A new species of Zelkova (Ulmaceae,Ulmoideae) with leaves and fruits from the Oligocene of South China and its biogeographical implications

A new fossil species of Zelkova is described from the Oligocene Ningming Formation of Guangxi, South China. Zelkova ningmingensis sp. nov. is characterized by leaves with craspedodromous venation pattern and drupaceous fruit type. The new species possesses elliptical to ovate leaves, bearing 7–12 pairs of secondary and simple toothed margin, as well as epidermal cells with straight or rounded anticlinal walls. The species is compared with extant and other fossil species hitherto reported of the genus. It is most similar to the living Zelkova schneideriana in the leaf gross morphology and epidermal characters, which may be suggested to be the ancestral type of Zelkova schneideriana. The discovery of Zelkova ningmingensis sp. nov. in Guangxi indicates that Zelkova has already existed in southern China as early as the Oligocene. In combination with Zelkova material from the Oligocene of Europe, it can be inferred that Eurasian Zelkova had begun to diversify by at least the Oligocene. Because China is the biodiversity centre of modern Zelkova, the fossil herein provides new insights into Zelkova biogeography.     

COMPARATIVE PALEOBIOCHEMISTRY OF SOME FOSSIL AND EXTANT FAGACEAE

A Fagus-like leaf fossil (cuticular compression) with an attached fruit, differing from any known Fagus species (fossil or extant) or other fagoid taxa, has been discovered from the Miocene Clarkia Lake deposits of northern Idaho. Because of its unusual morphology (especially the fruit) the fossil taxon has been described as a new genus and species, Pseudofagus idahoensis Smiley and Huggins. The successful previous use of paleobiochemistry in studies of fossil taxa from the Miocene Succor Creek Flora of Oregon suggested that chemical data might help clarify the taxonomic affinities of Pseudofagus. Indeed, examination of the chemistry of the fossil, Pseudofagus idahoensis, and comparison with extant Fagus species and related fagoid genera indicate that: 1) based on steroid chemistry, Pseudofagus idahonesis does belong in the Fagaceae; 2) like all extant species of Fagus, the fossil lacks the tannin component, ellagic acid, which separates it from other extant fagoid genera, and 3) its simple flavonoid pigment profile places it closest to the extant North American Fagus grandifolia or the European/Eurasian Fagus sylvatica. However, the exclusive presence of an isorhamnetin (3'-methoxyquercetin) 3-0-glycoside, onocerane, and 5α-cholestane imparts a species-specific chemical character to Pseudofagus idahoensis, which also sets it apart from extant species of Fagus. While the chemistry does not decide the taxonomic level to be accorded to the fossil, it certainly supports, along with morphology and anatomy, the distinctness of Pseudofagus and its proposed relationships within the Fagaceae.     

A Total Evidence Phylogeny of the Arctoidea (Carnivora: Mammalia): Relationships Among Basal Taxa

A total evidence phylogenetic analysis was performed for 14 extant and 18 fossil caniform genera using a data matrix of 5.6 kbp of concatenated sequence data from six independent loci and 80 morphological characters from the cranium and dentition. Maximum parsimony analysis recovered a single most parsimonious cladogram (MPC). The topology of the extant taxa in the MPC agreed with previous molecular phylogenies. Phylogenetic positions for fossil taxa indicate that several taxa previously described as early members of extant families (e.g., Bathygale and Plesictis) are likely stem taxa at the base of the Arctoidea. Taxa in the “Paleomustelidae” were found to be paraphyletic, but a monophyletic Oligobuninae was recovered within this set of taxa. This clade was closely related to the extant genera Gulo and Martes, therefore, nested within the extant radiation of the family Mustelidae. This analysis provides a resolution to several discrepancies between phylogenies considering either fossil taxa or extant taxa separately, and provides a framework for incorporating fossil and extant taxa into comprehensive combined evidence analyses.     

The fossil record of extant elasmobranchs

Sharks and their relatives (Elasmobranchii) are highly threatened with extinction due to various anthropogenic pressures. The abundant fossil record of fossil taxa has allowed the tracing of the evolutionary history of modern elasmobranchs to at least 250 MYA; nonetheless, exactly how far back the fossil record of living taxa goes has never been collectively surveyed. In this study, the authors assess the representation and extent of the fossil record of elasmobranchs currently living in our oceans by collecting their oldest records and quantifying first appearance dates at different taxonomic levels (i.e., orders, families, genera and species), ecological traits (e.g., body size, habitat and feeding mechanism) and extinction risks (i.e., threatened, not threatened and data deficient). The results of this study confirm the robust representation of higher taxonomic ranks, with all orders, most of the families and over half of the extant genera having a fossil record. Further, they reveal that 10% of the current global species diversity is represented in the geological past. Sharks are better represented and extend deeper in time than rays and skates. While the fossil record of extant genera (e.g., the six gill sharks, Hexanchus) goes as far back as c. 190 MYA, the fossil record of extant species (e.g., the sand shark, Carcharias taurus Rafinesque 1810) extends c. 66 MYA. Although no significant differences were found in the extent of the fossil record between ecological traits, it was found that the currently threatened species have a significantly older fossil record than the not threatened species. This study demonstrate that the fossil record of extant elasmobranchs extends deep into the geologic time, especially in the case of threatened sharks. As such, the elasmobranch geological history has great potential to advance the understanding of how species currently facing extinction have responded to different stressors in the past, thereby providing a deep-time perspective to conservation.     

INVESTIGATIONS OF ANGIOSPERMS FROM THE EOCENE OF NORTH AMERICA: A CATKIN WITH JUGLANDACEOUS AFFINITIES

Three specimens of one type of fossil catkin from the Middle Eocene of Tennessee are excellently preserved and have been investigated morphologically. The flowers on these catkins are subtended by elongate, three-lobed bracts, are exclusively staminate, and have three conspicuous, obovate, perianth parts that bear large peltate scales. The stamens are well preserved and contain triporate pollen grains that are equivalent to the dispersed pollen genus Momipites. Floral morphology, cuticular features, and pollen indicate close affinities with the extant genera Engelhardia, Oreomunnea, and Alfaroa of the Juglandaceae; but because the fossil catkins are distinct and are a dispersed plant organ, they are placed in a new form genus: Eokachyra. These fossil flowers represent a rare opportunity to correlate the micro- and macrofossil record and to compare the relative rates of evolution of these features. The fossil catkins also demonstrate that much structural information may be gained from the study of fossil angiosperm flowers. The similarities between the staminate flowers of the fossil catkins and the staminate flowers of Engelhardia, Oreomunnea, and Alfaroa confirm the idea that this complex has had a long evolutionary history and suggest that the pollination system of certain extant genera was well developed during Middle Eocene times.     

INVESTIGATIONS OF ANGIOSPERMS FROM THE EOCENE OF NORTH AMERICA: JUGLANDACEOUS WINGED FRUITS

The winged fruits of Engelhardia have been reported in many early Tertiary floras in the northern Hemisphere. Recent systematic revisions of the extant genera, Engelhardia, Oreomunnea, and Alfaroa provide a helpful guide to the study of the fossil fruits. Fruit form, venation, nut septations and flower parts are features that separate the extant Asian Engelhardia from the American Oreomunnea. Four types of fossil winged fruits can be recognized in Middle Eocene sediments of southeastern North America: Engelhardia, Paraoreomunnea gen. n., Paleooreomunnea gen. n., and Pararengelhardia. Both Paleooreomunnea and Paraengelhardia represent extinct lines; no living forms have similar winged fruits. The fossil fruits of Engelhardia and Paraoreomunnea are similar to the fruits of extant Engelhardia and Oreomunnea, respectively. The fossil taxa reported in this paper were sympatric. The present allopatric distribution of Engelhardia and Oreomunnea is the result of Neogene regional extinctions. Engelhardia and Oreomunnea have probably been distinct genera since early Paleogene time.     

Eocene handfishes from Monte Bolca,with description of a new genus and species,and a phylogeny of the family Brachionichthyidae (Teleostei: Lophiiformes)

The family Brachionichthyidae, commonly known as the handfishes, is a small group of lophiiform fishes, the living species of which are restricted in distribution mostly to shallow temperate and subtropical waters of Tasmania and southern and eastern Australia. Despite their narrow present‐day distribution, and the extreme rarity of lophiiforms in the fossil record, handfishes are well represented in the Eocene of Monte Bolca, Italy. A revision of the known fossil material shows the presence of two fossil species in two monotypic genera, ?Histionotophorus and ? Orrichthys gen. nov. Diagnoses of the family Brachionichthyidae, the two fossil genera, as well as two recognized extant genera Brachionichthys and Sympterichthys are provided. An osteological analysis of ?Histionotophorus bassani revealed many new features as well as reinterpretations of some previously described skeletal parts. A phylogenetic analysis of brachionichthyid genera and representatives of the antennarioid families Antennariidae, Tetrabrachiidae, and Lophichthyidae, using 36 morphological characters, strongly supported monophyly of brachionichthyids and antennarioids, the former taxon representing the sister group of the other families of the latter. Within the Brachionichthyidae, the two extant genera Brachionichthys and Sympterichthys form a species pair, as do the extinct genera ?Histionotophorus and ? Orrichthys gen. nov. Biogeographical considerations suggest that the present geographical range of handfishes can be considered a residual distribution of a temporally and spatially dynamic range shift. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 621–647.     

Evaluation of the fossil fish‐specific diversity in a chadian continental assemblage: Exploration of morphological continuous variation in Synodontis (Ostariophysi,Siluriformes)

In the fossil record, the quantification of continuous morphological variation has become a central issue when dealing with species identification and speciation. In this context, fossil taxa with living representatives hold great promise, because of the potential to characterise patterns of intraspecific morphological variation in extant species prior to any interpretation in the fossil record. The vast majority of catfish families fulfil this prerequisite, as most of them are represented by extant genera. However, although they constitute a major fish group in terms of distribution, and ecological and taxonomic diversity, the quantitative study of their past morphological variation has been neglected, as fossil specimens are generally identified based on the scarcest remains, that is, complete neurocrania that bear discrete characters. Consequently, a part of freshwater catfish history is unprospected and unknown. In this study, we explored the morphological continuous variation of the humeral plate shape in Synodontis catfishes using Elliptic Fourier Analysis (EFA), and compared extant members and fossil counterparts. We analysed 153 extant specimens of 11 Synodontis species present in the Chad basin, in addition to 23 fossil specimens from the Chadian fossiliferous area of Toros Menalla which is dated around 7 Ma. This highly speciose genus, which is one of the most diversified in Africa, exhibits a rich fossil record with several hundred remains mostly identified as Synodontis sp. The analysis of the outline of the humeral plate reveals that some living morphological types were already represented in the Chad Basin 7 My ago, and allows for the discovery of extinct species. Beside illuminating the complex Neogene evolutionary history of Synodontis, these results underline the interest in the ability of isolated remains to reconstruct a past dynamic history and to validate the relevance of EFA as a tool to explore specific diversity through time. J. Morphol. 277:1486–1496, 2016. © 2016 Wiley Periodicals, Inc.     

TRIPLOCHITIOXYLON (STERCULIACEAE): A NEW GENUS OF WOOD FROM THE EOCENE OF OREGON AND ITS BEARING ON XYLEM EVOLUTION IN THE EXTANT GENUS TRIPLOCHITON

A new sterculiaceous wood, Triplochitioxylon oregonensis gen. et sp. n., was collected from a Middle Eocene locality in the Clarno Formation of Oregon. Anatomical data indicate a close natural relationship between T. oregonensis and the living species of Triplochiton, a genus endemic to tropical Africa. The fossil is believed to represent a population of the group or complex from which Triplochiton evolved. The basic differences in the xylem organizations of the two genera are explained by a significant reduction of fusiform initial length and by a complete suppression of post-cambial parenchyma strand elongation in the extant genus. Paleobotanical and biogeographical evidence suggest that the xylem evolution has been strongly influenced by the increasing aridity of the African continent.     

On the phylogenetic position and systematics of extant and fossil Aclopinae (Coleoptera: Scarabaeidae)

The Aclopinae is a small subfamily within the family Scarabaeidae. It currently comprises five extant genera with 28 species, and eight fossil genera with 25 species. The systematic position of Aclopinae within the family Scarabaeidae is uncertain, particularly because representative species of Aclopinae have been absent in previous phylogenetic studies. Here we performed phylogenetic analyses using morphological and molecular data to investigate the phylogenetic position of fossil and extant Aclopinae. For this objective, we expanded and revised a former morphological data matrix (composed of 68 characters) including all extant genera of Aclopinae. We complemented our morphological investigations with a molecular phylogenetic analysis based on four genes of several extant taxa of Aclopinae and a wide sample of diverse Scarabaeoidea. Our phylogenetic analyses show that all the type species of the fossil genera formerly included within Aclopinae do not belong within the extant Aclopinae clade and support both the exclusion of those fossil taxa and the monophyly of the extant genera of Aclopinae: Aclopus Erichson, Desertaclopus Ocampo & Mondaca, Gracilaclopus Ocampo & Mondaca, Neophanaeognatha Allsopp and Phanaeognatha Hope. Our results also show that the fossil taxa Prophaenognatha robusta Bai et al. and Ceafornotensis archratiras Woolley are closely related to Ochodaeidae, while the remaining type species of fossils formerly included in Aclopinae (Cretaclopus longipes (Ponomarenko), Holcorobeus vittatus Nikritin, Juraclopus rodhendorfi Nikolajev, Mesaclopus mongolicus (Nikolajev), and Mongolrobeus zherikhini Nikolajev) belong to a distinct lineage closely related to Diphyllostomatidae. Based on these results, the subfamily Aclopinae appears monophyletic and sister to the ‘pleurostict’ lineage. Consequently, we propose the following changes to the current classification of the fossil taxa: Holcorobeus monreali (Gómez‐Pallerola) belongs to Carabidae (incertae sedis) as proposed by the original author, and we place Ceafornotensis Woolley, Cretaclopus Nikolajev, Holcorobeus Nikritin, Juraclopus Nikolajev, Mesaclopus Nikolajev, Mongolrobeus Nikolajev and Prophaenognatha Bai et al. in Scarabaeoidea (incertae sedis). Furthermore, we provide an identification key to, and diagnoses of, the genera, illustrations of diagnostic characters and checklists of their included species. The evolutionary perspective presented provides new insights into the evolution of the pleurostict condition in Scarabaeoidea and the biogeography of this group, which is now regarded as Gondwanan, probably evolving during the Cretaceous and not from the upper Jurassic as previously assumed.     

The postcranium of Miocene hominoids: Were dryopithecines merely “dental apes”?

This paper reviews the non-dental morphological configuration of Miocene hominoids with special reference to the hypothesis of linear relationships between certain fossil species and living analogues. Metrical analysis of the wrist shows thatDryopithecus africanus andPliopithecus vindobonensis are unequivocally affiliated with the morphological pattern of quadrupedal monkeys. Similar analyses of the fossil hominoid elbow shows that they are more cercopithecoid-like than hominoid-like. Multivariate analysis of theP. vindobonensis shoulder in the matrix of extant Anthropoidea indicate that this putative hylobatine fossil shows no indication of even the initial development of hominoid features. The total morphological pattern of theD. africanus forelimb as assessed by principal coordinates analysis of allometrically adjusted shape variables has little resemblance toPan. Likewise, the feet and proximal femora of the Miocene fossils are unlike any living hominoid species. Even theD. africanus skull is similar to extant cercopithecoids in several features. Although ancestors cannot be expected to resemble descendants in every way, the striking dissimilarity between Miocene and extant hominoids seems to eliminate the consideration of a direct ancestor-descendant relationship between specific Miocene and modern forms.     

INVESTIGATIONS OF ANGIOSPERMS FROM THE EOCENE OF NORTH AMERICA: STIPULATE LEAVES OF THE RUBIACEAE INCLUDING A PROBABLE POLYPLOID POPULATION

A comparative study was made of the gross morphology, fine venation and cuticular features of Leitneria fioridana Chapman, the single living representative of the order Leitneriales and Leitneria eocenica (Berry) Brown, presumbaly a related fossil species. In addition to the type material, newly collected fossil specimens were investigated from clay pits in the Middle Eocene, Claiborne Formation, of western Tennessee and Kentucky. Foliate stipules attached to the petioles of several specimens suggest the assignment of this fossil leaf type to the genus Leitneria is incorrect. The nature of the gross morphology, fine venation and cuticular features confirms the misidentification. Previously, various specimens of this fossil leaf type have been placed in eight species of seven genera in seven families of six angiosperm orders, none of which are correct systematically. The gross morphology, venation and cuticular characters of the fossil leaf are distributed among a few extant South American genera of arborescent Rubiaceae. The fossil is an extinct rubiaceous leaf type which cannot be placed within a single modern subfamily, tribe or genus of the family. The organ genus, Paleorubiaceophyllum is proposed for these leaves. Three varieties of a single fossil species, P. eocenicum, are recognized. One variety with epidermal cells nearly twice the size of the others may represent a polyploid population.     

Morphological phylogeny of gall‐forming aphids of the tribe Eriosomatini (Aphididae: Eriosomatinae)

Aphids of the tribe Eriosomatini are typically associated with the tree genera Ulmus and Zelkova as the primary host, on which they induce several types of leaf gall. To elucidate evolutionary changes in the aphid–host associations and gall morphology, phylogenetic relationships were inferred using 36 species (28 in the ingroup) and based on 52 morphological characters. Phylogenetic analysis with equal character weighting led to hundreds of most‐parsimonious trees, and the strict consensus of these was poorly resolved. However, the successive weighting of characters yielded three most‐parsimonious trees. The strict consensus of these supported the monophyly of the Eriosomatini and the monophyly of most genera. Reconstruction of the aphid–host associations on the consensus tree indicated that the ancestral Eriosomatini were associated with Zelkova and that the association with Ulmus evolved twice independently. Ancestral reconstruction suggests that galls of the leaf‐roll type are ancestral to those of the completely and incompletely closed pouch type, and that each type of pouch galls evolved independently. It is suggested that despite early diversification of the Eriosomatini on Zelkova species, Zelkova‐associated eriosomatines had become extinct or survived as relict parthenogens on the secondary host due to the elimination of Zelkova in most regions since the late Tertiary. In contrast, two large genera in the Eriosomatini, Eriosoma and Tetraneura, are associated with the largest elm section Ulmus whose elements are distributed widely in Eurasia, including boreal regions. Therefore, the available evidence suggests that the present species diversity and distribution patterns of the eriosomatines have been largely affected by the diversification and extinction of their host plants during the late Tertiary and Quaternary.     

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.     

Applications of Finite Element Analyses to Problems in Plant Morphology

Methods of describing the geometry and position of cells bymeans of the finite method are applied to selected extant (Fucus,Cutleria, and Calypogeia) and fossil plant genera (Protosaloinia,Spongiophyton, and Parka). The gross morphology and some aspectsof the ontogeny of these six genera are simulated by means ofcomputer; the pseudo- and parenchymatous tissue organizationsare mathematically characterized. Simulated representationsof living (Fucus and Cutleria) and fossil algae (Protosalcinia,Spongio phyton, and Parka) are shown to be distinctively differentfrom those of the bryophytes. The application of the finiteelement method to problems of plant morphology makes possiblethe dynamic simulations of some aspects of growth and developmentin living plants, and the extrapolation of missing stages inthe ontogeny of fossil plants.     

Phylogeny of the North American catfish family Ictaluridae (Teleostei: Siluriformes) combining morphology,genes and fossils

We performed the first combined‐data phylogenetic analysis of ictalurids including most living and fossil species. We sampled 56 extant species and 16 fossil species representing outgroups, the seven living genera, and the extinct genus ?Astephus long thought to be an ictalurid. In total, 209 morphological characters were curated and illustrated in MorphoBank from published and original work, and standardized using reductive coding. Molecular sequences harvested from GenBank for one nuclear and four mitochondrial genes were combined with the morphological data for total evidence analysis. Parsimony analysis recovers a crown clade Ictaluridae composed of seven living genera and numerous extinct species. The oldest ictalurid fossils are the Late Eocene members of Ameiurus and Ictalurus. The fossil clade ?Astephus placed outside of Ictaluridae and not as its sister taxon. Previous morphological phylogenetic studies of Ictaluridae hypothesized convergent evolution of troglobitic features among the subterranean species. In contrast, we found morphological evidence to support a single clade of the four troglobitic species, the sister taxon of all ictalurids. This result holds whether fossils are included or not. Some previously published clock‐based age estimates closely approximate our minimum ages of clades.     

Skull Shape, Masticatory Apparatus, and Diet of Vassallia and Holmesina (Mammalia: Xenarthra: Pampatheriidae): When Anatomy Constrains Destiny

The form and function of the masticatory apparatus of the fossil genera Vassallia and Holmesina are analyzed so that the possible dietary behaviors of these pampathere xenarthrans might be inferred. Analysis is based on comparisons of dental morphology and skeletal features (through RFTRA) associated with the masticatory musculature among the pampatheres, the extant dasypodids Euphractus and Dasypus, and the glyptodont Propalaeohoplophorus. A method is proposed for generating a moment arm of the massetericus independently of the muscle's line of action, which allows direct comparison among extant and fossil mammals. The masticatory apparatus of the pampatheres strongly resembles that of Euphractus among extant forms, but the development of muscular attachment sites indicates a more powerful musculature, particularly the massetericus; the taxa differ most markedly in dental morphology. Long moment arms about the jaw joint and large ratios of muscle to bite moments indicate forceful rather than quick movements. The various skeletal and dental features analyzed suggest that the masticatory apparatus of the pampatheres was more powerful and efficient in transverse chewing than in dasypodids and that they were primarily grazers consuming mainly coarse vegetation. These features, some shared with herbivorous ungulates, include wide, relatively flat mandibular condyles; condyles well dorsal to muscular insertion sites; expanded angular processes; unfused symphysis; a posteriorly extended tooth row; open-rooted teeth; mesial teeth that bear mainly transverse striations; distal teeth that are mesiodistally elongated, bear basined occlusal surfaces, and in Vassallia possess a central island of resistant dentine that acted as a functional analogue of an ectoloph; and teeth with a stepwise arrangement. The results of this study indicate that detailed analysis and comparison of morphology lead to useful predictions of behavior.     

Past and present diversity and distribution in the parasitic wasp family Megalyridae (Hymenoptera)

The phylogenetic relationships of extant and extinct Megalyridae are analysed at the genus level. The dataset comprises seven outgroup taxa, all eight extant genera and a number of extinct taxa that have been associated with Megalyridae, including two genera from Maimetshidae, whose affinity with Megalyridae is uncertain. Analytical results are unstable because some of the fossil taxa have many missing entries. The most stable results are produced when the maimetshid taxa and Cretodinapsis are excluded. When included, these taxa fall outside crown‐group Megalyridae, the maimetshid taxa being the sister of Orthogonalys (Trigonalidae). Based on the results of our analyses, we synonymize the fossil genera Rubes Perrichot n.syn . and Ukrainosa Perrichot & Perkovsky n.syn . with Prodinapsis, creating the new combinations Prodinapsis bruesi n.comb . and Prodinapsis prolata n.comb . When comparing past and present distributions of Megalyridae with the results of the phylogenetic analyses, it is evident that the genera radiated in the Mesozoic, and that the family as a whole was much more widespread then. The present‐day distribution is essentially relictual, with range contraction since the early Tertiary probably being the result of climate deterioration, which caused the disappearance of tropical forests throughout the Palaearctic.     

THE PLATYCARYA PERPLEX AND THE EVOLUTION OF THE JUGLANDACEAE

We report on the leaves, fruits, inflorescences, and pollen of two fossil species in the genus Platycarya. The association of these dispersed organs has been established by their repeated co-occurrence at a large number of localities, and for two of the organs (fruit and pistillate inflorescence, and pollen and staminate inflorescence) by apparent organic attachment of compression fossils. Each of the two species can be distinguished by characteristics of all the known megafossil organs. We also review the fossil record of dispersed platycaryoid fruits and inflorescences, recognizing three additional species of Platycarya and two of Hooleya. Two of the fossil Platycarya species are morphologically very different from the living Platycarya strobilacea Sieb. et Zucc., but they show the diagnostic features of the genus. Hooleya is a generalized member of the Platycaryeae that is probably close to the ancestry of Platycarya. The two Platycarya species known from multiple organs provide a remarkable example of mosaic evolution in which fertile and foliar structures have attained different levels of morphological specialization. The leaves, often considered the most plastic of plant organs, retain several features that are otherwise seen only in the Engelhardieae. These similarities in leaf architecture between the fossil Platycarya species and Engelhardieae are advanced features for the Juglandaceae, and thus indicate a sister-group relationship between the two lines. In contrast to the leaves, the fruits, inflorescences, and pollen of the fossil Platycarya species are almost as specialized as those of the extant P. strobilacea and bear little resemblance to the same structures in other genera of the family. The morphology, taphonomy, sedimentary setting, and geographic and stratigraphic distribution of three of the fossil platycaryoid species suggest that they were wind-dispersed, early successional plants that grew in thickets. This habit is retained by Platycarya strobilacea and is typical of many of the amentiferae (e.g. Myricaceae, Betulaceae). The r-selected life-history pattern of the Platycarya line may well have contributed to its low diversity through geologic time.