Gynoecium diversity and systematics of the Laurales

Carpel and ovule structure was comparatively studied in representatives of all eight families of the Laurales: Amborellaceae, Calycanthaceae, Chloranthaceae, Gomortegaceae, Hernandiaceae, Lauraceae, Monimiaceae, and Trimeniaceae. In all representatives the carpels are closed at anthesis. As in Magnoliales/winteroids, closure takes place in three different modes: (1) by postgenital fusion of the stylar (and ovarial) ventral slit (Calycanthaceae, Gomortegaceae, Lauraceae, Hernandiaceae); (2) by occlusion of the inner space by secretion (Amborellaceae, Chloranthaceae, Trimeniaceae, Mollinedioideae of Monimiaceae), all having extremely ascidiate carpels; (3) by a combination of (1) and (2), whereby the ventral slit in the style is postgenitally fused but a central canal remains open, which is filled by secretion (Monimiaceae except Mollinedioideae). The carpels have a single ovule in ventral median placentation; only Calycanthaceae have two lateral ovules, although the upper ovule degenerates. In contrast to Magnoliales/winteroids, several representatives have orthotropous or almost orthotropous ovules (Amborellaceae, Chloranthaceae, Gomortegaceae). Mature ovules vary in length between 425 μm (some Monimiaceae) and 1500 urn (some Calycanthaceae, Trimeniaceae). Although all ovules are crassinucellar, nucellus breadth varies between 60 μm (Chimonanthus, Calycanthaceae) and 500 μm (Hemandia, Hernandiaceae). In almost all representatives the single ovule (two in Calycanthaceae) tightly fills out the ovarial cavity. The micropyle is mostly formed by the inner integument. In a few cases there is no micropyle and the nucellar apex makes direct contact with the inner ovary surface or the funicle (Lauraceae p.p., Calycanthaceae p.p., Hernandiaceae p.p., Monimiaceae p.p.). The ovule is pachychalazal (or perichalazal) in Lauraceae, some Hernandiaceae, and Gomortegaceae. Both integuments are variously lobed or unlobed. The outer integument is semiannular or annular, and this may vary within a family (Calycanthaceae, Hernandiaceae, Monimiaceae); it is also exceedingly diverse in thickness (2–23 cell layers). Gynoecial traits support the association of Chloranthaceae, Trimeniaceae, and Amborellaceae, and also separately Gomortegaceae, Hernandiaceae, and Lauraceae. In addition, affinities of the first group with Schisandraceae, Illiciaceae and Austrobaileyaceae may also be supported.     

Trans‐Atlantic,trans‐Pacific and trans‐Indian Ocean dispersal in the small Gondwanan Laurales family Hernandiaceae

Aim To investigate the historical biogeography of the pantropical flowering plant family Hernandiaceae (Laurales), which today comprises 62 species in five genera. Location Hernandiaceae occur in Africa (9 species), Madagascar (4), the Neotropics (25), Australia (3), southern China, Indochina, Malesia, and on numerous Pacific Islands (32). These numbers include two widespread species, Hernandia nymphaeifolia, which ranges from East Africa to the Ogasawara Islands and New Caledonia, and Gyrocarpus americanus, thought to have a pantropical range. Methods We sampled 37 species from all genera, the widespread ones with multiple accessions, for a chloroplast DNA matrix of 2210 aligned nucleotides, and used maximum likelihood to infer species relationships. Divergence time estimation relied on an uncorrelated‐rates relaxed molecular clock calibrated with outgroup fossils of Lauraceae and Monimiaceae. Results The deepest split in the family is between a predominantly African–Madagascan–Malesian lineage comprising Hazomalania, Hernandia and Illigera, and an African–Neotropical lineage comprising Gyrocarpus and Sparattanthelium; this split may be 122 (110–134) Myr old. The stem lineages of the five genera date back at least to the Palaeocene, but six splits associated with transoceanic range disjunctions date only to the Oligocene and Miocene, implying long‐distance dispersal. It is inferred that Hernandia beninensis reached the West African islands of São Tomé and Bioko from the West Indies or the Guianas; Hernandia dispersed across the Pacific; and Illigera madagascariensis reached Madagascar from across the Indian Ocean. Main conclusions The disjunct ranges and divergence times of sister clades in the Hernandiaceae are partly congruent with the break‐up of West Gondwana, but mostly with later transoceanic dispersal. An exceptional ability to establish following prolonged oceanic dispersal may be largely responsible for the evolutionary persistence of this small clade.     

Chromosome evolution in the Laurales based on analyses of original and published data

We present a summary of currently available chromosome information for all seven families in the order Laurales on the basis of original and previously published data and discuss the evolution of chromosomes in this order. Based on a total of 53 genera for which chromosome data were available, basic chromosome numbers appear consistent within families: x = 11 (Calycanthaceae); x = 22 (Atherospermataceae and Siparunaceae); x = 19 (Monimiaceae); and x = 12 and 15 (Lauraceae). The Hernandiaceae have diverse numbers: x = 15 (Gyrocarpoideae) and x = 18 and 20 (Hernandioideae). Karyotype analyses showed that Hennecartia, Kibaropsis, and Matthaea (all Monimiaceae) contained two or three sets of four distinct chromosomes in 38 somatic chromosomes, suggesting that 2n = 38 was derived by aneuploid reduction from 2n = 40, a tetraploid of x = 10. In light of the overall framework of phylogenetic relationships in the Laurales, we show that x = 11 is an archaic base number in the order and is retained in the Calycanthaceae, which are sister to the remainder of the order. Polyploidization appears to have occurred from x = 11 to x = 22 in a common clade of the Siparunaceae, Atherospermataceae, and Gomortegaceae (although 2n = 42 in the Gomortegaceae), and aneuploid reduction from x = 11 to x = 10 occurred in a common clade of the Hernandiaceae, Lauraceae, and Monimiaceae. To understand chromosome evolution in the Lauraceae, however, more studies are needed of genera and species of Cryptocaryeae.     

Sex and age differences in reflectance and biochemistry of carotenoid‐based colour variation in the great tit Parus major

The plumage coloration in great tits (Parus major) is the subject of much behavioural and ecophysiological research, yet there is a lack of analyses of the natural colour variation and its mechanisms. We used reflectance spectrometry and high‐performance liquid chromatography to explore individual, sexual and age‐related variation in carotenoid coloration and pigmentation, paramount to the often presumed, but rarely substantiated, costs and ‘honesty’ of carotenoid displays. In adults, we found that sex was the strongest predictor of ‘brightness’ (higher in males) and of ‘hue’ (longer wavelength in females). There was no sex difference in ‘carotenoid chroma’ or carotenoid content of feathers which also was unrelated to adult age (1 or 2+ years) and condition. Similar patterns were revealed for nestlings. Regarding the biochemical ‘signal content’, ‘carotenoid chroma’, but not ‘hue’, was significantly related to the carotenoid content (lutein and zeaxanthin) of feathers. These results refute the previously assumed exaggeration of carotenoid pigmentation in male great tits, and question the condition‐dependence of carotenoid coloration in this species. However, the sexual dimorphism in total reflectance or ‘brightness’, most likely due to melanins rather than carotenoids, may have implications for signalling or other adaptive explanations that need to be explored. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 758–765.     

Embryology of Gomortegaceae (Laurales): characteristics and character evolution

The embryological characteristics of Gomortegaceae, which are poorly understood, were investigated on the basis of Gomortega nitida, the only species of the family, to understand better the evolution of this group within Laurales. Comparisons with other Laurales and Magnoliales (a sister group of Laurales) show that Gomortega has many embryological features in common with the other lauralean families. Notably, Gomortega shares a testa without or with at best only a poorly developed mesotesta as a synapomorphy with all other Laurales. The genus further shares anthers dehisced by valves as a synapomorphy with the other Laurales (except for Calycanthaceae and Monimiaceae), and a non-multiplicative testa and bisporangiate anther as synapomorphies with Atherospermataceae and Siparunaceae (although the non-multiplicative testa occurs as a homoplasy in Monimiaceae, and the bisporangiate anther in Monimiaceae pro parte, Lauraceae pro parte and Hernandiaceae, respectively). Gomortega shows simultaneous cytokinesis to form pollen grains, a one-celled ovule archesporium and non-specialized chalaza, all or part of which may be synapomorphies shared with Atherospermataceae. Gomortega appears to have no embryological autapomorphies, but further comparison with Atherospermataceae is required.Kweon Heo and Yukitoshi Kimoto contributed equally to this work.     

Plastid phylogenomics improve phylogenetic resolution in the Lauraceae

Abstract The family Lauraceae is a major component of tropical and subtropical forests worldwide, and includes some commercially important timber trees and medicinal plants. However, phylogenetic relationships within Lauraceae have long been problematic due to low sequence divergence in commonly used markers, even between morphologically distinct taxa within the family. Here we present phylogenetic analyses of 43 newly generated Lauraceae plastomes together with 77 plastomes obtained from GenBank, representing 24 genera of Lauraceae and 17 related families of angiosperms, plus nine barcodes from 19 additional species in 18 genera of Lauraceae, in order to reconstruct highly supported relationships for the Lauraceae. Our phylogeny supports the relationships: sisterhood of the Lauraceae and a clade containing Hernandiaceae and Monimiaceae, with Atherospermataceae and Gomortegaceae being the next sister groups, followed by Calycanthaceae. Our results highlight a monophyletic Lauraceae, with nine well‐supported clades as follows: Hypodaphnis clade, Beilschmiedia–Cryptocarya clade, Cassytha clade, Neocinnamomum clade, Caryodaphnopsis clade, Chlorocardium–Mezilaurus clade, Machilus–Persea clade, Cinnamomum–Ocotea clade, and Laurus–Neolitsea clade. The topology recovered here is consistent with the patterns of plastome structural evolution and morphological synapomorphies reported previously. More specifically, flower sex, living type, inflorescence type, ovary position, anther locus number, leaf arrangement, leaf venation, lateral vein number, tree height, and inflorescence location all represent morphological synapomorphies of different lineages. Our findings have taxonomic implications and two new tribes, Caryodaphnopsideae and Neocinnamomeae, are described, and the composition of four other tribes is updated. The phylogeny recovered here provides a robust phylogenetic framework through which to address the evolutionary history of the Magnoliids, the third‐largest group of Mesangiospermae.     

Interactions between Papilionidae and ancient Australian Angiosperms: evolutionary specialization or ecological monophagy?

The evolution of host range for insect herbivores involves many behavioral, physiological, and biochemical adaptations that often lead to locally specialized populations or species. Such specialization may be constrained by ecological factors (e.g., local host availability) or by evolutionary factors (e.g., phylogenetic divergence in behavioral, physiological, or biochemical detoxification enzymes; and potential inabilities to return to ancestral hosts). While insect adaptations to new hosts can be rapid, ancient detoxification systems may persist in some lineages of swallowtail butterflies (Papilionidae) for millions of years. Here, we test various species of specialized species/populations of Papilionidae (Lepidoptera) from North America and from Australia on an array of Australian host plant families in order to determine whether the current feeding constraints reflect loss of capabilities to recognize and use hosts other than their current (local) favorites. We selected two species of Lauraceae specialists (Papilio troilus L. and Papilio palamedes Drury) from North America and one locally specialized population of Papilio glaucus L. that only uses one plant species in the Magnoliaceae in Florida. We also examined three species/populations of Australian swallowtails for comparison, including the Monimiaceae‐specialized Graphium macleayanum moggana L. E. Couchman, the Rutaceae‐specialized Papilio aegeus Donovan, and the Annonaceae‐specialized Graphium eurypylus L. Our aim was to determine whether neonate larvae of these six specialists could survive on any plants other than their currently favored species. While the Lauraceae specialists could use nothing else and were thus evolutionarily constrained, the Magnoliaceae‐, Rutaceae‐, and Monimiaceae specialists all had common abilities to accept, feed, and grow on plants in the Lauraceae, Monimiaceae, Magnoliaceae, and Rutaceae families. Even the Annonaceae specialist was discovered using Magnoliaceae in the field, suggesting existence here also of both flexiblity in preferences and detoxification abilities and ‘ecological monophagy’.     

A second species in the endemic New Caledonian genus Gastrolepis (Stemonuraceae) and its implications for the conservation status of high‐altitude maquis vegetation: coherent application of the IUCN Red List criteria is urgently needed in New Caledonia

A new species in the previously monotypic, endemic New Caledonian genus Gastrolepis (Stemonuraceae) is described. Gastrolepis alticola differs from G. austrocaledonica by its shorter and thicker petioles, strongly coriaceous leaves with revolute margins, shorter inflorescences, and pubescent corollas. The new species is further distinguished by its ecology, occurring only in high‐altitude maquis on two massifs in southern New Caledonia, Mt. Kouakoué and the Montagne des Sources. Gastrolepis alticola is assigned a preliminary conservation status of ‘Endangered’ using the World Conservation Union (IUCN) Red List criteria. Comparison of the IUCN threat status for the 19 species endemic to this distinctive, restricted vegetation type reveals a striking lack of consistency and underscores the need for a reassessment of the entire New Caledonian flora. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157 , 775–783.     

Parallel evolution of termite‐egg mimicry by sclerotium‐forming fungi in distant termite groups

Among the great diversity of insect–fungus associations, fungal mimicry of termite eggs is a particularly fascinating consequence of evolution. Along with their eggs, Reticulitermes termites often harbour sclerotia of the fungus Fibularhizoctonia sp., called ‘termite balls’, giving the fungus competitor‐free habitat within termite nests. The fungus has evolved sophisticated morphological and chemical camouflage to mimic termite eggs. To date, this striking insect–fungus association has been found in eight temperate termite species, but is restricted to the lower termite genera Reticulitermes and Coptotermes. Here, we report the discovery of a novel type of termite ball (‘Z‐type’) in the subtropical termite, Nasutitermes takasagoensis. Phylogenetic analysis indicated that the Z‐type termite ball is an undescribed Trechisporoid fungus, Trechispora sp., that is phylogenetically distant from Fibularhizoctonia, indicating two independent origins of termite‐egg mimicry in sclerotium‐forming fungi. Egg protection bioassays using dummy eggs revealed that Reticulitermes speratus and N. takasagoensis differ in egg‐size preference. A comparative study of termite ball size and egg‐size preference of host termites showed that both fungi evolved a termite ball size that optimized the acceptance of termite balls as a unit investment. Termite‐egg mimicry by these fungi offers a model case of parallel evolution. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 531–537.     

Fruit structure and systematics of Monimiaceae s.s. (Laurales)

Fruit structure (anatomy) was studied in 27 species of 15 genera of Monimiaceae s.s. Almost all have apocarpous gynoecia, with the carpels more or less surrounded by a floral cup. The fruitlets are presented on the opened floral cup, which, depending on its pre‐ and post‐floral development, differentially contributes to the attractive part of the mature fruit. Morphologically similar fruits may differ conspicuously in anatomical structure. Based on anatomical characters two different fruit forms were found: drupe(let)s (with compact sclerenchymatic endocarp forming a stone: putamen) and berry(let)s (with parenchymatic endocarp, and mesocarp parenchyma containing isolated sclereid nests). Four types of drupelets differing by the endocarp structure were tentatively distinguished: (1) the Monimia‐type has a many‐cell‐layered putamen of large isodiametric sclereids, interrupted on the ventral side by few radial rows of small sclereids; (2) the Hortonia‐type has a few‐cell‐layered putamen of isodiametric, especially thick‐walled sclereids – it may be composed of two lateral halves, i.e. with the sclerenchyma partially interrupted on the ventral and dorsal sides (but without rows of small sclereids); (3) the Mollinedia‐type has a few‐cell‐layered putamen, with more or less radially elongate sclereids with wavy cell walls; and (4) the Hedycarya‐type has a one‐cell‐layered putamen of pronouncedly radially elongate sclereids with wavy cell walls. Drupelets of some taxa with a single‐cell‐layered endocarp with only weakly thickened cell walls may represent a transition from drupelets to berrylets. The fruit structure supports three major clades recognized earlier by morphological studies and by molecular phylogenetic analyses: (1) Monimioideae (Monimia‐type drupelets), (2) Hortonieae of Mollinedioideae (Hortonia‐type drupelets), and (3) the remainder of Mollinedioideae (Hedycarya‐ and Mollinedia‐types) and berrylets. Fruit structure also supports the close relationship of Monimiaceae and Lauraceae. © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society, 2007, 153 , 265–285.     

Ecophysiology of seed dormancy and the control of germination in early spring‐flowering Galanthus nivalis and Narcissus pseudonarcissus (Amaryllidaceae)

Seed dormancy induction and alleviation in the winter‐flowering, moist temperate woodland species Galanthus nivalis and Narcissus pseudonarcissus are complex and poorly understood. Temperature, light and desiccation were investigated to elucidate their role in the germination ecophysiology of these species. The effect of different seasonal temperatures, seasonal durations, temperature fluctuations, the presence of light during different seasons and intermittent drying (during the summer period) over several ‘years’ on seed germination was investigated with outdoor and laboratory experiments. Warm summer‐like temperatures (20 °C) were necessary for germination at subsequent cooler autumn‐like temperatures (greatest at 15 °C in G. nivalis and 10 °C in N. pseudonarcissus). As the warm temperature duration increased, so did germination at subsequent cooler temperatures; further germination occurred in subsequent ‘years’ at cooler temperatures following a second, and also third, warm period. Germination was significantly greater in darkness, particularly in G. nivalis. Dormancy increased with seed maturation period in G. nivalis, because seeds extracted from green capsules germinated more readily than those from yellow capsules. Desiccation increased dormancy in an increasing proportion of N. pseudonarcissus seeds the later they were dried in ‘summer’. Seed viability was only slightly reduced by desiccation in N. pseudonarcissus, but was poor and variable in G. nivalis. Shoot formation occurred both at the temperature at which germination was greatest and also if 5 °C cooler. In summary, continuous hydration of seeds of both species during warm summer‐like temperatures results in the gradual release of seed dormancy; thereafter, darkness and cooler temperatures promote germination. Cold temperatures, increased seed maturity (G. nivalis) and desiccation (N. pseudonarcissus) increase dormancy, and light inhibits germination. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 177 , 246–262.     

COMPARATIVE MORPHOLOGY AND PHYLOGENY OF THE RANALES

Comparative morphological studies of woody Ranales have established the primitive status of the group and hence their key place in angiosperm phylogeny. Significant advances in our knowledge of some ranalian families have been made in recent years. An attempt is made in the present review to bring together a range of morphological data (vegetative and floral anatomy, palynology and embryology) on the Ranales (sensu lato), with particular reference to research work published after the publication of Eames's (1961) book, and to discuss the relationships of the families. Recent ontogenetic studies have shown that the carpel of Drimys is ascidial and not conduplicate as earlier suggested. The inclusion of Degeneria in the Winteraceae is not supported by morphological data. Melville's gonophyll theory has been shown to be inapplicable to the magnoliaceous flower. The pollen of Schisandra is interpreted as derived and specialized rather than primitive as previously supposed. The removal of Schisandra from Magnoliaceae is upheld by morphological evidence. Recent morphological studies do not support a close relationship between Schisandraceae and Illiciaceae suggested by earlier authors. The Canellaceae shows similarities to Winteraceae, Magnoliaceae, Illiciaceae, Eupteleaceae and Myristicaceae. Transitional types of division of pollen mother cells found in Winteraceae, Schisandraceae and Annonaceae and their probable phylogenetic significance have been discussed. The Annonaceae, Winteraceae, Degeneriaceae, Magnoliaceae, Schisandraceae and Cercidiphyllaceae share several embryological features in addition to similarities in floral structure. Ruminate endosperm is regarded either as an archaic feature retained in some taxa or as a later and parallel development in others. Thus its value in assessing relationships seems to be doubtful. Myristicaceae has been shown to be closely related neither to the the Annonaceae nor to the Lauraceae. The suggested relationship of Eupomatiaceae to Annonaceae is not supported by palynology. Floral cortical vascular systems in Magnoliaceae, Annonaceae, Calycanthaceae and Myristicaceae have been compared and it is concluded that they may be vestigial structures. A great deal of similarity has been found between Lauraceae and Calycanthaceae in wood, node, flower structure and embryology. Further floral anatomical evidence has been adduced to support the removal of Scyphostegia from Monimiaceae. The Hernandiaceae show similarities to some members of Monimiaceae while the Gyrocarpaceae resemble the Lauraceae, Gomortegaceae and certain other genera of Monimiaceae. Available evidence from wood and floral anatomy and embryology indicates close relationships among Lauraceae, Monimiaceae and Hernandiaceae. Vegetative and floral anatomical and embryological data seem to indicate a place for the Chloranthaceae in the ranalian complex. Recent anatomical studies in the Nymphaeaceae show that the floral structure is of a primitive type with similarities to the woody Ranales. Available morphological evidence is considered inadequate to express an opinion on the splitting of the family. Ceratophyllaceae is regarded as a highly reduced ranalian family derived most probably from a nymphaeaceous stock. The gynoecium in Berberidaceae is interpreted as monocarpellate. No evidence has been found to support the tricarpellate view. Berberidaceae, Lardizabalaceae and Menispermaceae share several embryological features, while at the same time showing evidence of specialization, each in its own way. Thus they might have arisen from a common stock and early diverged along different lines. The occurrence of several types of embryo sac in Ranunculaceae may well be an indication of specialization, but their probable taxonomic value, if any, is not yet clear. The occurrence of numerous primitive features in Paeonia has been suggested as an argument for its retention in the Ranales. No evidence has been found to preclude the inclusion of Dilleniaceae in the Ranales. On the other hand, as opposed to similarities in wood and pollen characters between Dilleniaceae and Theaceae, floral anatomical and embryological features offer a sharp contrast between the two. The Ranales are believed to be polyphyletic. It has been tentatively suggested that two major phyletic lines may be recognized in each of the woody and herbaceous series: the magnolialian and lauralian lines in the former and the nymphaealian and berberidalian lines in the latter.     

Evaluation of Hydrangea macrophylla for Resistance to Leaf‐Spot Diseases

Garden hydrangea (Hydrangea macrophylla) is a popular ornamental plant that can be devastated by leaf‐spot diseases. Information is needed to determine susceptibility of commercial cultivars to leaf‐spot diseases. To address this need, 88 cultivars of H. macrophylla were evaluated for their resistance to leaf‐spot diseases in full‐shade (2007–2008), full‐sun (2007–2008) and partial‐shade (2009–2010) environments in McMinnville, TN, USA. Ten cultivars [‘Ami Pasquier’, ‘Ayesha’, ‘Blue Bird’, ‘Forever Pink’, ‘Fuji Waterfall’ (‘Fujinotaki’), ‘Miyama‐yae‐Murasaki’, ‘Seafoam’, ‘Taube’, ‘Tricolor’ and ‘Veitchii’] were rated resistant (R) or moderately resistant to leaf spot under each of the three environments. In 2007–2008, approximately 51% of the cultivars were rated R in full shade, but only 5% were R in full sun. In 2009–2010, only 1% of the cultivars were rated R in partial shade. Although environmental parameters including temperature and rainfall influence disease severity and host reaction, a shaded environment was least favourable for leaf‐spot disease development, which demonstrates that establishing hydrangea in shaded environment can be an effective tool along with cultivar selection for managing leaf‐spot diseases on hydrangea. Six pathogens, Corynespora cassiicola, Cercospora spp., Myrothecium roridum, Glomerella cingulata (Anamorph: Colletotrichum gloeosporioides), Phoma exigua and Botrytis cinerea, were associated with leaf‐spot diseases of garden hydrangea. Of the leaf‐spot pathogens, C. cassiicola was most frequently isolated (55% of all isolates), followed by Cercospora spp. (20%) and other pathogens (25%). Because symptoms attributed to each leaf‐spot pathogen were similar, cultivars were selected for resistance to multiple leaf‐spot pathogens.     

Craniological differentiation between European wildcats (Felis silvestris silvestris), African wildcats (F. s. lybica) and Asian wildcats (F. s. ornata): implications for their evolution and conservation

Intraspecific diversification of the wildcat (Felis silvestris), including the European wildcat (F. s. silvestris), the Asian wildcat (F. s. ornata) and the African wildcat (F. s. lybica), was examined based on 39 cranial morphology variables. The samples of free‐ranging cats originated from Britain, Europe, Central Asia and southern Africa, consisting of both nominal wildcat specimens (referred to henceforth as ‘wildcats’) and nominal non‐wildcat specimens (‘non‐wildcats’) based on museum labels. The skull morphology of ‘wildcats’ from Britain and Europe is clearly different from that of ‘wildcats’ of Central Asia and southern Africa. The latter are characterized especially by their proportionately larger cheek teeth. On the basis of principal component, discriminant function and canonical variate analyses, the skull morphology of British ‘non‐wildcats’ is less distinct than is that of British ‘wildcats’ from the skull morphologies of ‘wildcats’ of Central Asia and southern Africa. On the other hand, the skull morphology of southern African ‘non‐wildcats’ is as distinct from those of ‘wildcats’ of Britain and Europe as is that of southern African ‘wildcats’. We suggest that the evolution of the modern wildcat probably consisted of at least three different distribution expansions punctuated by two differentiation events: the exodus from Europe during the late Pleistocene, coinciding with the emergence of the steppe wildcat lineage (phenotype of Asian–African wildcat), followed by its rapid range expansion in the Old World. The second differentiation event was the emergence of the domestic cat followed by its subsequent colonization of the entire world with human assistance. Considering the recent evolutionary history of, and morphological divergence in, the wildcat, preventing hybridization between the European wildcat and the domestic cat is a high conservation priority. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83 , 47–63.     

The holotypes of the upper Pleistocene Crocuta crocuta spelaea (Goldfuss, 1823: Hyaenidae) and Panthera leo spelaea (Goldfuss, 1810: Felidae) of the Zoolithen Cave hyena den (South Germany) and their palaeo‐ecological interpretation

The rediscovered holotype skulls of Late Pleistocene Panthera leo spelaea ( Goldfuss, 1810 ) (Felidae) and Crocuta crocuta spelaea ( Goldfuss, 1823 ) (Hyaenidae) from the Zoolithen Cave at Burggeilenreuth, southern Germany, are discussed. The cave became famous mainly due to its rich cave bear bone remains from the late Saalian (OIS 6–8) to Eemian/Weichselian (OIS 3–6) including additionally a third holotype of Ursus spelaeus Rosenmüller, 1794 (Ursidae). The ‘Felis spelaea’ holotype represents an adult male with a strong bite mark on the saggital crest, which was in an early stage of healing. Compared with other European Late Pleistocene lion skulls and skeletons, and with modern African lions, it provides evidence of intraspecific conflict between male Ice Age lions. The holotype of ‘Hyaena spelaea’ is one of several hundred hyena remains from a well‐frequented hyena den cave. The cave was used intensively by Late Pleistocene hyena clans, for collecting lion carcasses in addition to their accustomed prey, as happened in many caves throughout Europe. Ice Age spotted hyena clans might have killed Ice Age steppe lions for many reasons, such as fights over prey and territory, and the protection of cubs, but they did not always scavenge on their carcasses. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154 , 822–831.     

Vicariance and convergence in Magellanic and New Zealand long‐looped brachiopod clades (Pan‐Brachiopoda: Terebratelloidea)

Phylogenetic reconstructions using parsimony, maximum likelihood (ML), and Bayesian relaxed molecular clock analyses of ～2850 nucleotides of nuclear‐encoded small and large subunit ribosomal DNAs (SSU and LSU rDNAs) from 14 long‐looped (terebratelloid) ingroup and four short‐looped (terebratulidine) outgroup brachiopod taxa, together with ML analyses of ～663 nucleotides of mitochondrial cytochrome oxidase subunit 1 (cox1) from 12 terebratelloid taxa, show that deep divergence separates taxa endemic to waters in the vicinity of New Zealand from those with a Magellanic distribution around South America and Antarctica. This deep divergence also separates Magellanic Terebratella dorsata from New Zealand Terebratella sanguinea, showing that they are not congeneric. Instead, they belong to separate ‘Magellanic’ (MAG) and ‘New Zealand’ (NZ) clades that first diverged about 82 Mya (95% highest posterior density, 48–120 Mya), correlating with separation between the NZ and Antarctic tectonic plates. Sequence analyses also reveal (1) that the Antarctic endemic taxa Magellania fragilis and Magellania joubini are not congeneric with Magellania venosa, suggesting that their previous placement in Aerothyris should be restored, and (2) that divergence between Antarctic and NZ species of the terebratulide Liothyrella occurred much later than plate separation, perhaps because of continuing gene flow caused by long‐lived larvae. The topology of the rDNA and cox1 gene trees implies that radial ornament of the shell (‘ribbing’) has been gained (and/or lost) independently within the MAG and NZ clades. Radial ribs are widespread in articulate brachiopods throughout the Phanerozoic, but no comparisons of brachiopod rib morphology and morphogenesis have been published. Our comparisons of transverse shell mid‐sections in the scanning electron microscope reveal no obvious evidence of differences in morphology between independently gained ribs. We also consider several ways in which ribs may affect fitness, including effects on hydrodynamics. Only scanty and inconclusive evidence is available, but we suggest that effects (if any) are likely to be of small magnitude; adaptive value of brachiopod shell ribs remains to be demonstrated. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162 , 631–645.     

Trubs,but no trianas: filled and empty regions of angiosperm stem length‐diameter‐mechanics space

Discrete plant habit categories such as ‘tree’, ‘shrub’, and ‘liana’ belie continuous variation in nature. To study the evolution of this continuous variation, we gathered data on stem length, diameter and tissue mechanical stiffness across a highly morphologically diverse highland xerophytic scrub on a lava flow in central Mexico. With stem allometric and mechanical data from 1216 segments from 50 species, we examined relationships between stem length–diameter proportions and tissue mechanical stiffness using linear mixed‐effects models. Rather than a series of discrete clouds in stem length–diameter–tissue stiffness space, corresponding to traditional habit categories, the plants of this xerophytic scrub formed a single continuous one. Within this cloud, self‐supporting plants had stems that became predictably longer and tissues that became stiffer for a given diameter increase, and there was no paucity of intermediates between trees and shrubs (‘trubs’). Non self‐supporting plants had a steeper stem length–diameter slope and their tissues did not increase in stiffness with stem size. The area between self‐ and non self‐supporting plants was sparsely occupied as stem size increased. We predict that this ‘empty’ space between lianas and trees is developmentally accessible but of low fitness, meaning that there should be few ‘trianas’ in nature. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 361–373.     

Horticulture,hybrid cultivars and exotic plant invasion: a case study of Wisteria (Fabaceae)

Exotic Wisteria species are highly favoured for their horticultural qualities and have been cultivated in North America since the early 1800s. This study determines the identity, genetic diversity and hybrid status of 25 Asian Wisteria cultivars using plastid, mitochondrial and nuclear DNA data. Fifteen (60%) hybrid cultivars were identified. All of the ‘Wisteria sinensis’ cultivars sampled are hybrids with W. floribunda. Although W. sinensis and W. floribunda are recognized invasive species in the southeastern USA, the relationships of horticultural cultivars to naturalized plants was previously unknown. Haplotype analysis of nuclear data identifies four haplotypes shared between cultivated stock and naturalized populations in the southeastern USA. In addition, US invasive haplotypes are present in New Zealand‐derived cultivars although, to date, naturalized Wisteria has not been documented in New Zealand. Finally, these data are used to make recommendations to horticulturalists of select species cultivars which may be less likely to invade US landscapes. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 593–601.