Embryonic shell structure of Early–Middle Jurassic belemnites,and its significance for belemnite expansion and diversification in the Jurassic

Early Jurassic belemnites are of particular interest to the study of the evolution of skeletal morphology in Lower Carboniferous to the uppermost Cretaceous belemnoids, because they signal the beginning of a global Jurassic–Cretaceous expansion and diversification of belemnitids. We investigated potentially relevant, to this evolutionary pattern, shell features of Sinemurian–Bajocian Nannobelus, Parapassaloteuthis, Holcobelus and Pachybelemnopsis from the Paris Basin. Our analysis of morphological, ultrastructural and chemical traits of the earliest ontogenetic stages of the shell suggests that modified embryonic shell structure of Early–Middle Jurassic belemnites was a factor in their expansion and colonization of the pelagic zone and resulted in remarkable diversification of belemnites. Innovative traits of the embryonic shell of Sinemurian–Bajocian belemnites include: (1) an inorganic–organic primordial rostrum encapsulating the protoconch and the phragmocone, its non‐biomineralized component, possibly chitin, is herein detected for the first time; (2) an organic rich closing membrane which was under formation. It was yet perforated and possessed a foramen; and (3) an organic rich pro‐ostracum earlier documented in an embryonic shell of Pliensbachian Passaloteuthis. The inorganic–organic primordial rostrum tightly coating the protoconch and phragmocone supposedly enhanced protection, without increase in shell weight, of the Early Jurassic belemnites against explosion in deep‐water environment. This may have increased the depth and temperature ranges of hatching eggs, accelerated the adaptation of hatchlings to a nektonic mode of life and promoted increasing diversity of belemnoids. This study supports the hypothesis that belemnite hatchlings were ‘a miniature of the adults’.     

Larval and early post-larval shell of three deep-sea Scaphopoda (Mollusca) from the southwest Atlantic

The larval shells of Antalis circumcincta (Watson, 1879) (order Dentaliida), Pertusiconcha callithrix (Dall, 1889) (order Gadilida) and of an undetermined species of uncertain systematic position are described. The material studied comprises mainly samples from deep waters, collected by expeditions along the southeast coast of Brazil. The larval shell of the three taxa matches other types previously described in the literature. Antalis circumcincta and P. callithrix have four regions (protoconch A, protoconch B, teleoconch A, teleoconch B), but differ in dimensions and sculpture from each other, while the undetermined species has three regions (protoconch A, protoconch B, teleoconch B). A morphometric approach combined with a discriminant analysis also indicates that the three taxa are significantly distinct. This study confirms patterns of larval shells at the taxonomic rank of orders but other supraspecific patterns remain uncertain.     

New Paleocene Sepiid Coleoids (Cephalopoda) from Egypt: Evolutionary Significance and Origin of the Sepiid ‘Rostrum’

New coleoid cephalopods, assignable to the order Sepiida, are recorded from the Selandian/Thanetian boundary interval (Middle to Upper Paleocene transition, c. 59.2 Ma) along the southeastern margin (Toshka Lakes) of the Western Desert in Egypt. The two genera recognised, Aegyptosaepia n. gen. and ?Anomalosaepia Weaver and Ciampaglio, are placed in the families Belosaepiidae and ?Anomalosaepiidae, respectively. They constitute the oldest record to date of sepiids with a ‘rostrum-like’ prong. In addition, a third, generically and specifically indeterminate coleoid is represented by a single rostrum-like find. The taxonomic assignment of the material is based on apical parts (as preserved), i.e., guard, apical prong (or ‘rostrum-like’ structure), phragmocone and (remains of) protoconch, plus shell mineralogy. We here confirm the shell of early sepiids to have been bimineralic, i.e., composed of both calcite and aragonite. Aegyptosaepia lugeri n. gen., n. sp. reveals some similarities to later species of Belosaepia, in particular the possession of a distinct prong. General features of the phragmocone and protoconch of the new form are similar to both Belocurta (Middle Danian [Lower Paleocene]) and Belosaepia (Eocene). However, breviconic coiling and the presence of a longer ventral conotheca indicate closer ties with late Maastrichtian–Middle Danian Ceratisepia. In this respect, Aegyptosaepia n. gen. constitutes a link between Ceratisepia and the Eocene Belosaepia. The occurrence of the new genus near the Selandian/Thanetian boundary suggests an earlier origin of belosaepiids, during the early to Middle Paleocene. These earliest known belosaepiids may have originated in the Tethyan Realm. From northeast Africa, they subsequently spread to western India, the Arabian Plate and, probably via the Mediterranean region, to Europe and North America.     

Usefulness of the opercular nucleus for inferring early development in neritimorph gastropods

The early ontogeny of gastropods (i.e., planktotrophic vs. nonplanktotrophic) may be inferable from the morphology of the protoconch in adult shells. The protoconch consists of both embryonic and larval shells in species with planktotrophic development; the embryonic shell forms in the intracapsular period and the succeeding larval shell gradually develops during the larval period. In nonplanktotrophic species, on the other hand, there is no additional growth of the larval shell and the protoconch consists exclusively of a relatively large embryonic shell formed prior to hatching. This "shell apex theory" has been applied to many species of shell-bearing gastropods, but biotic and abiotic erosion of the apex often prevents detailed examination of the protoconch and subsequent inferences about ontogeny. I examined the gastropod operculum to test its utility for predicting developmental mode, drawing on the Neritimorpha as model taxa. Most aquatic members of Neritimorpha were found to bear an operculum with a clearly demarcated nucleus; SEM observations reveal four types of nuclei, which correspond to different types of protoconch morphologies and observed ontogenies for the study species. The nucleus is secreted before metamorphosis, fits into the shell aperture of the larva, and reflects early ontogeny as morphology, as does the protoconch. Moreover, the apparently organic (rather than calcareous) composition of the nucleus makes it nearly invulnerable to erosion and very advantageous, compared to the protoconch, in this ecologically diverse group, whose habitats range from freshwater streams and mangrove swamps to rocky shores and deep-sea hydrothermal vents. The measurements of the nucleus are also valuable for taxonomic purposes, especially in the species identification of veliger larvae and juvenile snails. On the other hand, the opercular nuclei of the Caenogastropoda and Heterobranchia are often eroded away in adult individuals; even if present, the morphology of the nuclei does not seem to clearly reflect early ontogeny in those groups.     

Palaeoenvironmental significance of Miocene larger benthic foraminifera from the Xisha Islands,South China Sea

We studied 344 samples from Well XK-1 in Xisha Islands, South China Sea, and identified 66 species of larger benthic foraminifera, providing critical evidence for biostratigraphy and palaeoenvironmental interpretation of the Miocene reef carbonate sequence. Three assemblages are recognized, namely, Spiroclypeus higginsi–Borelis pygmaeus Assemblage (Letter Stage Te5, Early Miocene, 1256.28–1180.15 m), Nephrolepidina–Miogypsina Assemblage (Tf, Middle Miocene, 1031.10–577.04 m), and Cycloclypeus–Heterostegina Assemblage (Tg, Late Miocene, 468.13–380.42 m). On the basis of the palaeoecological preference of the larger foraminifera, we interpret that the Miocene carbonate sequence was deposited mainly in a warm tropical shallow water environment, characterized by five stages of continuous long-term evolution: backreef lagoon to shelf in the Early Miocene, normal to frontal reef in the early Middle Miocene, backreef lagoon to shelf in the later Middle Miocene, normal to frontal reef in the early Late Miocene, and proximal forereef shelf in the later Late Miocene.     

Lepadiform and scalpelliform barnacles from the Oligocene and Miocene of the Paratethys sea

Abstract: A review on the Oligocene and Miocene stalked barnacles of the Paratethys Sea is presented. The fauna comprises two lepadiform and eight scalpelliform species. Only two species are known from Oligocene deposits, four are documented from Lower Miocene formations, and another four species occur in the Middle Miocene. Only one Oligocene and one Miocene species occur in two different Paratethyan basins in roughly coeval environments, and only a single Middle Miocene species is also known from the adjacent Mediterranean Sea. This low degree of biogeographical similarity is probably a result from the still low number of publications dealing with this group. Stalked barnacles are often indicating bathyal or at least deep sublittoral environments. Especially, the abundance of the benthic deep‐water lepadiform Poecilasma in evaporitic deposits of the Middle Miocene Badenian Salinity Crisis may serve as new evidence for a deeper marine depositional environment. Scalpellum paratethyianum sp. nov. and Lepas sattmanni sp. nov. are described as new species; Arcoscalpellum renevieri (Mayer and Gümbel in Gümbel, 1861) is introduced as new combination.     

Dynamic diversification history with rate upshifts in Holarctic bell‐flowers (Campanula and allies)

Campanula s.l. is one of the most speciose flowering plant lineages of the Holarctic (ca. 600 species). In the present study we sequenced three regions of the plastid genome (petD, rpl16 and trnK/matK) across a broad sample of Campanula s.l., which markedly improved phylogenetic resolution and statistical support compared to previous studies. Based on this robust phylogenetic hypothesis we estimated divergence times using BEAST, diversification rate shifts using Bayesian Analysis of Macroevolutionary Mixture (BAMM) and TreePar, and ancestral ranges using Biogeography with Bayesian (and likelihood) Evolutionary Analyses in R. Campanula s.l. is estimated to have originated during the Early Eocene but the major diversification events occurred between the Late Oligocene and Middle Miocene. Two upward diversification rate shifts were revealed by BAMM, specific to the crown nodes of two Campanula clades: CAM17, a mostly South European‐SW Asian lineage originating during the Middle Miocene and containing nearly half of all known Campanula species; and CAM15B, a SW Asian–Sino‐Himalayan lineage of nine species originating in the early Pleistocene. The dynamic diversification history of Campanula and the inferred rate shifts are discussed in a geo‐historical context.     

Distinct protoconchs recognised in three of the larger Mediterranean Cerithium species (Caenogastropoda: Cerithiidae)

The gastropod genus Cerithium includes several polymorphic species which are hardly distinguishable using a morphological approach based on teleoconch characters. Here we show that protoconch characters can be reliably used to identify the larger Mediterranean species (Cerithium alucastrum, C. repandum and C. vulgatum), and to assess their intraspecific variability. Based on a large amount of morphological data, we show that a multispiral, strongly sculptured protoconch (traditionally associated with C. vulgatum) is found in C. alucastrum. This species originated in the Pliocene. A multispiral, weakly sculptured protoconch, not observed previously, is reported for C. vulgatum. A paucispiral protoconch, distinct from those observed previously for C. lividulum and C. renovatum, is reported for C. repandum and can be used for recognising its wide intraspecific variation and understanding its geographical distribution. Cerithium repandum, previously doubtfully recorded outside its type locality, extends back to the mid-Pliocene and is today rather widely distributed in the Mediterranean, although absent from the Aegean Sea and the Alboran Sea. We describe the larval development of C. repandum and C. vulgatum for the first time, confirming their respective non-planktotrophic and planktotrophic development previously inferred from protoconch morphology. Also, we report a possible case of morphological convergence between C. repandum and C. vulgatum occurring in harbour sites, where we found a distinct (possibly Anthropocene) shell form not described previously from other Recent or fossil material.     

Plankton stratigraphy of the Early and Middle Miocene Kareem and Rudeis Formations in the central part of the Gulf of Suez,Egypt

The planktonic foraminifera and nannofossils of three wells in the Gulf of Suez penetrating the Early to Middle Miocene Upper Rudeis and Kareem Formations are attributed (from top to base) to the Middle Miocene Globorotalia peripheroronda Partial Range Zone (M6), the earliest Middle Miocene Praeorbulina sicana–Orbulina suturalis Interval Zone (M5), subdivided into the Praeorbulina glomerosa s. strict.–O. suturalis Interval Subzone (M5b) and the P. sicana–P. glomerosa s. str. Interval Subzone (M5a) and the Early Miocene Globigerinoides bisphericus Partial Range Subzone (M4b). The appearance of O. suturalis at the base of Subzone M5b represents the final stage of evolution of the Globigerinoides trilobus–Praeorbulina–Orbulina Lineage. In addition, the calcareous nannoplankton assemblages indicate the Sphenolithus heteromorphus Zone (NN5) and the Helicosphaera ampliaperta Zone (NN4). These biozones are well correlatable with those established by El-Heiny and Martini (1981, Geol Mediterr. Tome, VIII(2): 101–108) from the southwestern flank of the Gulf of Suez.     

New data on molluscs and their shell microstructures from the Middle Cambrian Gowers Formation, Australia

Abstract: Numerous new cases of preserved shell microstructure were discovered in molluscs from the Middle Cambrian Gowers Formation (Ptychagnostus atavus/Peronopsis opimus Zone, Floran Stage) in the Georgina Basin, Australia. The new data provide further evidence that, by the Middle Cambrian, molluscan shell microstructures were diverse, and many molluscs had a complex shell with multiple types of shell microstructure. In addition, many new occurrences of laminar microstructures are described herein. For many, the nature of these laminar microstructures is not known, but in three species the microstructure is foliated calcite, and in at least two the microstructure is more likely to have been calcitic semi‐nacre, a type of microstructure known in brachiopods and bryozoans but unknown in modern molluscs. This commonality among these three closely related lophotrochozoans underscores a similar mechanism of biomineralization. Moreover, these observations suggest a prevalence of calcite‐shelled lineages among molluscs from the Middle Cambrian, a time of calcite seas. In addition, the broad occurrence of laminar, nacre‐like microstructures in many of these fossils reveals how widespread these strong (fracture‐resistant) microstructures were in Middle Cambrian molluscs. Additionally, a few specimens of Yochelcionella preserve imprints of a bilaterally symmetrical pair of muscle scars. New taxa described here include Corystos thorntoniensis gen. et sp. nov., Yochelcionella snorkorum sp. nov., Yochelcionella saginata sp. nov., and Anhuiconus? agrenon sp. nov.     

A new late Miocene Lymnaea with aberrant suture structure unique for the family (Gastropoda: Pulmonata: Lymnaeidae)

A new fossil species of the pond snails (family Lymnaeidae Rafinesque, 1815), Lymnaea bogatschevi sp. n., from the Neogene (Khersonian–Maeotian transition) of southern Russia is described. It is characterised by a peculiar structure of the shell suture not found elsewhere in the family Lymnaeidae (both in living and extinct taxa). The suture of L. bogatschevi sp. n. may be described as a narrow groove that penetrates deep into the shell matter. This feature is developed both in protoconch and teleoconch whorls and is presented in several 100s of specimens that precludes its origin as an individual abnormality. There are no environmental factors known to induce such groove-like sutures in aquatic pulmonates indicating a genetic basis for this structure in L. bogatschevi. Also, we report a single shell with the similar channelled suture from a sample of Lymnaea megarensis (Gaudry in Fischer, 1867) from the Pliocene of Greece. It seems, in this species the groove-like suture was represented as a rare individual aberration, and later this trait disappeared completely from the lymnaeid morphological repertoir. Possible phylogenetic relationships between L. bogatschevi and the recent L. (Kazakhlymnaea) taurica (Clessin, 1880) are suggested and briefly discussed.     

Provannid and provannid‐like gastropods from the Late Cretaceous cold seeps of Hokkaido (Japan) and the fossil record of the Provannidae (Gastropoda: Abyssochrysoidea)

A fauna of provannid and provannid‐like shells is described from Upper Cretaceous seep carbonates in Hokkaido, Japan. We describe two new provannid species, Provanna tappuensis sp. nov. and Desbruyeresia kanajirisawensis sp. nov. , with preserved protoconchs of unquestionable provannid type with decollate apex. This material confirms the occurrence of Provannidae as early as the Middle Cenomanian. We also describe Hokkaidoconcha gen. nov. and a new family Hokkaidoconchidae fam. nov. , with two named species, H. hikidai sp. nov. and H. tanabei sp. nov . Hokkaidoconchidae are possibly related to the Provannidae, judging from a similar, but not decollate larval shell, although the juvenile teleoconch whorls differ in being of a general cerithimorph appearance and the details of the aperture are unknown. Furthermore, we review the published fossil record of Provannidae and Abyssochrysidae, and we consider that in those older than the Eocene, there is no evidence preserved that unequivocally supports a position there. The Jurassic Acanthostrophia acanthica from Italy seems to be the oldest known record of Abyssochrysidae, and the most reliable occurrence of the family, older than from the Miocene. Other fossil, pre‐Miocene species that have been classified in the Abyssochryssidae are provisionally referred to Hokkaidoconchidae. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154 , 421–436.     

Ontogenetic Torsion and Protoconch Form in the Archaeogastropod Haliotis kamtschatkana: Evolutionary Implications

Ontogenetic torsion in archaeogastropods has strongly influenced theories about early gastropod evolution, but the seminal studies by Crofts (1937, 1955) remain the major source of information about tissue movements during this developmental process. Computer-generated reconstructions of histological sections indicate that the cephalopodium of Haliotis kamtschatkana Jonas, 1845 rotates by a full 180° relative to the shell and visceral lobe during the first quarter of pre-metamorphic development. However, a portion of pallial epithelium, including some of the shell field, accompanies the rotating cephalopodium; a process facilitated by detachment of the pallium from the apertural rim of the protoconch. Transmission electron microscopy indicates that a tract of the larval retractor muscle, which Crofts (1955) implicated in the mechanism of torsion, inserts on both pedal and pallial cells. A deep invagination of shell field epithelium is a major focus of rotational torque. As a result of pallial deformation during cephalopodial rotation, the anus and gill rudiment are restricted to the right half of the larval body for 2 days after cephalopodial rotation by 180° has been completed. Scanning and transmission electron microscopy indicate that grooves in the lateral flanks of the protoconch correspond to the deep invagination of shell field epithelium. The grooves are not created by a coiling type of accrelionary shell growth or by flexion of the protoconch. A calcareous shelf is secondarily added to the periostracal template of the protoconch along its visceral apertural rim. Morphogenetic movements during ontogenetic torsion in this species are more complex than a simple rotation between cephalopodium and visceropallium and the protoconch shows no evidence of exogastric coiling. © 1997 Published by Elsevier Science Ltd on behalf of The Royal Swedish Academy of Sciences.     

Endocochliate embryo model in the Mesozoic Ammonitida

An endocochliate embryo model for the Mesozoic Ammonitida is proposed based on scanning electron microscopy of the ammonitella (= embryonic shell) stage of well‐preserved Japanese Cretaceous specimens belonging to nine species of five superfamilies. As in other specimens described previously, the ammonitella wall succeeding from the initial chamber ("protoconch") in the species examined consists of the inner prismatic, middle subprismatic and outer prismatic layers, with minute tubercles resting on the outer. Developmental patterns of these structures and their comparison with primary shell formation in modern Nautilus and Spirula suggest that the outer thin prismatic layer with microtubercles was secreted by the exterior epithelium after the completion of the main ammonitella wall by the interior shell gland. Thus, the early ammonite embryo might have had an endocochliate structural plan as in coleoids, and at the time of hatching the overlying mantle epithelium had shifted anteriorly to become an ectocochliate condition.     

Evolution of an embryo

In fossil material from two welis from Java, one section from East Java and two sections from Madura, specimens belonging to the extinet genusLepidocyclina (Foraminifera) were examined. Attention was paid to the embryonic part of the megalospheric forms exclusively. The size of the protoconch and deuteroconch was measured, but in particular also their spatial relation as determined by the extent to which the latter encloses the former, or the extent to which the dividing-wall between them is curved (“degree of curvature”). This parameter was found to increase persistently from 10% in the Middle Oligocene to 67% in the Middle Miocene. The rate of evolution showed to be inconstant, changing from very slow to very fast to slow again and to fast again. The first period of rapid change coincides with the time (Middle Oligocene) during which the genus migrated from America to Europe and to the Far East and Australia. The second phase of rapid evolution took place from the beginning of the Miocene up to the moment of dying out of the genus. The progressive change of this parameter is considered from the point of view of genetic variation and selection.     

Zygopleuroidea (Gastropoda) aus dem Lias und Dogger Deutschlands und Nordwestpolens

16 taxa of gastropods are described from the Lower and Middle Jurassic of Germany and northwestern Poland. They belong to seven genera. Two species (Pommerozygia aspera, Costazygia bilzi) and two genera (Brevizygia, Costazygia) are new. The family Pommerozygiidae is new as well. Compared to the Zygopleuridae, the Pommerozygiidae have a rather short and broad shell with only few teleoconch whorls. The protoconch is broad conical with a rounded apex (because the first whorls are nearly planispiral). From the Zygopleuridae only members of the Zygopleurinae have been found. Most Jurassic species have a smooth protoconch. Within the Zygopleuridae, the development possibly began with protoconchs carrying collabral axial ribs (and spirals) and led to smooth protoconchs. The genera of the Pommerozygiidae are rather similar to each other. The planktotrophic larval shell has a subsutural row of nodes like many species of Triassic Zygopleuridae. Therefore, both families are closely related. The Pommerozygiidae are possibly a separate branch of the Zygopleuroidea without descendants. The main branch is probably the evolutionary line Zygopleuriidae — Janthinoidea.     

Taxonomic attribution of the La Grive hominoid teeth

The two hominoid teeth—a central upper incisor (NMB G.a.9.) and an upper molar (FSL 213981)—from the Middle Miocene site of La Grive‐Saint‐Alban (France) have been traditionally attributed to Dryopithecus fontani (Hominidae: Dryopithecinae). However, during the last decade discoveries in the Vallès‐Penedès Basin (Spain) have shown that several hominoid genera were present in Western Europe during the late Middle Miocene. As a result, the attribution of the dryopithecine teeth from La Grive is not as straightforward as previously thought. In fact, similarities with the upper incisor of Pierolapithecus have led to suggestions that either the latter taxon is present at La Grive, or that it is a junior synonym of Dryopithecus. Here, we re‐describe the La Grive teeth and critically revise their taxonomic assignment based on metrical and morphological comparisons with other Middle to Late Miocene hominoids from Europe and Turkey, with particular emphasis on those from the Vallès‐Penedès Basin. Our results suggest that the I¹ differs in several respects from those of Pierolapithecus and Hispanopithecus, so that an attribution to either Dryopithecus or Anoiapithecus (for which this tooth is unknown) seems more likely. The molar, in turn, most likely corresponds to the M¹ of a female individual. Compared to other Middle Miocene taxa, its occlusal morphology enables its distinction from Pierolapithecus, whereas relative crown height agrees well with Dryopithecus. Therefore, based on available evidence, we support the traditional attribution of the La Grive hominoid to D. fontani. Am J Phys Anthropol 151:558–565, 2013. © 2013 Wiley Periodicals, Inc.     

‘Out‐of‐Africa’ dispersal of tropical floras during the Miocene climatic optimum: evidence from Uvaria (Annonaceae)

Aim African–Asian disjunctions are common in palaeotropical taxa, and are typically explained by reference to three competing hypotheses: (1) ‘rafting’ on the Indian tectonic plate, enabling Africa‐to‐Asia dispersal; (2) migration via Eocene boreotropical forests; and (3) transoceanic long‐distance dispersal. These hypotheses are tested using Uvaria (Annonaceae), which is distributed in tropical regions of Africa, Asia and Australasia. Recent phylogenetic reconstructions of the genus show a clear correlation with geographical provenance, indicating a probable origin in Africa and subsequent dispersal to Asia and then Australasia. Ancestral areas and migration routes are inferred and compared with estimates of divergence times in order to distinguish between the prevailing dispersal hypotheses. Location Palaeotropics. Methods Divergence times in Uvaria are estimated by analysing the sequences of four DNA regions (matK, psbA–trnH spacer, rbcL and trnL–F) from 59 Uvaria species and 77 outgroup species, using a Bayesian uncorrelated lognormal (UCLD) relaxed molecular clock. The ancestral area of Uvaria and subsequent dispersal routes are inferred using statistical dispersal–vicariance analysis (s‐diva ). Results Uvaria is estimated to have originated in continental Africa 31.6 Ma [95% highest posterior density (HPD): 38.4–25.1 Ma] between the Middle Eocene and Late Oligocene. Two main migration events during the Miocene are identified: dispersal into Madagascar around 17.0 Ma (95% HPD: 22.3–12.3 Ma); and dispersal into Asia between 21.4 Ma (95% HPD: 26.7–16.7 Ma) and 16.1 Ma (95% HPD: 20.1–12.1 Ma). Main conclusions Uvaria fruits are widely reported to be consumed by primates, and are therefore unlikely candidates for successful long‐distance transoceanic dispersal. The other biogeographical hypotheses, involving rafting on the Indian tectonic plate, and dispersal via the European boreotropical forests associated with the Eocene thermal maximum, can be discounted due to incongruence with the divergence time estimates. An alternative scenario is suggested, involving dispersal across Arabia and central Asia via the tropical forests that developed during the late Middle Miocene thermal maximum (17–15 Ma), associated with the ‘out‐of‐Africa’ dispersal of primates. The probable route and mechanism of overland dispersal between Africa and Asia for tropical plant groups during the Miocene climatic optimum are clarified based on the Uvaria data.