EARLIEST MEGAFOSSIL EVIDENCE OF FAGACEAE: PHYLOGENETIC AND BIOGEOGRAPHIC IMPLICATIONS

Fossil evidence of Fagaceae from the Paleocene/Eocene boundary of western Tennessee is described and discussed. These fossils include a newly discovered pistillate inflorescence and dispersed fruits of subfamily Castaneoideae as well as a taxon that resembles modern trigonobalanoids (pistillate inflorescences and dispersed mature fruits). Fossil staminate catkins with fagaceous pollen, which we suggest may be conspecific with the trigonobalanoid infructescences, are also found at the locality. Two distinct types of fagaceous leaves are present at the locality. The reproductive structures are the oldest megafossils unequivocally assignable to Fagaceae and represent the oldest remains of subfamily Fagoideae and the oldest megafossil remains of Castaneoideae. In addition, the fossils provide insights into the chronology of diversification, biogeography, and phylogeny of Fagaceae. The trigonobalanoid remains may also provide insights into the timing and circumstances of the evolution of wind pollination in Fagaceae.     

Fossils of the Fagaceae and their implications in systematics and biogeography

The fossil history of the Fagaceae from China and its systematic and biogeographic implications are discussed based on revisionary studies of the fossil records. No creditable macrofossil record of the Fagaceae exists in the Cretaceous deposits and all the Cretaceous microfossil reports remain equivocal and require further study. The Paleocene fossils show the appearance and diversification of the two groups corresponding to the subfamilies Fagoideae and Castaneoideae sensu Nixon. By the Eocene, all modern genera had been present. The oldest fagaceous fossils represent subfamily Fagoideae with affinities to the extant genus Trigonobalanus. The leaf fossil genus Berryophyllum, with affinities to Quercus subg. Cyclobalanopsis, has been documented by the early Eocene and might have occurred earlier than other fossils assignable to Quercus. The appearance of evergreen sclerophyllous Ouercus with entire leaves might have occurred earlier than those with toothed leaves. Deciduous, urticoid-leaved oak fossils (Quercus subg. Quercus sect. Quercus) had not appeared until the Miocene. Fossil equivalents of Trigonobalanus, Castanopsis and Lithocarpus had occurred in Europe and North America by the early Tertiary, suggesting that continuous distributions were achieved via the northern hemisphere land bridges. Three groups of evergreen sclerophyllous oaks of apparent close phylogenetic relationships occurred in the Hengduan mountains, the Mediterranean area and northwestern North America. Their fossil forms have become dominant elements of those vegetation zones since the Miocene. A shared fossil history indicates a possible biogeographic boundary formed by the ancient Mediterranean. The evidence suggests that the oaks might arrive in North America during two distinct geologic periods: evergreen sclerophyllous entire-leaved oaks appeared by the Early Tertiary, whereas thedeciduous oaks with urticoid leaves appeared in the Late Tertiary.     

壳斗科的地质历史及其系统学和植物地理学意义

在收集整理现有壳斗科化石资料的基础上,讨论了壳斗科及其各属的起源时间、地史分布和地史 演替过程以及这些化石资料在系统学和植物地理学上的意义。白垩纪尚无壳斗科可靠的大化石记录, 微化石需要进一步研究才能确定亲缘关系以及古新世壳斗科已经分化出两个类群。从以上这些事实推 论壳斗科起源于白垩纪晚期,而壳斗科现代各属出现的时间应不晚于古新世。最早发现的壳斗科化石和现代栗亚科和水青冈亚科在形态结构上非常相似,这一事实表明,壳斗科分为两个亚科的观点更接近客观事实。在水青冈亚科中,三棱栎类的化石最早出现;在栎属中,青冈亚属更接近祖先类群;在地史中全缘栎类较具齿栎类出现早,粗齿的落叶栎类出现最晚。三棱栎属、栲属和石栎属的化石在老第三纪出现于北美和欧洲的事实说明,北美、欧洲和东亚在老第三纪时有一个相通的壳斗科植物区系。南美的三棱栎是通过北美进入南美的。中国横断山、欧洲地中海沿岸和北美西北部有一类形态特征相似、亲缘关系相近的硬叶栎类,它们之间有相同的地质演替历史,它们现代分布边界可能就是古地中海的边界。美洲的栎类有两个来源,常绿硬叶栎类是通过古地中海沿岸而经北美-欧洲陆桥到达的,落叶栎类则是在中新世以后通过白令海峡到达的。     

Two new fossil flowers of magnoliid affinity from the Late Cretaceous of New Jersey

Two taxa of cupulate magnoliid fossil flowers, Cronquistiflora and Detrusandra, are described from the Late Cretaceous (Turonian, ∼90 million years before present [MYBP]) Raritan (or lower Magothy) Formation of New Jersey. The fossil taxa are represented by flowers at various stages of development, associated fragments of cup-shaped floral receptacles with attached anthers, and isolated anthers. Both taxa have laminar stamens with adaxial thecae and valvate dehiscence. Pollen is boat-shaped and foveolate in anthers associated with Cronquistiflora and spherical with reticulate ornamentation in Detrusandra. Cup-shaped receptacles are externally bracteose in both taxa. The receptacle of Cronquistiflora is broader than the campanulate one of Detrusandra. Cronquistiflora also has more carpels (∼50 in a spiral vs. ∼5 in a whorl or tight spiral). In Detrusandra the carpels are surrounded by dorsiventrally flattened structures (pistillodes?) that are remote from the attachment of the stamens near the distal rim of the receptacular cupule. Detrusandra stigmas are rounded and bilobed, while those of Cronquistiflora, although bilateral in symmetry, are somewhat peltate. The fossil taxa share prominent characters with extant cupulate magnoliids (e.g., Eupomatia, Calycanthus), but also share characters with other magnoliids including Winteraceae. These fossils represent taxa that are character mosaics relative to currently recognized families. Inclusion of these fossils in existing data matrices and ensuing phylogenetic analyses effect changes in tree topologies consistent with their mosaicism relative to modern taxa. But such analyses do not definitively demonstrate the affinities of the fossils other than illustrating that these fossils are generalized magnoliids. Additional analysis of modern and fossil magnoliids is necessary to fully appreciate the phylogenetic significance and positions of these fossil taxa. However, the results of the phylogenetic analyses do introduce the possibility that extinct taxa of Magnoliales with cupulate floral receptacles were transitional between basal angiosperms and those with tricolpate pollen. The fossils provide insights into the timing of evolution of character complexes now associated with coleopteran pollination.     

The earliest fossil evidence of the Hamamelidaceae: Late Cretaceous (Turonian) inflorescences and fruits of Altingioideae

Fossilized pistillate inflorescences, fruits, and pollen grains from the Turonian (～90 million years before present) of New Jersey are described as a new genus, Microaltingia, in the family Hamamelidaceae. The fossils are remarkably preserved in exceptional detail. Several morphological and anatomical characters suggest affinities with Hamamelidaceae. These include capitate inflorescences, florets with a hypanthium, two-carpellate gynoecia, perigynous flowers, tricolpate reticulate pollen, a three-layered carpel wall, scalariform perforation plates with oblique end walls, and scalariform and opposite/alternate intervascular pitting. The gross morphology of pistillate inflorescences, unisexual flowers, phyllome structure, numerous ovules per carpel, and mode of carpel dehiscence indicate affinities with subfamily Altingioideae, which includes the modern genera Liquidambar and Altingia. Cladistic analysis using a previously published morphological matrix and scoring the fossil for available characters supports the position of the fossil as a sister taxon of modern Altingioideae. Although the fossil exhibits a mosaic of characters found within modern Hamamelidaceae, it is not identical to any modern taxon. Based on cladistic analysis, the fossil appears to be a basal "altingioid" that lacks the derived pollen found in extant Altingioideae and retains the more plesiomorphic tricolpate pollen found in the rest of Hamamelidaceae. The floral characters of the fossils, including phyllomes with stomata, short and straight styles, and small perprolate pollen grains, also indicate the possibility of insect pollination.     

Elucidating the affinities and habitat of ancient, widespread Cyperaceae: Volkeria messelensis gen. et sp. nov., a fossil mapanioid sedge from the Eocene of Europe

The sedges (family Cyperaceae) are an economically and ecologically important monocot group dating back at least to the Paleocene. While modern genera are mostly unknown before the Oligocene, several extinct taxa are recognized as the earliest sedges. Their affinities have been unclear until now, because they are found as isolated, often abraded fruits or endocarps. Exceptionally preserved sedge fossils from the Middle Eocene of Messel, Germany yield more characters for identification. Fossil cyperacean infructescences with in situ pollen are recognized for the first time and show features of the early-divergent mapanioid sedges. Pollen resembles that of tribe Hypolytreae. Comparisons with extant taxa suggest the closest affinities with Hypolytrum and Mapania. However, the Messel fossils represent a distinct taxon, Volkeria messelensis gen. et sp. nov. Without the additional characters of infructescence and pollen, the Messel fruits would have been placed in the extinct genus Caricoidea, a typical Eocene sedge that was widespread across Eurasia. Similarities of fruit structure suggest that Caricoidea was also a mapanioid sedge. Mapanioid sedges are found today in tropical wet forests and swamps, a distribution suggesting that early sedges occupied a similar habitat, unlike many modern sedges, and were not precursors to open grassland vegetation.     

PSEUDOFAGUS IDAHOENSIS,N. GEN. ET SP. (FAGACEAE) FROM THE MIOCENE CLARKIA FLORA OF IDAHO

A new genus and species of Fagaceae are established for compressed leaves and attached fruits which represent one of the dominant taxa in the rich Miocene Clarkia flora of northern Idaho. The leaf and fruit morphology, cuticular and pericarp anatomy, and leaf phytoliths fall within the range of variation of the subfamily Fagoideae. The unique fruit consists of a large, exposed, distinctly keeled, trigonous nut and a diminutive basal cupule. The cupule has an apical frill of free, petiolate, leaflike appendages but lacks other ornamentation, and there is no evidence of a distinct Valvation. As no other genus of the Fagaceae has fruits with this combination of characters, a new genus is established and is referred to the subfamily Fagoideae. The Miocene genus seems most closely related to the extant genus Fagus.     

Sclerosperma fossils from the late Oligocene of Chilga,north-western Ethiopia

The palm family, Arecaceae, is notoriously depauperate in Africa today, and its evolutionary, paleobiogeographic, and extinction history there are not well documented by fossils. In this article we report the pollen of two new extinct species of the small genus, Sclerosperma (Arecoideae), from a late Oligocene (27–28 Ma) stratum exposed along the Guang River in Chilga Wereda of north-western Ethiopia. The pollen are triporate, and the two taxa can be distinguished from each other and from modern species using a combination of light and scanning electron microscopy, which reveals variations in the finer details of their reticulate to perforate exine sculpture. We also report a palm leaf fragment from a stratum higher in the same section that is in the Arecoideae subfamily, and most likely belongs to Sclerosperma. The implications of these discoveries for the evolutionary history of this clade of African arecoid palms is that their diversification was well underway by the middle to late Oligocene, and they were much more widespread in Africa at that time than they are now, limited to West and Central Africa. Sclerosperma exhibits ecological conservatism, as today it occurs primarily in swamps and flooded forests, and the sedimentology of the Guang River deposits at Chilga indicate a heterogeneous landscape with a high water table. The matrix containing the fossil pollen is lignite, which itself indicates standing water, and a variety of plant macrofossils from higher in the section have been interpreted as representing moist tropical forest or seasonally inundated forest communities.     

Triuridaceae fossil flowers from the Upper Cretaceous of New Jersey

We report here on a series of fossil flowers exhibiting a mosaic of characters present in the extant monocot family Triuridaceae. Phylogenetic analyses of morphological data from a broad sample of extant monocots confirm the affinities of the fossils with modern Triuridaceae. The fossil flowers were collected from outcrops of the Raritan Formation (Upper Cretaceous, ～90 million years before present), New Jersey, USA. These are the oldest known unequivocal monocot flowers. Because other reports of "earliest" monocots are all based on equivocal character suites and/or ambiguously preserved fossil material, the Triuridaceae fossils reported here should also be considered as the oldest unequivocal fossil monocots. Flowers are minute and unisexual (only male flowers are known); the perianth is composed of six tepals, lacking stomata. The unicyclic androecium is of three stamens with dithecal, monosporangiate, extrorse anthers that open by longitudinal slits. The endothecium has U-shaped type thickenings. Pollen grains are monosulcate. The triurid fossil flowers can be separated into three distinctive species. On the basis of phylogenetic analyses of morphological characters, the fossil taxa nest within the completely saprophytic achlorophyllous Triuridaceae supporting the interpretation that the extinct plants were also achlorophyllous and saprophytic. If so, this represents the earliest known fossil occurrence of the saprophytic/mycotrophic habit in angiosperms.     

Marsileaceaephyllum, a new genus for marsileaceous macrofossils: leaf remains from the Early Cretaceous (Albian) of southern Gondwana

Three new taxa from Albian, Early Cretaceous assemblages in Gondwana (Australia and Antarctica) and two previously described fossils from the Late Cretaceous and Eocene of North America are attributable to the heterosporous semi-aquatic fern family Marsileaceae. They are assigned to Marsileaceaephyllum, a morphotaxon erected here for sterile remains (whole plants, and isolated leaves and leaflets) of Marsileaceae. The Gondwanan taxa, Marsileaceaephyllum lobatum and Marsileaceaephyllum spp. B-C, have either a cruciform leaflet arrangement or dichotomous and anastomosing venation characteristic of modern Marsileaceae. Two previously established taxa, Marsilea johnhallii and Marsilea sp., which represent sterile Marsileaceae, are also transferred to the new genus (now Marsileaceaephyllum johnhallii and Marsileaceaephyllum sp. A, respectively). Examination of all fossil venation patterns reveals four new venation types not present in extant taxa, suggesting that most fossil Marsileaceae (leaves) are distinct from extant genera, and are likely members of extinct lineages. This is further supported by the absence of modern megaspore types in the Early Cretaceous.     

Phylogeny of extant and fossil Juglandaceae inferred from the integration of molecular and morphological data sets

It is widely acknowledged that integrating fossils into data sets of extant taxa is imperative for proper placement of fossils, resolution of relationships, and a better understanding of character evolution. The importance of this process has been further magnified because of the crucial role of fossils in dating divergence times. Outstanding issues remain, including appropriate methods to place fossils in phylogenetic trees, the importance of molecules versus morphology in these analyses, as well as the impact of potentially large amounts of missing data for fossil taxa. In this study we used the angiosperm clade Juglandaceae as a model for investigating methods of integrating fossils into a phylogenetic framework of extant taxa. The clade has a rich fossil record relative to low extant diversity, as well as a robust molecular phylogeny and morphological database for extant taxa. After combining fossil organ genera into composite and terminal taxa, our objectives were to (1) compare multiple methods for the integration of the fossils and extant taxa (including total evidence, molecular scaffolds, and molecular matrix representation with parsimony [MRP]); (2) explore the impact of missing data (incomplete taxa and characters) and the evidence for placing fossils on the topology; (3) simulate the phylogenetic effect of missing data by creating "artificial fossils"; and (4) place fossils and compare the impact of single and multiple fossil constraints in estimating the age of clades. Despite large and variable amounts of missing data, each of the methods provided reasonable placement of both fossils and simulated "artificial fossils" in the phylogeny previously inferred only from extant taxa. Our results clearly show that the amount of missing data in any given taxon is not by itself an operational guideline for excluding fossils from analysis. Three fossil taxa (Cruciptera simsonii, Paleoplatycarya wingii, and Platycarya americana) were placed within crown clades containing living taxa for which relationships previously had been suggested based on morphology, whereas Polyptera manningii, a mosaic taxon with equivocal affinities, was placed firmly as sister to two modern crown clades. The position of Paleooreomunnea stoneana was ambiguous with total evidence but conclusive with DNA scaffolds and MRP. There was less disturbance of relationships among extant taxa using a total evidence approach, and the DNA scaffold approach did not provide improved resolution or internal support for clades compared to total evidence, whereas weighted MRP retained comparable levels of support but lost crown clade resolution. Multiple internal minimum age constraints generally provided reasonable age estimates, but the use of single constraints provided by extinct genera tended to underestimate clade ages.     

三棱栎的幼苗形态解剖及系统位置

对三棱栎幼苗的形态和解剖学研究表明,三棱栎幼苗与壳斗科其它属区别明显,并与水青冈属及南水青冈属较为接近,而与栎属关系甚远,最后提出了三棱栎属与水青冈属,南水青冈属共同组成水青冈亚科,为壳斗科中较原始的类群。     

Morphological and molecular evidence for a stepwise evolutionary transition from teeth to baleen in mysticete whales

The origin of baleen in mysticete whales represents a major transition in the phylogenetic history of Cetacea. This key specialization, a keratinous sieve that enables filter-feeding, permitted exploitation of a new ecological niche and heralded the evolution of modern baleen-bearing whales, the largest animals on Earth. To date, all formally described mysticete fossils conform to two types: toothed species from Oligocene-age rocks ( approximately 24 to 34 million years old) and toothless species that presumably utilized baleen to feed (Recent to approximately 30 million years old). Here, we show that several Oligocene toothed mysticetes have nutrient foramina and associated sulci on the lateral portions of their palates, homologous structures in extant mysticetes house vessels that nourish baleen. The simultaneous occurrence of teeth and nutrient foramina implies that both teeth and baleen were present in these early mysticetes. Phylogenetic analyses of a supermatrix that includes extinct taxa and new data for 11 nuclear genes consistently resolve relationships at the base of Mysticeti. The combined data set of 27,340 characters supports a stepwise transition from a toothed ancestor, to a mosaic intermediate with both teeth and baleen, to modern baleen whales that lack an adult dentition but retain developmental and genetic evidence of their ancestral toothed heritage. Comparative sequence data for ENAM (enamelin) and AMBN (ameloblastin) indicate that enamel-specific loci are present in Mysticeti but have degraded to pseudogenes in this group. The dramatic transformation in mysticete feeding anatomy documents an apparently rare, stepwise mode of evolution in which a composite phenotype bridged the gap between primitive and derived morphologies; a combination of fossil and molecular evidence provides a multifaceted record of this macroevolutionary pattern.     

Establishing a fossil record for the perianthless Piperales: Saururus tuckerae sp. nov. (Saururaceae) from the Middle Eocene Princeton Chert

Investigations of small permineralized flowers from the Middle Eocene Princeton Chert, British Columbia, Canada have revealed that they represent an extinct species of Saururus. Over 100 flowers and one partial inflorescence were studied, and numerous minute perianthless flowers are borne in an indeterminate raceme. Each flower is subtended by a bract, and flowers and bracts are borne at the end of a common stalk. Five stamens are basally adnate to the carpels. Pollen is frequently found in situ in the anthers. Examined under SEM and TEM, pollen grains are minute (6-11 μm), monosulcate, boat-shaped-elliptic, with punctate sculpturing and a granulate aperture membrane. The gynoecium is composed of four basally connate, lobed carpels with recurved styles and a single ovule per carpel. Flower structure and pollen are indicative of Saururaceae (Piperales), and in phylogenetic analyses using morphological characters, the fossils are sister to extant Saururus. The fossil flowers are described here as Saururus tuckerae sp. nov. These fossil specimens add to the otherwise sparse fossil record of Piperales, represent the oldest fossils of Saururaceae as well as the first North American fossil specimens of this family, and provide the first evidence of saururaceous pollen in the fossil record.     

Combined LM and SEM study of the middle Miocene (Sarmatian) palynoflora from the Lavanttal Basin,Austria: part IV. Magnoliophyta 2 – Fagales to Rosales

An ongoing investigation of the middle Miocene (Sarmatian) palynoflora from the Lavanttal Basin continues to show that it contains an extremely rich assemblage of angiosperm taxa. The Fagales to Rosales pollen record documented here contains 34 different taxa belonging to the Betulaceae (Alnus, Betula, Carpinus, Corylus, Ostrya), Fagaceae (Castanea, Fagus, Quercus Groups Cerris, Ilex, Cyclobalanopsis, Quercus/Lobatae), Juglandaceae (Engelhardioideae, Carya, Juglans, Pterocarya), Myricaceae (Morrella vel Myrica), Cannabaceae (Celtis), Elaeagnaceae (Elaeagnus), Rhamnaceae, Rosaceae (Prunus) and Ulmaceae (Cedrelospermum, Ulmus, Zelkova). Two of the pollen types represent extinct genera, Trigonobalanopsis and Cedrelospermum, and are also reported for the first time from the Lavanttal Basin along with pollen of Rhamnaceae and Prunus. The different types of Quercus pollen are now affiliated with Groups Cerris, Cyclobalanopsis, Ilex and Quercus/Lobatae based on sculpturing elements observed using scanning electron microscopy (SEM). Köppen signatures of potential modern analogues of the fossil Fagales and Rosales suggest a subtropical (Cfa, Cwa) climate at lower elevation and subsequent subtropical to temperate climate with altitudinal succession (Cfa → Cfb/Dfa→ Dfb; Cwa → Cwb → Dwb) in the Lavanttal area during accumulation of the palynoflora. Most of the fossil taxa have potential modern analogues that can be grouped as nemoral and/or merido-nemoral vegetation elements, and the diversity of Fagales indicates a varying landscape with a high variety of niches.     

INVESTIGATIONS OF TERTIARY ANGIOSPERMS: A NEW FLORA INCLUDING EOMIMOSOIDEA PLUMOSA FROM THE OLIGOCENE OF EASTERN TEXAS

A new fossil flower and inflorescence-bearing locality has been discovered in the Oligocene of the Texas Gulf Coast. The new flora is similar to the Middle Eocene Claiborne Flora of the southeastern USA, but the quality of preservation is sometimes better in the Oligocene fossils. One component of the new flora, a mimosoid legume inflorescence, appears identical with Eomimosoidea plumosa, first reported from the Claiborne Formation of western Tennessee. Investigations of these younger specimens indicate that the taxon had changed little during the Middle Eocene-Oligocene interval, and the better quality of preservation of the Texas specimens has provided further insights into the structure of the fossils. Comparisons of the fine structural details of the pollen of Eomimosoidea with similar pollen of extant mimosoids has confirmed that the fossil genus is indeed extinct and suggests that tetrahedral tetrads of columellate, tricolporate pollen grains are ancient, possibly primitive, in the Mimosoideae.     

The Pollen Morphology in Relation to the Taxonomy and Phylogeny of Fagaceae

The present paper deals with the pollen morphology of 103 species belonging to sixgenera-Castanea, Castanopsis, Lithocarpus, Quercus, Fagus and Trigonobalanus in threesubfamilies-Castaneoideae, Quercoideae and Fagoideae. All pollen grains were examinedunder light microscope and scanning electron microscope, and those of some species were ex-amined under transmission electron microscope. The results may be summarised as follows:1. Pollen morphology of Fagaceae, seems to support division of the family into threesubfamilies. Fagoideae, Castaneoideae and Quercoideae.2. Four types of pollen grains are recognized in Fagaceae:1) Fagus-type (representative genus: Fagus): pollen granis are oblate-sphaeroidal,(31.5-39.9) x (35.7 46.2) μm in size, 3(-4)-colporate, peritreme or goniotreme, granulate-ornate under LM., granulate or verrucate under SEM.2) Trigonobalanus-type (T. doichangensis): pollen grains are suboblate-sphaeroidal,(23.1-29.4) ×(25.2-29.4) μm in size, 3-colporate, goniotreme, obscurely granulate-ornateunder LM, densely granulate or verrucate under SEM.3) Quercus-type (Quercus): pollen grains are subspheroidal-subprolate, (21-44.3)× (16.8-39.9) μm in size. 3-colporoidate (-3-colpate), peritreme, crassgranulate or finely-gra-nulate under LM, tuberculate verrucate or spinate under SEM.4) Castanea-type (including Castanea, Castanopsis, Lithocarpus): Pollen grains areprolate-supraprolate, (14.7-23.1)×(10.5-16.8)μm in size; 3-colporate, peritreme, obscu-rely ornate or subpsilate, under LM, rugulose, striate-rugulate or crass-striate under SEM.3. Pollen grains of Cyclobalanopsis age very similar to those of Quercus, and there-fore we support the treatment of Cyclobalanopsis as a subgenus of Quercus.4. On the basis of shape, type of aperture and exine structure, pollen of Trigonobalanusis distinguishable from those of the other genera in Fagaceae and it may be a new type of Fagaceae;5. On the basis of pollen morphology, morphological characters and geological stratification a scheme of phylogeny of Fagaceae is here presented.     

Ancestral state reconstruction of body size in the Caniformia (Carnivora, Mammalia): the effects of incorporating data from the fossil record

A recent molecular phylogeny of the mammalian order Carnivora implied large body size as the ancestral condition for the caniform subclade Arctoidea using the distribution of species mean body sizes among living taxa. "Extant taxa-only" approaches such as these discount character state observations for fossil members of living clades and completely ignore data from extinct lineages. To more rigorously reconstruct body sizes of ancestral forms within the Caniformia, body size and first appearance data were collected for 149 extant and 367 extinct taxa. Body sizes were reconstructed for four ancestral nodes using weighted squared-change parsimony on log-transformed body mass data. Reconstructions based on extant taxa alone favored large body sizes (on the order of 10 to 50 kg) for the last common ancestors of both the Caniformia and Arctoidea. In contrast, reconstructions incorporating fossil data support small body sizes (< 5 kg) for the ancestors of those clades. When the temporal information associated with fossil data was discarded, body size reconstructions became ambiguous, demonstrating that incorporating both character state and temporal information from fossil taxa unambiguously supports a small ancestral body size, thereby falsifying hypotheses derived from extant taxa alone. Body size reconstructions for Caniformia, Arctoidea, and Musteloidea were not sensitive to potential errors introduced by uncertainty in the position of extinct lineages relative to the molecular topology, or to missing body size data for extinct members of an entire major clade (the aquatic Pinnipedia). Incorporating character state observations and temporal information from the fossil record into hypothesis testing has a significant impact on the ability to reconstruct ancestral characters and constrains the range of potential hypotheses of character evolution. Fossil data here provide the evidence to reliably document trends of both increasing and decreasing body size in several caniform clades. More generally, including fossils in such analyses incorporates evidence of directional trends, thereby yielding more reliable ancestral character state reconstructions.     

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.