Change in diversity,ecological significance and biogeographical relationships of the Mediterranean Miocene toothed whale fauna

The main evolutionary trend in the Mediterranean Miocene toothed whale fauna is related (1) to the change in diversity and (2) to the turnover in community structure. Diversity increases from Upper Aquitanian–Lower Burdigalian to Burdigalian–Langhian, when it reaches its maximum. Starting from this time, diversity decreases progressively. The Early Miocene (Upper Aquitanian–Lower Burdigalian) Mediterranean toothed whale fauna, as well as the extramediterranean ones, is characterised by a high number of endemic taxa and by the prevalence of longirostral forms living in estuarine-neritic environments. A more diversified fauna spreading in neritic and pelagic environments characterises the Burdigalian–Langhian age, while an increase in pelagic forms and the nearly complete disappearance of some archaic longirostral taxa is typical of the Serravallian–Messinian fauna. Decrease in diversity and disappearance of archaic longirostral taxa are also recorded, at more general scale, in the Late Miocene extramediterranean fossil bearing deposits. These events can be related to the progressive global climatic deterioration, starting from Middle Miocene. From a biogeographic point a view, we can outline some relationships between the Mediterranean and western North Atlantic Miocene faunas. Closer affinities are observed between the Baltringen fauna and the northern Atlantic one, because of the presence of the genera Pomatodelphis and Zarhachis (platanistids) in both areas. In the Miocene Mediterranean and in North Atlantic, the delphinids are apparently absent as well as other extant delphinoid groups even if erroneously recorded in the past.     

Neogene history of Sporolithon Heydrich (Corallinales,Rhodophyta) in the Mediterranean region

Most modern species of Sporolithon live in tropical and subtropical areas and only one species of the genus, S. ptychoides, occurs in the Mediterranean Sea. The scarce present-day populations of Sporolithon in the Mediterranean region are relics of a long history of the genus in this area since the Early Cretaceous. Analysis of data from the palaeontological literature, combined with the study of both fossil samples and Recent ones collected from Italy and Spain, shows that during the Miocene variations in the number of Sporolithon species in the Mediterranean region parallel changes in global temperature. After a maximum species richness in the Langhian (early Mid Miocene), coincident with the Miocene climatic optimum, the number of species decreased to just two before the Messinian Salinity Crisis. This marked decline follows the global cooling event that began at around 14 Ma. The closure of the connections of the Mediterranean region with the Indian Ocean during the Langhian left Mediterranean Sporolithon populations isolated from the main dispersal area of the genus. After the Messinian desiccation, a single species, S. ptychoides, re-invaded the Mediterranean Basin during the Early Pliocene and continues to inhabit this temperate sea today. The Atlantic Ocean is the most probable source of the re-invading Sporolithon plants.     

Ostracodes et stratigraphie du néogène et du quaternaire méditerranéens

A stratigraphical chart of marine ostracoda from Lower Miocene to Recent is established. Selected species (approximatively 220) are those morphologicaly well characterized and known from different parts of the Mediterranean area. It appears that: • lower Miocene ostracodes are still poorly known; • specific diversity is high during the Tortonian and the Lower Messinian before the complete disappearance of marine Mediterranean species during the Upper Messinian évaporitic episodes; • during the early Pliocene, about half of the Upper Miocene marine species are reintroduced with the Atlantic waters; other species migrate for the first time in the Mediterranean Sea by the same way; • at the end of the Pliocene or at the beginning of the Pleistocene several species known in Mediterranean since the Middle Miocene or before, such as Cytherella sp. gr. transversa and Ruggieria tetraptera, as well some “nordic guests” such as Hemicythere villosa and Cythere lutea, appear. This work is an opportunity to confirme a Late Miocene age for the Neogene of Skyros (Aegean Sea), to assign the “Upper Pliocene” of Terquem to the Lower Pleistocene and to refute the existence of a pliocene psychrosphere.     

Biogéographied'un groupe d'Échinides cénozoiques (Echinolampas et ses sous-genres Conolampas et Hypsoclypus)

Echinolampas is a subtropical genus living in rather shallow water; one may regards it as a climatic marker. The theory of continental drift affords a rather good explanation for its distribution in space during Cenozoic era. It appears in Old Wolrd during Paleocene and it occurs in Central America during Middle Eocene; that implies it had to cross the already broad Atlantic Ocean; but at that time this ocean is not as broad at it is now. Migration along the shelf area which rimmed North Atlantic might have been impossible, owing to disruption of land connection between Europe and North America. Probably the migration occured in low latitudes and pelagic larvae were transported by one of the two equatorial currents. Diversity of the genus has much decreased during Late Eocene. The cause may be chiefly due to climatic deterioration, resulting from marine communication between North Atlantic and Arctic Ocean. Echinolampas occurs for the first time in Australia during Oligocene. One may suggest the possibility of a link between this late evidence and the quite remote position till then of Australian continent. During Miocene, the relative decrease in Echinolampas diversity in the Mediterranean Basin occurs as a result of the welding between Asia and Africa.     

Upper Eocene to Middle Miocene ostracode faunas and paleo-oceanography of the North Coastal Belt,Jamaica, West Indies

The ostracode fauna of the Montpelier Group (Upper Eocene-Middle Miocene) exposed in the western part of the North Coastal Belt of Jamaica contains representatives of three paleo-environments. Allochthonous shelf and littoral species, characterized by abraded carapaces and eye tubercles, and a diverse group of archibenthal (slope) forms are present throughout. Species restricted to the World Ocean Psychrosphere (> 1000 m), notably Bradleya dictyon, Australoecia tipica, Agrenocythere hazelae and Macrocypris spp., first occur in sediments of earliest Miocene age (planktonic foraminiferal zone N4). Their presence establishes a minimum age for the entry into the Cayman Trench of frigid (<8–10°C) water masses drawn from the Atlantic thermohaline stratification.Upper Eocene and Oligocene assemblages are dominated by smooth, blind genera such as Krithe, Messinella and Bythocypris and lived in lower thermospheric conditions (10°C) at or near the base of the thermocline (700– 900 m). During the Early Miocene, downfaulting of the north island slope of Jamaica in response to southward tilting of the Nicaraguan Rise brought the Montpelier depositional site into contact with progressively deeper and colder water-masses. This subsidence culminated in the late Early Miocene; faunas of this age contain large numbers of Bradleya dictyon, Agrenocythere hazelae, Macrocypris sp. 1, Bairdia oarion and Bairdoppilata cassida. They suggest paleodepths of 1500–2000 m and the local presence of a 3–4° water-mass tentatively identified as the North Atlantic deep water. If this interpretation is correct, then the maximum sill-depth and water-mass configuration of the Miocene Cayman Trench have persisted to the present. Middle Miocene assemblages indicate a return to paleodepths of 1000–1500 m heralding the pronounced Upper Miocene to mid-Pleistocene tectonism which elevated Montpelier strata above sea level. An alternative view attributes the Miocene ostracode succession in the Montpelier to episodes in the development of the Atlantic circulation as recognized in the northeast Atlantic.Montpelier deposition took place either on a broad step intervening between the northern edge of the Clarendon Platform and the abyssal basins of the Cayman Trench or on a Tertiary analog of the existing north island slope. These results have emphasized the value of detailed taxonomic and morphologic studies of deep-sea ostracodes in geohistorical reconstruction.     

The Mediterranean hydrologic budget from a Late Miocene global climate simulation

During the Late Miocene the Mediterranean experienced a period of extreme salinity fluctuations known as the Messinian Salinity Crisis (MSC). The causes of these high amplitude changes in salinity are not fully understood but are thought to be the result of restriction of flow between the Mediterranean and Atlantic, eustatic sea level change and climate. Results from a new Atmospheric General Circulation Model (AGCM) simulation of Late Miocene climate for the Mediterranean and adjacent regions are presented here. The model, HadAM3, was forced by a Late Miocene global palaeogeography, higher CO₂ concentrations and prescribed sea surface temperatures. The results show that fluvial freshwater fluxes to the Mediterranean in the Late Miocene were around 3 times greater than for the present day. Most of this water was derived from North African rivers, which fed the Eastern Mediterranean. This increase in runoff arises from a northward shift in the intertropical convergence zone caused by a reduced latitudinal gradient in global sea surface temperatures. The northwards drainage of the Late Miocene Chad Basin also contributes. Numerical models designed to explore Late Miocene salt precipitation regimes in the Mediterranean, which typically make use of river discharge fluxes within a few tens of percent of present-day values, may therefore be grossly underestimating these fluxes.Although the AGCM simulated Late Miocene river discharge is high, the model predicts a smaller net hydrologic budget (river discharge plus precipitation minus evaporation) than for present day. We discuss a possible mechanism by which this change in the hydrologic budget, coupled with a reduced connection between the Mediterranean and the global ocean, could cause the salinity fluctuations of the MSC.     

Integrated stratigraphy and paleoceanography of the Messinian (latest Miocene) across the North Atlantic Ocean

The Messinian was a time of major climatic and paleoceanographic change during the late Cenozoic. It is well known around the Mediterranean region because of the giant anhydrite/gypsum sequence and the suggested desiccation of the Mediterranean Sea. However, this interval is less constrained outside the Mediterranean region, where several paleoceanographic changes could have taken place because of the desiccation. Hence, we present an integrated stratigraphic framework for future Messinian paleoceanographic studies, determination of the effect of the Mediterranean desiccation on deep-water paleoceanography, and comparison of intra-Mediterranean paleoceanographic changes with those in the open oceans during the Messinian Stage.Four DSDP/ODP Holes (552A, 646B, 608, and 547A) from the North Atlantic Ocean and one land borehole from Morocco have been studied to integrate bio-, magneto-, and stable isotope Messinian stratigraphy with high resolution sampling. Our results produce the best assessment of the Tortonian/Messinian boundaries in all holes because they do not rely on any one signal.In paleomagnetic Subchronozone CSAn.lr in the Salé borehole and DSDP Site 609, a S/D coiling direction change of Neogloboquadrina pachyderma/acostaensis appears to indicate PMOW entering the northeastern Atlantic Ocean, at least reaching 50°N. Diachrony and synchrony of some important Messinian planktic foraminifera from these Atlantic DSDP/ODP holes and the Salé borehole, such as the LO of Gq. dehiscens, the LO of Gt. lenguaensis, the FO and LO of Gt. conomiozea, the FO of Gt. margaritae s.s., the FO of Gt. puncticulata, and the FO of Gt. crassaformis are discussed for understanding some of the paleoceanographic changes. This integrated stratigraphie framework presented here allows much better North Atlantic correlations at this critical point in Messinian geologic history.     

Late Miocene paleoenvironmental changes in North Africa and the Mediterranean recorded by geochemical proxies (Monte Gibliscemi section,Sicily)

The astronomically tuned marls of the Monte Gibliscemi section, Sicily, constitute an archive to trace the late Miocene palaeoenvironmental conditions (~ 9.7–7.0 Ma) in North Africa. Here we have utilised carbonate content and Al-normalised geochemical proxies to trace changes in terrigenous source area and bottom-water ventilation. The terrigenous input into the section is dominated by North African river systems draining into the Eastern Mediterranean. The proxy parameters indicate that the palaeoenvironmental conditions in North Africa were humid from 9.5 Ma onward with high fluvial input to the Mediterranean. Increases in the Si/Al and Mg/Al ratios occurred from 8.4 to 8.2 Ma and from 8.05 to 7.75 Ma, with maximum values similar to those of Messinian diatomites. These peaks indicate conditions of enhanced biosiliceous productivity and the presence of authigenic clay formation. Sluggish water circulation in the Mediterranean during those times is inferred from the Mn/Al and V/Al behaviour. Late Miocene changes of the Betic (southern Spain) and Rifian (Morocco) Mediterranean–Atlantic gateways are interpreted as the driving force for the changes in water circulation and the transgression associated with the opening of the Rifian corridor is interpreted to occur at 7.8 Ma.     

Multilocus Bayesian Estimates of Intra-Oceanic Genetic Differentiation,Connectivity, and Admixture in Atlantic Swordfish (Xiphias gladius L.)

Previous genetic studies of Atlantic swordfish (Xiphias gladius L.) revealed significant differentiation among Mediterranean, North Atlantic and South Atlantic populations using both mitochondrial and nuclear DNA data. However, limitations in geographic sampling coverage, and the use of single loci, precluded an accurate placement of boundaries and of estimates of admixture. In this study, we present multilocus analyses of 26 single nucleotide polymorphisms (SNPs) within 10 nuclear genes to estimate population differentiation and admixture based on the characterization of 774 individuals representing North Atlantic, South Atlantic, and Mediterranean swordfish populations. Pairwise F _ST values, AMOVA, PCoA, and Bayesian individual assignments support the differentiation of swordfish inhabiting these three basins, but not the current placement of the boundaries that separate them. Specifically, the range of the South Atlantic population extends beyond 5°N management boundary to 20°N-25°N from 45°W. Likewise the Mediterranean population extends beyond the current management boundary at the Strait of Gibraltar to approximately 10°W. Further, admixture zones, characterized by asymmetric contributions of adjacent populations within samples, are confined to the Northeast Atlantic. While South Atlantic and Mediterranean migrants were identified within these Northeast Atlantic admixture zones no North Atlantic migrants were identified respectively in these two neighboring basins. Owing to both, the characterization of larger number of loci and a more ample spatial sampling coverage, it was possible to provide a finer resolution of the boundaries separating Atlantic swordfish populations than previous studies. Finally, the patterns of population structure and admixture are discussed in the light of the reproductive biology, the known patterns of dispersal, and oceanographic features that may act as barriers to gene flow to Atlantic swordfish.     

Introduced brown algae in the North East Atlantic, with particular respect toUndaria pinnatifida (Harvey) suringar

The recent introduction of the macroalgaUndaria pinnatifida (Harvey) Suringar into the North Atlantic is the latest of a large number of introductions, which have occurred over many years. Some have been deliberate introductions for mariculture or research, while most have been accidental, via vectors such as shipping and shellfish imports. Not all newly recorded species are introductions; some are thought to be merely extensions of distribution, e.g.Laminaria ochroleuca, while others may have been overlooked previously, e.g.Scytosiphon dotyi. Subsequent to its accidental introduction into the waters around the Mediterranean French coast at Sete, most likely with imported oysters,Undaria was deliberately introduced into the North Atlantic, to Brittany, in 1983 by IFREMER for commercial exploitation.Undaria has since spread from the original sites in Brittany, and is now established at several sites on the south coast of England. This paper discusses the introduced brown algae in the North Atlantic and outlines the establishment ofUndaria in the UK.     

Conservation genetics of the short-beaked common dolphin (<Emphasis Type="Italic">Delphinus delphis</Emphasis>) in the Mediterranean Sea and in the eastern North Atlantic Ocean

Mediterranean Sea common dolphins have recently been listed as ‘endangered’ in the IUCN Red list, due to their reported decline since the middle of the 20th century. However, little is know about the number or distribution of populations in this region. We analysed 118 samples from the Black Sea, Mediterranean Sea and eastern North Atlantic at nine microsatellite nuclear loci and for 428 bps of the mtDNA control region. We found small but significant population differentiation across the basin between the eastern and the western Mediterranean populations at both nuclear and mtDNA markers (microsatellite F _ST = 0.052, mtDNA F _ST = 0.107, P values ≤ 0.001). This matched the differential distribution and habitat use patterns exhibited by this species in the eastern and the western parts of the Mediterranean Sea. The assignment test of a small number of samples from the central Mediterranean could not exclude further population structure in the central area of the basin. No significant genetic differentiation at either marker was observed among the eastern north Atlantic populations, though the Alboran population (inhabiting the Mediterranean waters immediately adjacent the Atlantic ocean) showed significant mtDNA genetic differentiation compared to the Atlantic populations. Directional estimates of gene flow suggested movement of females out of the Mediterranean, which may be relevant to the population decline. Phylogenetic analysis suggested that the observed population structure evolved recently.     

New Occurrences of Flying Fishes <Emphasis Type="Italic">Cheilopogon melanurus</Emphasis> and <Emphasis Type="Italic">C. heterurus</Emphasis> (Exocoetidae) in the North Central Atlantic and Eastern Part of the Mediterranean Sea

Cheilopogon melanurus, found for the first time in the central part of the North Atlantic (26°34′6 N 33°55′2 W), and C. heterurus from the eastern part of the Mediterranean Sea (Mahmutlar, Alanya, Turkey) are described. C. heterurus is widely distributed in the eastern part of the Mediterranean Sea occurring here in all seasons.     

Natural or Naturalized? Phylogeography Suggests That the Abundant Sea Urchin Arbacia lixula Is a Recent Colonizer of the Mediterranean

We present the global phylogeography of the black sea urchin Arbacia lixula, an amphi-Atlantic echinoid with potential to strongly impact shallow rocky ecosystems. Sequences of the mitochondrial cytochrome c oxidase gene of 604 specimens from 24 localities were obtained, covering most of the distribution area of the species, including the Mediterranean and both shores of the Atlantic. Genetic diversity measures, phylogeographic patterns, demographic parameters and population differentiation were analysed. We found high haplotype diversity but relatively low nucleotide diversity, with 176 haplotypes grouped within three haplogroups: one is shared between Eastern Atlantic (including Mediterranean) and Brazilian populations, the second is found in Eastern Atlantic and the Mediterranean and the third is exclusively from Brazil. Significant genetic differentiation was found between Brazilian, Eastern Atlantic and Mediterranean regions, but no differentiation was found among Mediterranean sub-basins or among Eastern Atlantic sub-regions. The star-shaped topology of the haplotype network and the unimodal mismatch distributions of Mediterranean and Eastern Atlantic samples suggest that these populations have suffered very recent demographic expansions. These expansions could be dated 94–205 kya in the Mediterranean, and 31–67 kya in the Eastern Atlantic. In contrast, Brazilian populations did not show any signature of population expansion. Our results indicate that all populations of A. lixula constitute a single species. The Brazilian populations probably diverged from an Eastern Atlantic stock. The present-day genetic structure of the species in Eastern Atlantic and the Mediterranean is shaped by very recent demographic processes. Our results support the view (backed by the lack of fossil record) that A. lixula is a recent thermophilous colonizer which spread throughout the Mediterranean during a warm period of the Pleistocene, probably during the last interglacial. Implications for the possible future impact of A. lixula on shallow Mediterranean ecosystems in the context of global warming trends must be considered.     

Palaeoceanographic record of the last deglaciation in the Strait of Sicily

The palaeoceanographic evolution of the Levantine waters during the last deglacial time is investigated using the sedimentary record of a deep sea core, CS 70-5, from the Linosa basin (35° 44.4′N/13° 11.0′E, 1486 m water depth). Radiocarbon dating and oxygen isotope stratigraphy based on¹⁸O changes inGlobigerina bulloides allow us to recognize and to date the different steps of the deglaciation. These steps are synchronous with those reported in the North Atlantic, but correspond to a δ¹⁸O decrease of higher amplitude than in the Alboran sea or in the North Atlantic Ocean. Major faunal events permit the establishment of a local biozonation which differs from those reported either for the Alboran sea or the Eastern Mediterranean basin. Major breaks in the faunal assemblages occurred during Termination I_A, within the first step of the deglaciation around 14 kyr B.P., and near 10.6 kyr B.P. within the Younger Dryas. The onset of the last deglaciation induced important changes in the characteristics of the Levantine and Atlantic water masses which occupied the Strait of Sicily. The δ¹³C records ofGlobigerina bulloides andCibicidoides pachyderma indicate that the modifications observed in the assemblages of deep faunas are controlled by the oxygenation of the water column. δ¹³C records ofGlobigerina bulloides are similar throughout the West Mediterranean as well as in sediments located below the present Mediterranean outflow. The distributional pattern as well as its δ¹³C record suggest that this species could be a good recorder of the upper Levantine waters and, more precisely, of the mixing layer of the overlying Atlantic waters with the Levantine ones.Major influxes of Atlantic waters during Terminations I_A and I_B could have slowed down the vertical mixing of the different water layers present in the Strait of Sicily and caused a decrease in the oxygenation of the water column. During Termination I_B the effect of Atlantic influxes was reinforced by the occurrence of a low salinity layer in the eastern basin which led to the stagnation of the deep waters. The two episodes of decreased oxygenation in the Levantine waters also favored the precipitation of inorganic magnesium calcite.     

New Nuclear SNP Markers Unravel the Genetic Structure and Effective Population Size of Albacore Tuna (Thunnus alalunga)

In the present study we have investigated the population genetic structure of albacore (Thunnus alalunga, Bonnaterre 1788) and assessed the loss of genetic diversity, likely due to overfishing, of albacore population in the North Atlantic Ocean. For this purpose, 1,331 individuals from 26 worldwide locations were analyzed by genotyping 75 novel nuclear SNPs. Our results indicated the existence of four genetically homogeneous populations delimited within the Mediterranean Sea, the Atlantic Ocean, the Indian Ocean and the Pacific Ocean. Current definition of stocks allows the sustainable management of albacore since no stock includes more than one genetic entity. In addition, short- and long-term effective population sizes were estimated for the North Atlantic Ocean albacore population, and results showed no historical decline for this population. Therefore, the genetic diversity and, consequently, the adaptive potential of this population have not been significantly affected by overfishing.     

EVOLUTIONARY RELATIONSHIPS BETWEEN FOUR SPECIES OF CLADOPHORA (CLADOPHORALES,CHLOROPHYTA) BASED ON DNA–DNA HYBRIDIZATION1

Analysis of the reassociation kinetics of the DNA from Cladophora pellucida (Huds.) Kütz. indicates that the genome of this benthic alga is comprised of approximately 75% repetitive sequences. Single-copy sequences reassociated with a rate constant of 1.8 × 10^?3 M^?1· s^?1, which corresponds to a haploid genome size of 4.7 × 10⁸ bp. Genotypic relationships between members of the form section Longiarticulatae were determined by the method of DNA–DNA hybridization. No significant divergence was observed between the single-copy sequences of C. pellucida isolates from the East Atlantic coast and Mediterranean Sea. Cladophora feredayi Harv. and C. att. ad pellucida from Australia and C. pellucidoidea van den Hoek from the West Atlantic coast were highly and about equally divergent from C. pellucida. The data support the hypothesis that the West Atlantic–West Pacific divergence reflects the middle Miocene closure of the Mediterranean–Indo-Pacific seaways, and the hypothesis that the Northwest Atlantic–Northeast Atlantic divergence reflects the middle Miocene thermal separation of these coasts.     

New data reinforcing the taxonomic status of Lepidion eques as synonym of Lepidion lepidion (Teleostei,Gadiformes)

The Mediterranean morid codling Lepidion lepidion is thought to be endemic, yet its taxonomic distinctiveness from the morphologically similar and more wide-ranging Atlantic Lepidion eques is unresolved, and has been controversial since the beginning of the twentieth century. Despite the abundant taxonomic literature questioning the interspecific relationship between these taxa, their current status remains unchanged. To elucidate the differentiation of the specimens identified as L. lepidion and L. eques collected across much of their geographic ranges, the sequence divergence of the cytochrome oxidase I “DNA barcode” gene of the mitochondrial genome was evaluated. A network analysis indicates that the most observed haplotypes are common to both species throughout their Mediterranean and North Atlantic distribution areas. This molecular evidence suggests the absence of biogeographical barriers and is insufficient to support the different species designations, giving L. eques the taxonomic status of junior synonym of L. lepidion.     

Miocene bristlemouths (Teleostei: Stomiiformes: Gonostomatidae) from the Makrilia Formation,Ierapetra, Crete

Bristlemouths of the genus Cyclothone are currently regarded as the most abundant vertebrates on Earth. The fossil record seems to suggest that these fishes diversified during the Miocene in the Pacific Ocean, but there is no evidence of their presence in the Miocene of the Atlantic Ocean and Mediterranean basin. A new bristlemouth, Cyclothone gaudanti sp. nov. (Teleostei, Stomiiformes, Gonostomatidae), is described herein based on 16 specimens from the Upper Miocene Makrilia Formation (late Tortonian of Crete, Greece). The small sized species is characterized by light pigmentation, 30–31 (14–15 + 15–16) vertebrae, dorsal fin with 10–13 rays, anal fin with 10–14 rays, premaxilla bearing seven closely spaced teeth, maxilla with 42–55 teeth, epipleurals, and autogenous parhypural. The presence of epipleurals appears to be unique of this Miocene species, and the re-establishment of this ancestral character state may be possibly interpreted as related to a phylogenetic character reversal. Morphological and paleoecological considerations suggest that this species possibly inhabited the upper mesopelagic layer, at depths ranging from 2–300 and 500 meters.     

Evolution of the Madrean–Tethyan disjunctions and the North and South American amphitropical disjunctions in plants

Abstract The present paper reviews advances in the study of two major intercontinental disjunct biogeographic patterns: (i) between Eurasian and western North American deserts with the Mediterranean climate (the Madrean–Tethyan disjunctions); and (ii) between the temperate regions of North and South America (the amphitropical disjunctions). Both disjunct patterns have multiple times of origin. The amphitropical disjunctions have largely resulted from long‐distance dispersal, primarily from the Miocene to the Holocene, with available data indicating that most lineages dispersed from North to South America. Results of recent studies on the Mediterranean disjuncts between the deserts of Eurasia and western North America support the multiple modes of origin and are mostly consistent with hypotheses of long‐distance dispersal and the North Atlantic migration. Axelrod's Madrean–Tethyan hypothesis, which implies vicariance between the two regions in the early Tertiary, has been favored by a few studies. The Beringian migration corridor for semiarid taxa is also supported in some cases.