narrowly winged). Furthermore, the same fruit shape type is distributed among different clades. For instance, fruits with prominent horns at apex are found in Thlaspi s. s. ( T. ceratocarpum) and Thlaspiceras (T oxyceras). These results clearly indicate convergence in fruit characters previously used for sectional classification in Thlaspi s. l.     

FOSSIL CERATOPHYLLUM (CERATOPHYLLACEAE) FROM THE TERTIARY OF NORTH AMERICA

Fossil fruits and a vegetative axis assignable to the extant genus Ceratophyllum are described from four North American Tertiary localities. Fossil fruits assignable to the extant species C. muricatum and C. echinatum are reported from the Eocene Green River and Claiborne formations, and the Miocene Esmerelda Formation, respectively. An extinct species, C. furcatispinum, is described from the Paleocene Fort Union Formation and represents the oldest published report of Ceratophyllum in the fossil record. The existence of extant angiosperm species in the Eocene is very unusual and may be attributable in this case to slow evolutionary rates and unusual evolutionary properties associated with hydrophily in the genus Ceratophyllum.     

Two new species of Echinobothrium van Beneden, 1849 (Cestoda: Diphyllidea) from the Persian Gulf

Two new species of diphyllidean cestodes are described from the Persian Gulf, Echinobothrium persiense n. sp. from Rhinobatos punctifer Compagno & Randall and Echinobothrium hormozganiense n. sp. from Mustelus mosis Hemprich & Ehrenberg. E. persiense is the first record of a species of Echinobothrium van Beneden, 1849 from R. punctifer and these two new species increase the number of diphyllideans known from the Persian Gulf from one to three. The number of apical hooks of E. persiense (hook formula: {5-6 6/5 5-6}) is distinct from all other species in the genus except for E. affine Diesing, 1863, E. harfordi McVicar, 1976, E. bonasum Williams & Campbell, 1980, E. fautleyae Tyler & Caira, 1999, E. syrtense (Neifar, Tyler & Euzet, 2001) Tyler, 2006 (emend), E. chisholmae Jones & Beveridge, 2001, E. tetabuanense Ivanov & Caira, 2012, E. sematanense Ivanov & Caira, 2012 and E. weipaense Ivanov & Caira, 2012. Echinobothrium persiense can be distinguished from all other species of the genus with 11 apical hooks by a combination of the following features: armed cephalic peduncle, testes arranged in a single column, lateral hooklets arranged in two groups, U-shaped ovary, cephalic peduncle length (124-181 μm), genital pore and cirrus-sac position, and by having 10-14 spines per row on the cephalic peduncle. Echinobothrium hormozganiense has a hook formula of {12-15 16/15 12-15} and is similar to E. musteli Pintner, 1889, E. notoguidoi Ivanov, 1997 and E. diamanti Ivanov & Lipshitz, 2006 by possessing additional spines between the rostellum and the bothria. It differs from E. musteli by having an H-shaped ovary and lateral hooklets arranged in two lateral groups, and the number of spines per row on the cephalic peduncle (18-21) readily differentiates it from E. notoguidoi (24-26) and E. diamanti (95-118). With these two new species, Echinobothrium van Beneden, 1849 now includes 45 valid species.     

23种伞形科植物果实形态及其分类学意义

该实验采用徒手切片法对伞形科17属23种植物的果实进行了外部形态和解剖特征的观察,结果表明:伞形科果实有背腹压扁、两侧压扁和不压扁3种类型;侧棱有宽、有窄;油管的分布有棱槽单油管型和棱槽多油管型;花柱基多数为圆锥状,少数种的花柱基为扁平垫状;萼齿明显或不明显。通过进一步对属间和属内果实解剖特征的比较得出:(1)果实表面被钩刺或刚毛及果棱特征在属间差异明显,在属内表现出一致性,可作为伞形科属间分类的依据。(2)果实横切面的形状、胚乳腹面的凹凸以及萼齿的形态特征在属内种间的分类研究中有重要的意义。基于果实形态特征,编制了17属23种植物的分种检索表。     

Highly specialized Cretaceous beetle parasitoids (Ripiphoridae) identified with optimized visualization of microstructures

Extremely miniaturized longipedes insects (body length c. 0.3 mm) embedded in two pieces of Cretaceous amber from Myanmar are described and interpreted. Using inverted fluorescence and light microscopy for detailed analysis of microstructures, the inclusions were identified as primary larvae of the beetle family Ripiphoridae, subfamily Ripidiinae. While the structure of thoracic and abdominal segments including appendages corresponds well with the groundplan known in recent members of Ripidiinae, a curved prosternal ridge with prominent spines (each c. 5 μm), the reduced condition of stemmata and antennae and the lack of sharp mandibles are unique features within the entire family, apparently apomorphies of the longipedes larvae. A sinuate prosternal edge with a dense row of spines (prosternoctenidium) might be homologous with ‘head ctenidia’ in some previously described miniaturized conicocephalate larvae, but further investigation is needed. The morphological differences between the head of longipedes larvae and extant Ripidiinae are interpreted as adaptations to different groups of hosts and life strategies. Palaeoethology of the longipedes larvae is briefly discussed. In addition, the systematic placement of conicocephalate larvae from Canadian, Myanmar and Russian Cretaceous ambers, already interpreted by various authors as primary instars within Coleopterida (assigned to either Strepsiptera or to the coleopteran Tenebrionoidea: Ripiphoridae), is discussed.     

中国安息香科一新属──长果安息香属

本文描述了安息香科－新属──长果安息香属,并对新组合长果安息香的原始描述作了修订。本属与秤锤树属近缘,但村干无茎刺,具鳞芽,总状花序伞形,花无苞片,四基数,花萼管顶端平截,花冠４深裂,雄蕊８,等长,药隔不突出,子房半下位,柱头不分裂,果实特长,密被长柔毛,上部果喙细长,中部倒卵形,具８条纵梭,下部渐狭,延长成柄状,可以区别。本属的芽具鳞片,花无苞片,四基数,与银钟花属相似,但后者的花萼管顶端具４裂片,雄蕊８～１６,不等长,子房下位,柱头４分裂,果实无毛,具４或２翅和缩存花柱基部,无果喙,与前者明显不同。文中还给出了安息香科分属检索表。     

Numerical taxonomy of Palaeocarya (Juglandaceae) from the Mangbang Formation of West Yunnan, China

Specimens of Palaeocarya (Juglandaceae) were collected from the upper Pliocene of western Yunnan Province, which is a new geographical and age range record of engelhardioid winged fruits in China. A numerical taxonomic investigation based on 14 morphological characters of fossil and extant specimens of engelhardioid winged fruits in China was conducted using multivariate methods. Phenetic clustering shows that there are four morphotypical groups which we assign to four species. Two groups are assigned to the previously described fossil species, Palaeocarya guangxiensis Li Hao-Min et Chen Yun-Fa, 2003 and Palaeocarya koreanica (Oishi) Manchester, 1987. Two new species are recognized, Palaeocarya yunnanensis and Palaeocarya longialata. These Neogene Engelhardieae fruits provide fossil data for understanding the origin and evolution of the Engelhardia species that now inhabit South China.     

Fruit features in relation to the ecology and distribution of

Fruit features of 17 Acaena species in New Zealand were investigated to determine the relations between reproductive allocation patterns, mode of dispersal and species ecology. Three basic morphological types, corresponding to major sections within the genus, are represented: spineless fruits (Sect. Pteracaena—one species), spined fruits lacking barbs (Sect. Microphyllae—6 species), and spined fruits with barbs (Sect. Ancistrum—10 species). The presence of terminal barbs on spines enhances fruit adhesion to animals, and is associated with the development of longer scapes, globular many-fruited capitula, and smaller single-seeded fruits. Barb-spined species generally have the broadest geographical range and habitat distribution within New Zealand, and are the only species of Acaena to occur in forests or on offshore islands. These species are generally strongly stoloniferous. Acaena species with barbless spines and/or no spines are often regional endemics confined to open and/or localised habitats, are rhizomatous, and may be dispersed by water, ingestion or wind. There is a tight correlation between intrageneric classification and species ecology and fruit features which requires testing via independent phylogenetic analysis based on molecular genetic characters. Although avian dispersers have declined since human settlement, introduced mammals may be performing a similar function for barbed Acaena species.     

Inflorescence bracts of fossil and extant Tilia in North America,Europe, and Asia: patterns of morphologic divergence and biogeographic history

The enlarged inflorescence bract diagnostic of extant Tilia has an extensive Tertiary fossil record in the Northern Hemisphere. Diversity of bract morphology, and the extent of adnation between peduncle and bract, is reviewed for fossil and extant species of Tilia. An extinct type of bract with an orbicular outline and palmate venation is documented by the fossil species Tilia circularis (Chaney) comb. nov. from the early Oligocene of Oregon and is designated Type A. Living species of the genus have elongate bracts with predominately pinnate venation that are borne in two basic configurations: Type B, with the peduncle fused only to the extreme base of the bract lamina, as in extant Tilia endochrysea Hand.-Mzt. of southern China; and Type C with the peduncle fused medially along the basal one-third of the bract lamina, as in most extant species. Bracts of Type B were widely distributed in the Tertiary of western North America (late Eocene to Miocene) and Europe (early Miocene to Pliocene), while those of Type C are known in the fossil condition only from the middle and late Tertiary of Asia and Pliocene of Europe. The bracts of T. circularis, like those of type B, are borne on relatively long stalks and have the peduncle fused only at the extreme base. The fossil record supports recognition of the following characters as apomorphic in Tilia bract evolution: bracts sessile, peduncle adnate to the upper surface of the bract, and pinnate bract venation.     

Evolution of fruit characters and dispersal modes in the tropical family Rubiaceae

We investigated the evolution of fruit characters, animals versus abiotic dispersal modes, life forms and geographical distribution, in the large, mostly tropical, family Rubiaceae. As a basis for our analysis we used a phylogenetic tree derived from chloroplast DNA variation. Fleshy fruits have evolved independently at least 12 times in the family. Most of these originations appear to have occurred during Eocene to Oligocene, i.e. the radiation period for some animal taxa (bird families, mammal orders) comprising most extant dispersers of Rubiaceae fruits. Changes of dispersal modes may be of more recent origin in a few cases, e.g. evolution of drupes in some lineages, and shifts from drupes to nuts. The distribution of fruit characters suggested that in several lineages animal-dispersed fruits, such as berries and drupes have remained largely unaltered since the time of origination. This is in contrast to the occurrence of winged seeds in capsules, and pterophylls, i.e. enlarged calyx lobes promoting wind dispersal of fruits, which apparently have shifted more frequently during evolution, indicating a difference in 'phylogenetic plasticity' between modes of animal and wind dispersal.
Animal dispersal was over-represented among genera dominated by shrubs, whereas abiotic dispersal was most prevalent among herbaceous genera. Drupes were over-represented in groups with transoceanic distributions, and on islands, indicating dispersal over long distances, probably by birds. In contrast, no evidence was found to support the view that animal dispersal in general enhances long distance dispersal. We also analysed geographical patterns on the tribal level but these were too complex to yield any resolved area cladograms due to the occurrence of many widespread taxa and area redundancy.     

Phylogenetic Distribution and Identification of Fin-winged Fruits

Fin-winged fruits have two or more wings aligned with the longitudinal axis like the feathers of an arrow, as exemplified by Combretum, Halesia, and Ptelea. Such fruits vary in dispersal mode from those in which the fruit itself is the ultimate disseminule, to schizocarps dispersing two or more mericarps, to capsules releasing multiple seeds. At least 45 families and more than 140 genera are known to possess fin-winged fruits. We present an inventory of these taxa and describe their morphological characters as an aid for the identification and phylogenetic assessment of fossil and extant genera. Such fruits are most prevalent among Eudicots, but occur occasionally in Magnoliids (Hernandiaceae: Illigera) and Monocots (Burmannia, Dioscorea, Herreria). Although convergent in general form, fin-winged fruits of different genera can be distinguished by details of the wing number, texture, shape and venation, along with characters of persistent floral parts and dehiscence mode. Families having genera with fin-winged fruits and epigynous perianth include Aizoaceae, Apiaceae, Araliaceae, Asteraceae, Begoniaceae, Burmanniaceae, Combretaceae, Cucurbitaceae, Dioscoreaceae, Haloragaceae, Lecythidiaceae, Lophopyxidaceae, Loranthaceae, and Styracaceae. Families with genera having fin-winged fruits and hypogynous perianth include Achariaceae, Brassicaceae, Burseraceae, Celastraceae, Cunoniaceae, Cyrillaceae, Fabaceae, Malvaceae, Melianthaceae, Nyctaginaceae, Pedaliaceae, Polygalaceae, Phyllanthaceae, Polygonaceae, Rhamnaceae, Salicaceae sl, Sapindaceae, Simaroubaceae, Trigoniaceae, and Zygophyllaceae. This survey has facilitated the identification of fossil winged fruits such as Combretaceae and Araliaceae in the late Cretaceous of western North America and provides additional evidence toward the identification of various Cenozoic fossils including Brassicaceae, Fabaceae, Polygonaceae, Rutaceae, and Sapindaceae.