Elena Conti

Clades, Clocks, and Continents: Historical and Biogeographical Analysis of Myrtaceae, Vochysiaceae, and Relatives in the Southern Hemisphere

Some of the most interesting but still most contentious disjunct biogeographical distributions involve Southern Hemisphere tropical and warm temperate families. The PHMV clade of Myrtales includes four families (Psiloxylaceae, Heteropyxidaceae, Myrtaceae, and Vochysiaceae) that exhibit a number of these biogeographical patterns. The related Psiloxylaceae and Heteropyxidaceae are small families restricted in distribution to the recent volcanic Mascarene Islands to the east of Madagascar and to southeast Africa, respectively. Myrtaceae are found on three major Gondwanan regions (South America, Australasia, and Africa). Because the New World taxa are almost exclusively fleshy fruited, it is unclear whether the family distribution is a classic Gondwanan vicariance pattern or results from one or more long‐distance dispersal events over ocean barriers. The Vochysiaceae represent one of a handful of families with amphi‐Atlantic distributions vigorously argued to support both long‐distance dispersal over the Atlantic and vicariance of western Gondwanan biota by Atlantic seafloor spreading. Molecular phylogenetic relationships, fossil dating of nodes, and penalized likelihood rate smoothing of maximum likelihood trees were employed for a Myrtales‐wide analysis using rbcL and ndhF and an analysis of the PHMV analysis using ndhF and matK. The results indicate that the PHMV differentiated during the late Cretaceous. The African lineage of Vochysiaceae is nested within a South American clade and probably arose via long‐distance dispersal in the Oligocene at a time when the Atlantic had already rifted 80 m.yr. at the equatorial region. The African/Mascarene Island families, most closely related to Myrtaceae, differentiated during the late Eocene, with subsequent but recent long‐distance dispersal from Africa to the Mascarenes. Myrtaceae show a rapid differentiation of a basal, paraphyletic subf. Leptospermoideae in Australasia. Fleshy‐fruited taxa (subf. Myrtoideae) are not monophyletic. Vicariance of a widespread warm temperate Southern Hemisphere distribution is likely in explaining the South American–Australasian disjunction, with subsequent dispersal events between the two and to Africa and the Mediterranean basin.

EARLY TERTIARY OUT-OF-INDIA DISPERSAL OF CRYPTERONIACEAE: EVIDENCE FROM PHYLOGENY AND MOLECULAR DATING

Abstract Phylogenetic analyses and molecular dating estimates based on chloroplast DNA sequences were used to establish the relationships of the southern and Southeast Asian Crypteroniaceae and elucidate their biogeographic history. Maximum parsimony and likelihood analyses of rbcL sequences suggested that Crypteroniaceae should be restricted to Crypteronia, Axinandra, and Dactylocladus and that Crypteroniaceae, so defined, are sister to a clade formed by three small African taxa (Oliniaceae, Penaeaceae, and Rhynchocalycaceae) and the monotypic Central and South American Alzateaceae. Three molecular dating approaches (maximum-likelihood under a molecular clock, Langley-Fitch, and penalized-likelihood) were used to infer the age of Crypteroniaceae using both paleobotanic and geologic calibrations. Comparisons among these three methods revealed significant lineage effects in rbcL sequences. Clock-independent dating estimates suggested that divergence of Crypteroniaceae from its African and South American relatives coincided with the breakup of Gondwana, and that India likely served as a “raft” transporting Crypteroniaceae to Asia, with later expansion to Southeast Asia. To our knowledge, Crypteroniaceae are the first plant group for which the out-of-India hypothesis is well corroborated by molecular-based estimates of divergence times.

Assessing calibration uncertainty in molecular dating : The assignment of fossils to alternative calibration points

Although recent methodological advances have allowed the incorporation of rate variation in molecular dating analyses, the calibration procedure, performed mainly through fossils, remains resistant to improvements. One source of uncertainty pertains to the assignment of fossils to specific nodes in a phylogeny, especially when alternative possibilities exist that can be equally justified on morphological grounds. Here we expand on a recently developed fossil cross-validation method to evaluate whether alternative nodal assignments of multiple fossils produce calibration sets that differ in their internal consistency. We use an enlarged Crypteroniaceae-centered phylogeny of Myrtales, six fossils, and 72 combinations of calibration points, termed calibration sets, to identify (i) the fossil assignments that produce the most internally consistent calibration sets and (ii) the mean ages, derived from these calibration sets, for the split of the Southeast Asian Crypteroniaceae from their West Gondwanan sister clade (node X). We found that a correlation exists between s values, devised to measure the consistency among the calibration points of a calibration set (Near and Sanderson, 2004), and nodal distances among calibration points. By ranking all sets according to the percent deviation of s from the regression line with nodal distance, we identified the sets with the highest level of corrected calibration-set consistency. These sets generated lower standard deviations associated with the ages of node X than sets characterized by lower corrected consistency. The three calibration sets with the highest corrected consistencies produced mean age estimates for node X of 79.70, 79.14, and 78.15 My. These timeframes are most compatible with the hypothesis that the Crypteroniaceae stem lineage dispersed from Africa to the Deccan plate as it drifted northward during the Late Cretaceous.

Nucleotide Sequences of the rbcL Gene Indicate Monophyly of Mustard Oil Plants

Nucleotide sequences for the chloroplast rbcL gene were obtained from representatives of 11 of the 15 plant families known to produce glucosinolates (mustard oil glucosides). Parsimony analyses indicate that these constitute two widely separated groups and thus imply two independent origins of the mustard oil/myrosin cell syndrome. A strongly supported, major group comprises the mustard family Brassicaceae and its near relatives Capparaceae, Resedaceae, and Tovariaceae along with such morphologically diverse taxa as Bataceae, Bretschneideraceae, Caricaceae, Limnanthaceae, Moringaceae, and Tropaeolaceae. The second mustard oil group is restricted to the euphorbiaceous genus Drypetes

Phylogenetic Analysis Informed by Geological History Supports Multiple, Sequential Invasions of the Mediterranean Basin by the Angiosperm Family Araceae

Despite the remarkable species richness of the Mediterranean flora and its well-known geological history, few studies have investigated its temporal and spatial origins. Most importantly, the relative contribution of geological processes and long-distance dispersal to the composition of contemporary Mediterranean biotas remains largely unknown. We used phylogenetic analyses of sequences from six chloroplast DNA markers, Bayesian dating methods, and ancestral area reconstructions, in combination with paleogeographic, paleoclimatic, and ecological evidence, to elucidate the time frame and biogeographic events associated with the diversification of Araceae in the Mediterranean Basin. We focused on the origin of four species, Ambrosina bassii, Biarum dispar, Helicodiceros muscivorus, Arum pictum, subendemic or endemic to Corsica, Sardinia, and the Balearic Archipelago. The results support two main invasions of the Mediterranean Basin by the Araceae, one from an area connecting North America and Eurasia in the Late Cretaceous and one from the Anatolian microplate in western Asia during the Late Eocene, thus confirming the proposed heterogeneous origins of the Mediterranean flora. The subendemic Ambrosina bassii and Biarum dispar likely diverged sympatrically from their widespread Mediterranean sister clades in the Early-Middle Eocene and Early-Middle Miocene, respectively. Combined evidence corroborates a relictual origin for the endemic Helicodiceros muscivorus and Arum pictum, the former apparently representing the first documented case of vicariance driven by the initial splitting of the Hercynian belt in the Early Oligocene. A recurrent theme emerging from our analyses is that land connections and interruptions, caused by repeated cycles of marine transgressions-regressions between the Tethys and Paratethys, favored geodispersalist expansion of biotic ranges from western Asia into the western Mediterranean Basin and subsequent allopatric speciation at different points in time from the Late Eocene to the Late Oligocene.

Tribal relationships in Onagraceae: implications from rbcL sequence data

The evolutionary relationships among the seven tribes of Onagraceae, the most intensively studied family of intermediate size, have been examined from morphological and molecular perspectives. Previous cladistic analyses of chloroplast (cp) and nuclear ribosomal (nr) DNA restriction sites, nuclear-encoded rbcS amino acid sequences, nrRNA nucleotide sequences, and morphological characters, produced trees that agree in defining the tribe Jussiaeeae as the sister group to the rest of the family. However, the relationships among the rest of the tribes are not completely resolved by these analyses. Cladistic analyses of sequence data from the chloroplast encoded rbcL gene produced results that bear on two malor issues: (i) intertribal relationships within Onagraceae, and (ii) congruence with other cladistic analyses

Phylogenetic Relationships in Primula L. and Related Genera (Primulaceae) Based on Noncoding Chloroplast DNA

We sequenced the trnL and rpl16 introns of the chloroplast DNA from 95 of the ca. 425 species (30 of 37 sections, seven of eight subgenera) of Primula L. in order to reconstruct the phylogenetic history of the group. Among the 24 additional taxa sampled are representatives of all genera that are likely to be embedded in Primula, as well as outgroups from the Maesaceae, Theophrastaceae, and Myrsinaceae. In the strict consensus of the most parsimonious trees, Primula and the genera embedded in it (Dionysia Fenzl., Sredinskya [Stein] Fedorov, Dodecatheon L., and Cortusa L.) are sister to a clade of several genera previously suspected to be embedded in Primula (Hottonia L., Omphalogramma [Franchet] Franch., and Soldanella L.). In recognition of this, two new rankless names are defined for these clades (/Primula and /Soldanella). Close relationships are inferred between Dionysia and Primula subgenus Sphondylia (Duby) Rupr., Sredinskya and Primula subgenus Primula, Dodecatheon and Primula subgenus Auriculastrum Schott, and Cortusa and Primula subgenus Auganthus (Link) Wendelbo. The largest subgenus, Aleuritia (Duby) Wendelbo, is dispersed among three clades that are not each others closest relatives. Primula sections Muscarioides Balf. f., Soldanelloides Pax, Denticulata Watt, Armerina Lindley, and Aleuritia Duby are resolved as para‐ or polyphyletic with moderate to strong support. Throughout, we consider the striking morphological and cytological variation seen in Primula within a phylogenetic context, particularly as it relates to the close relationship implied here between Dionysia and Primula subgenus Sphondylia. The homology of involute leaf vernation in Primula is reconsidered in light of its two independent origins, and we come to the conclusion that vernation in subgenus Sphondylia is better characterized as conduplicate.

Interfamilial Relationships in Myrtales: Molecular Phylogeny and Patterns of Morphological Evolution

Numerous phenotypic (morphological, palynological, cytological, and anatomical) studies have been conducted on Myrtales, yet the detailed relationships among the families of the order remain elusive. In this paper, the rbcL sequences of 50 taxa (39 representatives of Myrtales and 11 rosid outgroups) were analyzed using parsimony and maximum likelihood to provide a phylogenetic hypothesis of intraordi- nal relationships in Myrtales. The congruence between the phenotypic data from an earlier study and the rbcL topology was assessed to identify the potential synapomorphies that would corroborate the clades supported by the molecular tree. The rbcL consensus tree defined two major clades in the order. The first clade comprised a Myrtaceae lineage sister to a Melastomataceae lineage and the second clade included Onagraceae, a Lythraceae lineage, and Combretaceae. Phenotypic characters suggest that the ancestor of the first clade was characterized by the acquisition of fibrous seed exotegmen, while the ancestor of the second clade had flowers with stamens inserted directly on the rim of the hypanthium. However, branch support for the basal split of Myrtales is weak, possibly as a result of rapid early radiation in the order.

Evolutionary biology in biodiversity science, conservation, and policy: A call to action

Evolutionary biologists have long endeavored to document how many species exist on Earth, to understand the processes by which biodiversity waxes and wanes, to document and interpret spatial patterns of biodiversity, and to infer evolutionary relationships. Despite the great potential of this knowledge to improve biodiversity science, conservation, and policy, evolutionary biologists have generally devoted limited attention to these broader implications. Likewise, many workers in biodiversity science have underappreciated the fundamental relevance of evolutionary biology. The aim of this article is to summarize and illustrate some ways in which evolutionary biology is directly relevant. We do so in the context of four broad areas: (1) discovering and documenting biodiversity, (2) understanding the causes of diversification, (3) evaluating evolutionary responses to human disturbances, and (4) implications for ecological communities, ecosystems, and humans. We also introduce bioGENESIS, a new project within DIVERSITAS launched to explore the potential practical contributions of evolutionary biology. In addition to fostering the integration of evolutionary thinking into biodiversity science, bioGENESIS provides practical recommendations to policy makers for incorporating evolutionary perspectives into biodiversity agendas and conservation. We solicit your involvement in developing innovative ways of using evolutionary biology to better comprehend and stem the loss of biodiversity.

Tracing the temporal and spatial origins of island endemics in the Mediterranean region: a case study from the citrus family (Ruta L., Rutaceae)

Understanding the origin of island endemics is a central task of historical biogeography. Recent methodological advances provide a rigorous framework to determine the relative contribution of different biogeographic processes (e.g., vicariance, land migration, long-distance dispersal) to the origin of island endemics. With its complex but well-known history of microplate movements and climatic oscillations, the Mediterranean region (including the Mediterranean basin and Macaronesia) provides the geographic backdrop for the diversification of Ruta L., the type genus of Rutaceae (citrus family). Phylogenetic, molecular dating, and ancestral range reconstruction analyses were carried out to investigate the extent to which past geological connections and climatic history of the Mediterranean region explain the current distribution of species in Ruta, with emphasis on its island endemics. The analyses showed that Ruta invaded the region from the north well before the onset of the Mediterranean climate and diversified in situ as the climate became Mediterranean. The continental fragment island endemics of the genus originated via processes of land migration/vicariance driven by connections/disconnections between microplates, whereas the oceanic island endemics were the product of a single colonization event from the mainland followed by in situ diversification. This study emphasizes the need for an integrative, hypothesis-based approach to historical biogeography and stresses the importance of temporary land connections and colonization opportunity in the biotic assembly of continental fragment and oceanic islands, respectively.