Molecular phylogenetic relationships of Melanthiaceae (Liliales) based on plastid DNA sequences

Melanthiaceae (Liliales) comprise 17 genera of rhizomatous or bulbous perennials and are distributed across the Northern Hemisphere. The relationships among the five tribes in this family have been evaluated in many molecular and morphological studies. In this study, we performed a phylogenetic analysis of the 17 genera, including 106 species of Melanthiaceae sensu APG III and nine related species as outgroups, based on sequences of five plastid regions (atpB, rbcL, matK, ndhF and trnL‐F). Support values for the monophyly of the family (BS_MP = 96%, BS_ML = 100%, PP_BI = 1.00) and each tribe were improved in comparison with previous studies. Among the tribes, Melanthieae were sister to the remainder of the family and sister relationships between Xerophylleae and Parideae (BS_MP = 96%, BS_ML = 100%, PP_BI = 1.00) and Chionographideae and Heloniadeae (BS_MP = 96%, BS_ML = 100%, PP_BI = 1.00) were confirmed. Notably, the generic concept of Veratrum s.l. including Melanthium was not supported in the present study and these genera should be treated as distinct. In the case of Parideae, the relationship of Trillium govanianum to the other species remains uncertain and requires further studies. Finally, we mapped seven representative morphological characters onto the molecular phylogenetic tree for Melanthiaceae.     

The tribal position of Fockea and Cibirhiza (Apocynaceae: Asclepiadoideae): evidence from pollinium structure and cpDNA sequence data

The pollinium morphology of the two members of the Asclepiadoideae, tribe Fockeeae, Fockea Endl. and Cibirhiza Bruyns, has been studied in detail and compared with that of eight genera of Marsdenieae, the tribe in which Fockea and Cibirhiza were previously accommodated and thus their putative closest relatives, as well as nine genera of Asclepiadeae. Both Fockea and Cibirhiza have several morphological characteristics in common, the most important of which is the absence of well-developed caudicula, which distinguishes them from all other genera of Asclepiadoideae known. The pollinium structure of these two genera, however, differs significantly. Whereas the pollinium of Cibirhiza consists of single pollen grains and is covered by a pollinium wall, as is typical for other Asclepiadoideae, the pollinium of Fockea consists of tetrads and is not covered by a pollinium wall, a condition otherwise typical of Secamonoideae. Fockea, however, has only two pollinia per anther, as does Cibirhiza and all other Asclepiadoideae, whereas the Secamonoideae have four pollinia per anther. Sequence data from two intergenic spacers, trnT-L and trnL-F and the trnL intron of cpDNA was analyzed. The ingroup included three species of Fockea and one species of Cibirhiza. The outgroup taxa consisted of three representatives each of Periplocoideae, and Secamonoideae and 24 species of Asclepiadoideae, including representatives of all tribes, of which eight genera belong to Marsdenieae, as outgroups. The results of the DNA analysis provide strong support for Fockeeae as a monophyletic tribe, distinct from Marsdenieae and, to the rest of the Asclepiadoideae. With the exception of pollen data, all morphological and molecular evidence clearly support recognition of the tribe Fockeeae. The occurrence of two such significantly different types of pollinia structure – characters elsewhere in the family used to distinguish subfamilies – within the small tribe Fockeeae was unexpected, and can perhaps best be understood as yet another attestment to the basal position of the Fockeeae in the nascence of the Asclepiadoideae.     

The taxonomy and systematics of Apocynaceae: where we stand in 2012

In their most recent classification of Apocynaceae in 2000, Endress and Bruyns recognized five subfamilies of Apocynaceae (Rauvolfioideae, Apocynoideae, Periplocoideae, Secamonoideae and Asclepiadoideae). Subsequently, through various studies using molecular data, it has been shown that most tribes and subtribes of Rauvolfioideae were not monophyletic, and new tribes and subtribes have been erected to reflect improved phylogenetic understanding of the family: Aspidospermeae in Rauvolfioideae; Nerieae, Odontadenieae and Baisseeae in Apocynoideae; Fockeeae in Asclepiadoideae; and Orthosiinae in Asclepiadeae. Several genera in Rauvolfioideae have been reassigned to different tribes in order to improve the monophyly of these tribes. The sister group of Asclepiadoideae plus Secamonoideae is not Periplocoideae, as formerly assumed, but tribe Baisseeae. Periplocoideae are nested in Apocynoideae. However, tribal composition remains unclear in some parts of the family. Clade structure in Apocynaceae is now generally well understood. The principal challenges now lie in identifying characters that can reflect and articulate these clades in a formal classification. Species‐rich, recent radiations such as core Asclepiadinae in Africa and the Metastematinae in Latin America present particular problems in this regard. © 2013 The Linnean Society of London     

Origin and early evolution of Ceropegieae (Apocynaceae-Asclepiadoideae)

The first-branching lineages of the tribe Ceropegieae, Anisotominae, Leptadeniinae, and Heterostemminae, are the focus of the present study. In the ingroup we molecularly analysed 34 samples from southern and (north)eastern Africa and Asia for the nuclear internal transcribed spacer (ITS) region, and five plastid markers (trnT-L, trnL-F and trnH-psbA intergenic spacers, trnL, and rps16 introns). Maximum Parsimony and Bayesian analyses were conducted for phylogenetic reconstruction and Bayesian binary MCMC (BBM) analysis as implemented in RASP for testing biogeographic inference. Sister to the well-known hyperdiverse subtribe Stapeliinae (around 700 species) are the Anisotominae, an African subtribe of c. 30 species, with the tropical Neoschumannia in sister-group position to the remaining genera, Anisotoma, Emplectanthus, Riocreuxia, and Sisyranthus. Emplectanthus is sequenced in full for the first time, and its monophyly as sister to Anisotoma is documented. As in Sisyranthus, new sequences in Riocreuxia, especially in Riocreuxia torulosa s.l., reveal very low genetic variation pointing to an actively radiating and speciating group, possibly in adaptation to pollinator pressure. Molecular and morphological data necessitate the transfer of Brachystelma natalense to Anisotoma, and the new combination Anisotoma natalensis is proposed. Leptadeniinae are sister to the Stapeliinae–Anisotominae clade. The position of the strictly Asian, rheophytic Pentasachme could be resolved as sister to the remaining genera of Leptadeniinae, Conomitra, Leptadenia, and Orthanthera. The position of the S Asian/Australasian wet forest lianas of the genus Heterostemma as sister to the remaining Ceropegieae is confirmed, indicating a humid Asian origin for this tribe known for its arid-adapted African succulents.     

Flavonoid glycosides from Egyptian species of the tribe Asclepiadeae (Apocynaceae,subfamily Asclepiadoideae)

The flavonoids of 11 Egyptian species of the tribe Asclepiadeae (Apocynaceae, subfamily Asclepiadoideae) were studied: Pentatropis nivalis, Pleurostelma schimperi (subtribe Astephaninae), Glossonema boveanum, Solenostemma arghel (subtribe Glossonematinae), Cynanchum acutum, Oxystelma esculentum (subtribe Metastelmatinae), Calotropis procera, Gomphocarpus fruticosus, Gomphocarpus sinaicus, Pergularia tomentosa and Pergularia daemia (subtribe Asclepiadinae). These 11 species were found to produce flavonol glycosides. In addition, flavonol sulphates and disulphates were found in a specimen of P. nivalis. The flavonoids may provide useful taxonomic characters at several levels of classification.     

ON THE POSITION OF THE GENUS KARIMBOLEA (ASCLEPIADACE AE)

The position of the rare and monotypic genus Karimbolea is clarified. Its tribal affinities are examined using mainly floral characters; the former tentative placement in the tribe Asclepiadeae (Asclepiadaceae) is confirmed. Characters of vegetative and floral morphology are used to determine the position of the genus within the tribe Asclepiadeae. Sarcostemma is identified as the most likely sister taxon of Karimbolea. Triterpenoid data obtained from gas-liquid chromatographic analysis of latex are used to test this hypothesis. This paper forms the first study within the Asclepiadaceae using two independent data sets to clarify the position of a taxon of uncertain affinities.     

A molecular phylogeny and generic rearrangement of the stapelioid Ceropegieae (Apocynaceae-Asclepiadoideae)

 Representatives of nearly all genera of the taxon-rich stem-succulent stapeliads and most of the few related, leafy genera were analyzed. Sequence data from two non-coding molecular markers (ITS region of nrDNA and trnT-L and trnL-F spacers as well as the trnL intron of cpDNA) support the traditional tribal affiliation of the genera, which form a monophyletic group. This monophylum breaks into a basal Neoschumannia/Anisotoma/Riocreuxia/Sisyranthus nk;clade, from which the core Ceropegieae are derived. The four Ceropegia species included are not monophyletic, and their relationship to Brachystelma changes depending on the marker studied. The stem succulent taxa fall in a number of well supported, but unresolved clades, the most prominent being the predominantly southern African clade comprising Orbea, Stapelia and some other genera. The most derived taxa of NE Africa, Duvaliandra and White-sloanea, are basal to this southern African clade. The other clades comprise the more basal genera of stem-succulent stapeliads, including the members of the Caralluma complex. Of the 17 genera accepted by Plowes for the Caralluma complex, seven are recognized: Caralluma, Apteranthes, Australluma, Boucerosia, Caudanthera, Desmidorchis and Monolluma. New combinations are proposed in 15 cases; Caralluma adscendens var. geniculata is raised to specific rank. Anomalluma is reinstated, and Pseudolithos mccoyi is transfered to it. A broadened concept for Orbea (incl. Angolluma and Orbeopsis) is recognized, but Orbeanthus is kept separate. The monotypic Ballyanthus, recently separated from Orbea, is nested within Duvalia. Piaranthus (incl. Huerniopsis) is monophyletic. The bitypic Notechidnopsis is reduced to the type species, N. tessellata, while N. columnaris is transferred to a new genus, Richtersveldia. Received February 25, 2002; accepted June 17, 2002 Published online: November 7, 2002 Address of the authors: Dr. Ulrich Meve (e-mail: ulrich.meve@uni-bayreuth.de) and Prof. Dr. Sigrid Liede (e-mail: sigrid.liede@uni-bayreuth.de), Universit?t Bayreuth, Lehrstuhl für Pflanzensystematik, Universit?tsstrasse 30, D-95440 Bayreuth, Germany.     

Systematics of Amaryllidaceae based on cladistic analysis of plastid sequence data

Cladistic analyses of plastid DNA sequences rbcL and trnL-F are presented separately and combined for 48 genera of Amaryllidaceae and 29 genera of related asparagalean families. The combined analysis is the most highly resolved of the three and provides good support for the monophyly of Amaryllidaceae and indicates Agapanthaceae as its sister family. Alliaceae are in turn sister to the Amaryllidaceae/Agapanthaceae clade. The origins of the family appear to be western Gondwanaland (Africa), and infrafamilial relationships are resolved along biogeographic lines. Tribe Amaryllideae, primarily South African, is sister to the rest of Amaryllidaceae; this tribe is supported by numerous morphological synapomorphies as well. The remaining two African tribes of the family, Haemantheae and Cyrtantheae, are well supported, but their position relative to the Australasian Calostemmateae and a large clade comprising the Eurasian and American genera, is not yet clear. The Eurasian and American elements of the family are each monophyletic sister clades. Internal resolution of the Eurasian clade only partially supports currently accepted tribal concepts, and few conclusions can be drawn on the relationships of the genera based on these data. A monophyletic Lycorideae (Central and East Asian) is weakly supported. Galanthus and Leucojum (Galantheae pro parte) are supported as sister genera by the bootstrap. The American clade shows a higher degree of internal resolution. Hippeastreae (minus Griffinia and Worsleya) are well supported, and Zephyranthinae are resolved as a distinct subtribe. An Andean clade marked by a chromosome number of 2n = 46 (and derivatives thereof) is resolved with weak support. The plastid DNA phylogenies are discussed in the context of biogeography and character evolution in the family.     

Phylogenetic analysis of the Malvadendrina clade (Malvaceae s.l.) based on plastid DNA sequences

Phylogenetic relationships within Malvaceae s.l., a clade that includes the traditional families Bombacaceae, Malvaceae s.str., Sterculiaceae, and Tiliaceae, have become greatly clarified thanks to recent molecular systematic research. In this paper, we use DNA sequences of four plastid regions (atpB, matK, ndhF, and rbcL) to study relationships within Malvadendrina, one of the two major clades of Malvaceae s.l. The four data sets were generally in agreement, but five terminal taxa manifested highly unexpected affinities in the rbcL partition, and the non-coding sequences of the trnK intron were found to provide limited phylogenetic information for resolving relationships at the base of Malvadendrina. The remaining data strongly support the existence of six major clades within Malvadendrina: Brownlowioideae, Dombeyoideae, Helicteroideae, Malvatheca (comprising Bombacoideae and Malvoideae), Sterculioideae, and Tilioideae. These data also resolve the placement of two problematic taxa: Nesogordonia (in Dombeyoideae) and Mortoniodendron (in Tilioideae). The relationships among the six clades are not definitively resolved, but the best-supported topology has Dombeyoideae as sister to the remainder of Malvadendrina (posterior probability PP=80%) and Sterculioideae as sister to Malvatheca (PP=86%). This early branching position of Dombeyoideae is supported by similarities in floral characters between members of that clade and outgroup taxa in Byttnerioideae. Similarly, the sister-group relationship of Sterculioideae and Malvatheca receives support from androecial characteristics, like subsessile or sessile anthers and an absence of staminodes, shared by these two clades.     

Neurocranial osteology and systematic relationships of Varanus (Megalania) prisca Owen, 1859 (Squamata: Varanidae)

The neurocranial osteology of the giant monitor lizard Varanus (Megalania) prisca Owen, 1859 is described in detail for the first time. Optimization of neurocranial characters onto phylogenetic topologies for varanoids, including Lanthanotus, Heloderma, and Varanus species nests V. prisca within an Indo‐Australian clade of Varanus on the basis of characters of the otic capsule. A sister‐taxon relationship between V. prisca and Varanus komodoensis Ouwens, 1912 is proposed based on apomorphies of the crista prootica, fenestra vestibuli, occipital recess, and supraoccipital. These results support a monophyletic clade of giant monitors among Indo‐Australian species, and unambiguously synonymize Megalania with Varanus at both generic and subgeneric levels. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 445–457.     

Phylogenetic relationships within the South American fish family Anostomidae (Teleostei,Ostariophysi, Characiformes)

Analysis of a morphological dataset containing 152 parsimony‐informative characters yielded the first phylogenetic reconstruction spanning the South American characiform family Anostomidae. The reconstruction included 46 ingroup species representing all anostomid genera and subgenera. Outgroup comparisons included members of the sister group to the Anostomidae (the Chilodontidae) as well as members of the families Curimatidae, Characidae, Citharinidae, Distichodontidae, Hemiodontidae, Parodontidae and Prochilodontidae. The results supported a clade containing Anostomus, Gnathodolus, Pseudanos, Sartor and Synaptolaemus (the subfamily Anostominae sensu Winterbottom) albeit with a somewhat different set of relationships among the species within these genera. Anostomus as previously recognized was found to be paraphyletic and is split herein into two monophyletic components, a restricted Anostomus and the new genus Petulanos gen. nov. , described herein. Laemolyta appeared as sister to the clade containing Anostomus, Gnathodolus, Petulanos, Pseudanos, Sartor and Synaptolaemus. Rhytiodus and Schizodon together formed a well‐supported clade that was, in turn, sister to the clade containing Anostomus, Gnathodolus, Laemolyta, Petulanos, Pseudanos, Sartor and Synaptolaemus. Anostomoides was sister to the clade formed by these nine genera. Leporinus as currently defined was not found to be monophyletic, although certain clades within that genus were supported, including the species with subterminal mouths in the former subgenus Hypomasticus which we recognize herein as a genus. Abramites nested in Leporinus, and Leporellus was found to be the most basal anostomid genus. The presence of cis‐ and trans‐Andean species in Abramites, Leporellus, Leporinus and Schizodon, all relatively basal genera, suggests that much of the diversification of anostomid species pre‐dates the uplift of the Andean Cordilleras circa 11.8 million years ago. Several important morphological shifts in anostomid evolution are illustrated and discussed, including instances of convergence and reversal. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154 , 70–210.     

Global phylogeny of Hadrosauridae (Dinosauria: Ornithopoda) using parsimony and Bayesian methods

The late Cretaceous hadrosaurids were the most specialized and diverse clade of ornithopod dinosaurs. Parsimony and Bayesian methods were implemented to elucidate the phylogenetic relationships of all hadrosaurid species. Traditional and geometric morphometrics were applied to discover patterns of variation containing phylogenetic information. In total, 286 phylogenetically informative characters (196 cranial and 90 postcranial) were defined and documented: the most extensive character data set ever constructed for hadrosaurid dinosaurs. Of these, 136 characters were used for the first time in phylogenetic analysis of these ornithopods, and 93 were modified from those of other authors. Parsimony and the Bayesian analysis (using the Mk model without the gamma parameter) confirmed the split of hadrosaurids into Saurolophinae and Lambeosaurinae. Saurolophines included a major clade composed of the Prosaurolophus–Saurolophus and the Kritosaurus–Gryposaurus–Secernosaurus subclades. Edmontosaurus and Shantungosaurus were recovered outside the major clade of saurolophines. The Brachylophosaurus clade was recovered as the most basal clade of saurolophines in the parsimony analysis, whereas following the Bayesian analysis it was recovered as the sister clade to the Kritosaurus–Gryposaurus–Secernosaurus clade. These two analyses resulted in a Lambeosaurinae composed of a succession of Eurasian sister taxa to two major clades: the Parasaurolophus clade and the Hypacrosaurs altispinus–Corythosaurus clade. In contrast, the Bayesian analysis using the Mk model with the gamma parameter included, resulted in an unbalanced hadrosauroid tree, with a paraphyletic Saurolophinae, and with the Prosaurolophus clade, Edmontosaurus, and Shantungosaurus as successively closer sister taxa to Lambeosaurinae. Based on the strict reduced consensus tree derived from the parsimony analysis, Hadrosauridae was redefined as the clade stemming from the most recent common ancestor of Hadrosaurus foulkii and Parasaurolophus walkeri. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 159 , 435–502.     

Comparative morphology,phylogenetic relationships,and historical biogeography of plesiolebiasine seasonal killifishes (Teleostei: Cyprinodontiformes: Rivulidae)

Phylogenetic and biogeographical analyses were performed for the Plesiolebiasini, a group of 20 small and rare species of South American annual killifishes, some threatened with extinction, occurring in river basins of Brazil, Bolivia, Paraguay, and Argentina. The results of a maximum parsimony analysis of 142 morphological characters highly corroborate monophyly of the Plesiolebiasini. Monophyly of each plesiolebiasine genus is supported and Plesiolebias is hypothesized to be the sister group to a clade comprising the remaining plesiolebiasine genera (Papiliolebias, Pituna, Maratecoara, and Stenolebias), corroborating studies based on mitochondrial genes. The phylogenetic analysis supports sister group relationships between Papiliolebias and the clade containing Pituna, Maratecoara, and Stenolebias, and between Maratecoara and Stenolebias. The biogeographical analysis indicates a complex historical biogeographical scenario for plesiolebiasines. A vicariance event between the western Paraguay area and northern river basins may be related to the isolation of the Chaco region from the Amazon between the Late Oligocene and Early Miocene. A vicariance event involving the Paraguay River basin and northern rivers, and the multiple occurrence of area hybridism along the Araguaia depression are tentatively identified as a consequence of tectonic subsidence events occurring during the Pleistocene. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162 , 131–148.     

What is the subtribe Glossonematinae (Apocynaceae: Asclepiadoideae)? A phylogenetic study based on cpDNA spacer

Evidence from the chloroplast trn T-L spacer, trn L intron and trn L-F spacer shows the subtribe Glossonematinae of the tribe Asclepiadeae, hitherto composed of the Arabian and North African genera Glossonema , Odontanthera and Solenostemma , not to be monophyletic. While the affinities of Solenostemma cannot be determined with certainty at present, molecular, karyological and morphological evidence suggests that Glossonema and Odontanthera are closely allied to Pentarrhinum , an African genus of five species, belonging to the Cynanchinae.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 139 , 145–158.     

Molecular relationships in Genista L. Sect. Spartocarpus Spach (Fabaceae)

 A molecular investigation on the species of Genista belonging to sect. Spartocarpus was undertaken with the aim of inferring phylogenetic relationships within the section. The combined cladistic analysis of ITS 1 and 2 and of the chloroplast DNA trnL^(UAA) intron yielded a single most parsimonious cladogram with G. aetnensis as sister group to the rest of the section, which is in turn divided into two major clades, one of which includes G. nissana, G. aucheri, G. sessilifolia, G. spartioides and G. haenseleri, and the other the remaining species of the section. This latter group has G. holopetala and G. radiata forming a clade, which is a sister group to a not entirely resolved monophylum including the G. ephedroides species complex. The latter shows a basal collapse of G. cilentina, G. dorycnifolia and G. numidica, and then a clade including G. ephedroides and G. valsecchiae, as well as a collapsed monophylum with G. demarcoi, G. gasparrini and G. thyrrena. The clade including the G. ephedroides species group has the lowest within-group molecular variation and appears to be recent in origin, as compared to the rest of the section. The pattern of cladogeny in the section seems to be related to geographical isolation, and is broadly congruent with palynological and biogeographical evidence. Received March 3, 2001 Accepted October 8, 2001     

Phylogeny and divergence times of the Coluteoid clade with special reference to Colutea (Fabaceae) inferred from nrDNA ITS and two cpDNAs,matK and rpl32-trnL(UAG) sequences data

This study reconstructed the phylogeny of the Coluteoid clade using nrDNA ITS and plastid matK and rpl32-trnL_(UAG) sequences data. The analyses resolve a well-supported Coluteoid clade, as sister to Astragalus s.str. + Oxytropis, nested within the larger, strongly supported Astragalean clade. The Coluteoid clade is now composed of 12 genera including Podlechiella, Swainsona, Carmichaelia, Clianthus, Montigena, Phyllolobium, Lessertia, Sutherlandia, Sphaerophysa, Smirnowia, Eremosparton and Colutea. Within this clade, Podlechiella is the first diverging lineage followed by successive subclades of Carmichaelia + Clianthus + Swainsona, Phyllolobium, Lessertia + Sutherlandia, Sphaerophysa + Smirnowia + Eremosparton, and Colutea. We assigned the formal tribal name to this clade and redefined the tribe Coluteae. A diagnostic key to the genera of the tribe is presented. Astragalus cysticalyx and A. sinicus have no relationship with the Coluteoid clade, instead, they are nested in Astragalus s. str. Resolution within Colutea is rather low, but several smaller subclades with low to high supports are found in the genus. None of the large sections in Colutea are monophyletic. Divergence time estimates revealed that the Coluteoid clade originated in the Early Miocene (20.4 Mya). Most of its members were diverged during the Late Miocene to Pliocene. Colutea and Podlechiella form the youngest lineages where the diversification occurred in the Pliocene-Pleistocene.     

An osteology‐based appraisal of the phylogeny and evolution of kangaroos and wallabies (Macropodidae: Marsupialia)

Macropodids are the most diverse group of marsupial herbivores ever to have evolved. They have been the subject of more phylogenetic studies than any other marsupial family, yet relationships of several key clades remain uncertain. Two important problem areas have been the position of the merrnine (Lagostrophus fasciatus) and the phylogenetic proximity of tree‐kangaroos and rock‐wallabies. Our osteological analysis revealed strong support for a plesiomorphic clade ( Lagostrophinae subfam. nov. ) containing Lagostrophus and Troposodon, which is likely to have originated in the early Miocene. The extinct short‐faced kangaroos (Sthenurinae) emerged in the middle Miocene as the sister lineage to a clade containing all other living kangaroos and wallabies (Macropodinae). New Guinea forest wallabies ( Dorcopsini trib. nov. ) are the most plesiomorphic macropodines; the other two main lineages include tree‐kangaroos and rock‐wallabies (Dendrolagini), and ‘true’ kangaroos and wallabies (Macropodini). These phylogenetic outcomes are broadly consistent with the results of recent molecular studies, although conflicts remain over the relative positions of some macropodins (e.g. Setonix, Onychogalea, and Wallabia). Given the presence of derived dendrolagins and macropodins in early Pliocene localities, it is probable that most macropodine genera originated in the late Miocene. Key functional–adaptive trajectories within the craniodental and locomotory systems of the dominant macropodid lineages represent varying responses to the spread of drier, open habitats following the Miocene Climatic Optimum. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 159 , 954–987.