A synopsis of Goniothalamus species (Annonaceae) in Peninsular Malaysia, with a description of a new species

The species of Goniothalamus (Annonaceae) occurring in Peninsular Malaysia and Singapore are revised, updating the previous taxonomic treatment by James Sinclair, published in 1955. A total of 18 species is recognized, including a new species, G. tomentosus . Collections referrable to G. tomentosus were previously determined as ' G. marcanii '; examination of the types of G. marcanii reveals that the two taxa are not conspecific, however, and a new name is accordingly validated here. Goniothalamus tomentosus has closest affinities with the Javan/Sumatran species G. costulatus . Other important nomenclatural changes include the reduction of G. umbrosus to synonymy with G. tapis .  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 142 , 321−339.     

Floristics of the Dunbar Valley serpentinite site, Songimvelo Game Reserve, South Africa

The Dunbar Valley serpentinite outcrop has a flora comprising 254 taxa (species and below) in 172 genera and 63 families. Dunbar Valley has more species than other studied serpentinite sites in the Barberton Greenstone Belt (BGB). The genus Senecio is the most speciose genus in the BGB. The level of species endemism at Dunbar Valley is 2.0%. Most of the serpentinite endemics in the BGB show phytogeographical affinities with the Sudano-Zambezian Region. Six modified-Whittaker plots, three on serpentinite and three on non-serpentinite soils, were sampled. Sørenson's index was 0.312, indicating low similarity in species between serpentinite and non-serpentinite sites (β-diversity) at Dunbar Valley, as at other sites in the BGB. α-diversity (using the Shannon–Wiener index) for the serpentinite was 2.631 ± 0.901 and for the non-serpentinite, 2.886 ± 0.130. However, a Student's t -test showed no significant difference in α-diversity between the two habitats. There was also no significant difference in species richness between serpentinite and non-serpentinite sites. Total species showed negative correlations with total nickel, altitude and serpentinite outcrop size (area) for six sites in the BGB. Number of endemic taxa showed no correlation with environmental variables.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 271–285.     

Analysis of morphological variation of the Acer tschonoskii complex in eastern Asia: implications of inflorescence size and number of flowers within sect. Macrantha

Flower and fruit specimens of 184 individuals were sampled to investigate patterns of intraspecific variation and to evaluate recognition of taxa within the Acer tschonoskii complex using morphometric analysis. Previous taxonomic treatments have considered A. tschonoskii var. rubripes (=  A. komarovii ) and A. tschonoskii var. tschonoskii to be separate species. The morphological discontinuity between these two taxa was evident in peduncle and pedicel length, and in number of flowers. In addition, the delimitations of some species within sect. Macrantha were clarified using these diagnostic characters. In view of the geographical distribution of the A. tschonoskii complex, which includes many taxa of sect. Macrantha from China to Japan through Korea, the long raceme with many flowers ( A. sikkimense ) and unlobed leaf are considered more primitive than the short raceme with a small number of flowers and five-lobed leaf ( A. maximowiczii and A. komarovii ). However, many intermediate taxa were present. This study also suggests that several Chinese taxa, such as A. metcalfii, A. taronense, A. hookeri and A. grosseri , may be subject to different taxonomic interpretation and should be reinvestigated morphologically.  © 2003 The Linnean Society of London. Botanical Journal of the Linnean Society, 2003, 143 , 29−42.     

Pollinator sharing in dioecious figs (Ficus: Moraceae)

As one of the most specialized pollination syndromes, the fig (Ficus)–fig wasp (Agaonidae) mutualism can shed light on how pollinator behaviour and specificity affect plant diversification through processes such as reproductive isolation and hybridization. Pollinator sharing among species has important implications for Ficus species delimitation and the evolutionary history of the mutualism. Although agaonid wasp pollinators are known to visit more than one host species in monoecious figs, pollinator sharing has yet to be documented in dioecious figs. The present study investigated the frequency of pollinator sharing among sympatric, closely‐related dioecious figs in Ficus sections Sycocarpus and Sycidium. Molecular and morphological species identification established the associations between pollinating agaonid wasp species and host fig species. Cytochrome oxidase I was sequenced from 372 Ceratosolen pollinators of Ficus section Sycocarpus and 210 Kradibia pollinators of Ficus section Sycidium. The association between fig species and morphologically distinct clades of pollinator haplotypes was predominantly one‐to‐one. In Ceratosolen, six of 372 pollinators (1.5%) visited fig species other than the predominant host. No pollinator sharing was detected between the two Sycidium host species, although a rare hybrid shared Kradibia pollinators with both parental species. These findings point to low rates of pollinator sharing among closely‐related dioecious fig species in sympatry, and perhaps lower rates than among monoecious figs. Such rare events could be evolutionarily important as mechanisms for gene flow among fig species. Differences in rates of pollinator sharing among fig lineages might explain the conflicting phylogenetic patterns inferred among monoecious figs, dioecious figs, and their respective pollinators. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 546–558.     

An inquiline fig wasp using seeds as a resource for small male production: a potential first step for the evolution of new feeding habits?

The processes allowing evolutionary transitions in resources used by parasitic wasps are largely unknown. Microhymenopteran communities associated with figs could provide a model system to investigate such transitions. We investigate here a species of genus Idarnes . The larvae generally develop as inquilines of the pollinating wasp larvae. However, in figs where the parasitic pressure is high, eggs are laid in developing seeds. These eggs turn into small males. This is the first report of seed consumption by a fig wasp. Using an alternative resource to produce small males could provide a pathway to select for increased ability to use this resource and hence provide an intermediate step for evolving the capacity to use new resources.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 9–17.     

Ecological diversity and adaptive tendencies in the tropical fern Trichomanes L. (Hymenophyllaceae) with special reference to climbing and epiphytic habits

Among the basal fern families, the Hymenophyllaceae, with more than 600 species, display a high diversity in terms of their morphology and the habitats that they occupy. We have chosen to focus on Trichomanes L., a clearly defined genus for which a phylogeny is presently being developed, to investigate the appearance of the climbing and epiphytic habits, as well as the related supposed adaptive characters. In this study we present the first review of the different ecological types within the genus: terrestrial, climbing (divided into hemi-epiphytic forms and true lianas), and epiphytic types. The study of several features concerning stem morphology and leaf size allows a proposal on relationships between ecology and plant morphology. Terrestrial species display a thick monocaulous rhizome with robust roots and short internodes. Climbing species are characterized by a branched, thick, creeping rhizome with long internodes. Epiphytic species also exhibit long, creeping and branching stems with long internodes but the rhizome is fine to filiform. Under these circumstances, there is a reduction of root system and frond size leading to dwarfism in numerous instances. This may be related to an extreme hygrophilous epiphytic strategy. Finally, hypotheses on the evolution of these habits and hence on the evolutionary relationships between ecology and characters are presented and discussed.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 41–63.     

Phylogenetic diversity of forest trees in the Usambara mountains of Tanzania: correlations with altitude

The relationship between diversity of large trees and altitude was investigated in a Tanzanian tropical rain forest. In total, 231 samples of 20 trees of ≥ 20 cm d.b.h. from the East and West Usambara mountains, covering an elevation range from 280 m to 2180 m a.s.l., were analysed. An ordination demonstrated a constant turnover of species, genera, family and orders with elevation. There were no obvious zones or discontinuities. There was no decline in plot richness with respect to altitude for species, genera or orders. Family richness was shown to increase with altitude. A measure of genetic diversity, the avalanche index, was calculated for each plot to investigate the effect of incorporating phylogenetic relatedness of individuals into the diversity measure. Distances between taxa were extracted from a recent molecular phylogeny of the angiosperms. Incorporation of phylogenetic diversity at family level enhanced the positive correlation between plot diversity and altitude.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 149 , 217–228.     

Facultative apomixis and hybridization in arctic Potentilla section Niveae (Rosaceae) from Svalbard

The degree of sexual seed set and the ability to cross were investigated in three taxa of Potentilla section Niveae ( P. chamissonis , P. insularis and P. nivea ) from the Svalbard Archipelago. Emasculated and bagged flowers had little or no seed set, while 71% of the emasculated, bagged and pollinated flowers had some. The taxa are interpreted as pseudogamous apomicts. Parental plants and offspring were subjected to isozyme analysis. Most of the offspring were clones of the mother plant, but 1.7% had bands from the pollen donor. The sexual offspring were all produced by mother plants of P. insularis , which is interpreted as a facultative apomict. Surprisingly, crosses between different taxa gave higher seed to ovule ratios and numbers of sexually produced offspring than crosses within taxa. Some of the sexually produced hybrid offspring had the same kind of hairs on the petioles as their mother taxon, showing that hybrids may not be intermediate in hair characters, which are considered important in the delimitation of these taxa. The results of the present study indicate very close relationships among the investigated plants.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 373−381.     

Resistance to temperature extremes in sub-Antarctic weevils: interspecific variation, population differentiation and acclimation

Much of the work on the responses of terrestrial arthropods to high and low temperatures has been done on model organisms such as Drosophila . However, considerable variation in thermotolerance is partitioned at the family level and above, raising questions about the broader applicability of this work to other taxa. Here we investigate resistance to high and low temperatures, following different temperature treatments, in ten species and 31 populations of weevils found on sub-Antarctic Heard Island and Marion Island, which have substantially different climates. In these weevils there is considerable interspecific and among-population variation in critical thermal minimum (CTmin) and critical thermal maximum (CTmax), but most of this variation in critical limits can be ascribed to phenotypic plasticity. We find no relationship between CTmin and CTmax at the species level, and this is true also of populations and of responses to the temperature treatments. In general, plastic (acclimation) changes in CTmin are larger than those in CTmax. Our data therefore provide support for the idea that resistance to heat and to cold are decoupled in terrestrial arthropods. Furthermore, our results suggest that investigations of physiological limits to species borders should incorporate the effects of phenotypic plasticity on physiological capabilities.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 78, 401–414.     

A new species in the genus Dioon (Zamiaceae) from north-central Oaxaca, Mexico

Dioon argenteum sp. nov. (Zamiaceae) is described from northern Oaxaca, México. Flat leaves, and persistently tomentose, slightly imbricate leaflets with marginal prickles characterize this species. The specific epithet argenteum was chosen to describe the silver appearance of the persistent tomentum covering the new leaves. D. argenteum appears to have affinities with D. purpusii and D. califanoi.   © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141 , 471–476.     

How to be an ant on figs

Mutualistic interactions are open to exploitation by one or other of the partners and a diversity of other organisms, and hence are best understood as being embedded in a complex network of biotic interactions. Figs participate in an obligate mutualism in that figs are dependent on agaonid fig wasps for pollination and the wasps are dependent on fig ovules for brood sites. Ants are common insect predators and abundant in tropical forests. Ants have been recorded on approximately 11% of fig species, including all six subgenera, and often affect the fig–fig pollinator interaction through their predation of either pollinating and parasitic wasps. On monoecious figs, ants are often associated with hemipterans, whereas in dioecious figs ants predominantly prey on fig wasps. A few fig species are true myrmecophytes, with domatia or food rewards for ants, and in at least one species this is linked to predation of parasitic fig wasps. Ants also play a role in dispersal of fig seeds and may be particularly important for hemi-epiphytic species, which require high quality establishment microsites in the canopy. The intersection between the fig–fig pollinator and ant–plant systems promises to provide fertile ground for understanding mutualistic interactions within the context of complex interaction networks.     

Little genetic differentiation across Europe between early-flowering and late-flowering populations of the rapidly declining orchid Neotinea ustulata

Amplified fragment length polymorphism (AFLP) was used to characterize genetic diversity of the endangered Burnt Orchid, Neotinea (formerly Orchis ) ustulata . Fingerprinting of Estonian and British populations revealed surprisingly little genetic differentiation between populations but larger amounts of diversity within populations, especially in Britain. The resulting mean F _st value of 0.51 is unusually high for an orchid species. Much of the variation follows a west–east cline across Europe, whereas the much-discussed early- and late-flowering taxa of N. ustulata are considered insufficiently distinct to be viewed as separate subspecies. The later flowering N. ustulata var. aestivalis probably evolved independently on two or three occasions, each time diverging from the earlier flowering nominate race. The identity of the genes underpinning phenology in the species, and the potential selective advantages of phenological divergence, merit further study. Overall genetic diversity within populations is sufficiently high to render impoverishment an unlikely cause of their recent, precipitous decline.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 13–25.     

Evolutionary and ecological significance of Lepidaster grayi, the earliest multiradiate starfish

Lepidaster grayi Forbes, 1850, from the Much Wenlock Limestone Formation (Silurian: Wenlock) of England, is the earliest species of starfish (Echinodermata: Asteroidea) to deviate from pentaradial symmetry, having 13 rays rather than five. Based on the patterns of supernumerary ray development seen in extant multiradiate asteroids, two possible models are evaluated for the origin of the eight additional rays seen in L. grayi . In the 'all-in-one' model, all rays were added in the same interradius, whereas in the 'quadrants' model generations of rays would have been added in each of four interradii. The smallest specimen of L. grayi , apparently having only nine rays, suggests that the 'quadrants' model is most probable for the species. The presence of supernumerary rays in Silurian starfish, coupled with the existence of numerous other Palaeozoic multiradiate taxa, shows that asteroids have been able to deviate from pentamerism for most of their evolutionary history, and the variety of methods of supernumerary ray addition indicates that the multiradiate condition is homoplastic. The ecological significance of multiradiate Palaeozoic starfish is reviewed: the mouth frame of L. grayi had considerably greater flexibility than that of contemporaneous five-rayed species and, in combination with its supernumerary rays, enabled L. grayi to manipulate and consume larger food items. It is probable that Silurian starfish utilized a similar range of trophic guilds as those exploited by extant taxa.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 150 , 743–754.     

A new diploid Hieracium (Asteraceae: Lactuceae) from Bulgaria

Hieracium kittanae Vladimirov, a new diploid (2 n  = 18) species restricted to a few gorges in the Central Rhodope Mountains, south Bulgaria, is described and illustrated. It occurs in crevices of limestone rock in shady to sunny places with high atmospheric humidity. Morphologically, it to some extent resembles taxa from the H. glaucinum and H. schmidtii groups, but is otherwise very distinct, without close relatives in the Rhodope and neighbouring mountains.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003 , 143 , 213−218.     

Genetic relatedness assessment through individual odour similarities in mice

Previous research has demonstrated that closer genetic relatedness between individuals (from kin to across species) is associated with greater similarity in the qualities of their individual odours ('odour-genes covariance'). This predictable relationship between individual genotypes and individual odours could enable animals to assess their degree of genetic relatedness to other individuals by comparing the degree of similarity between another individual's odour and their own odour. In two-choice tests with odours of unfamiliar mice from different populations and species, subjects from two species of wild mice, Mus spicilegus and M. musculus , that had been raised in mixed litters of both species spent significantly more time investigating the ano-genital odour of the more genetically similar individual. This differential interest was not affected by common rearing with heterospecifics. These responses are consistent with a self-referencing mechanism enabling differential responses across a wide spectrum of genetic relatedness from kin through populations to heterospecifics. These assessments depend on the degree of similarity between the donor's and the subject's odours rather than on differences between them. Such a parsimonious mechanism may provide a basis for differential responses to conspecifics as opposed to heterospecifics that may function as a premating isolating mechanism.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 78, 595–603.     

Some remarks on the genus Astragalus sect. Incani in Iran

New findings are given for the Astragalus sect. Incani , which occurs within the area covered by the Flora of Iran. Astragalus wojciechowskianus Ranjbar is described as a new species. A. procerus Boiss. & Hausskn . , A. cottonianus Aitch. & Baker, A. achundovii Grossh., A. orduabadensis Grossh. and A. glaucophyllus Bunge are reported as new records for the flora of Iran .   © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 443–447.     

Genetic variation within and between Iliamna corei and I. remota (Malvaceae): implications for species delimitation

Iliamna corei and I. remota are classified as endangered species; however, their designation as separate species has been questioned. In order to address this problem, intersimple sequence repeat (ISSR) data were generated to examine patterns of genetic differentiation within and between these two taxa. ISSRs were used to screen individuals for genetic diversity of I. corei from the single known natural population and two garden populations, and individuals of I. remota from the six natural populations and four garden populations. Using ten primers, 140 informative markers were generated. Ninety-four percent of loci detected revealed polymorphisms. Cluster analysis (neighbour-joining, NJ) revealed that the two species are genetically distinct and that the Illinois populations of I. remota were genetically distinct from the Virginia populations of I. remota . Ordination by Principal Coordinate Analysis (PCoA) supported the findings of the NJ, with a separation of I. corei and I. remota . Analysis of Molecular Variance (AMOVA) revealed that the majority of variation detected was within populations, which is consistent with other self-incompatible plants. The results indicate a correlation between the geographical distributions of the species and gene flow.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 345–354.     

Polyploidy and karyotype variation in Turkish Bellevalia (Hyacinthaceae)

There are 20 Bellevalia species in Turkey, half of which are endemic. Chromosome numbers are known for 15 species. A chromosome survey of 145 Bellevalia individuals showed that the karyotype is remarkably stable. All are based on x  = 4. The majority are diploid with 2 n  = 8, but there is also a polyploid series of 2 n  = 16, 24 and 32. Aneuploidy occurs only at the octoploid level. Eleven individuals had metacentric B chromosomes, one had acrocentric Bs and one had telocentric Bs. Bellevalia pycnantha and B. paradoxa are morphologically similar, with B. pycnantha reduced to a synonom of B. paradoxa .  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 87–98.     

Avoiding injury and surviving injury: two coexisting evolutionary strategies in lizards

Congenital morphological differences between injured and intact individuals in a population may reflect adaptations to avoid injury, to survive injury, or both. We explore the possible occurrence of such adaptations, analysing the relationship between tail state (original, O; regenerated, R) and morphology (20 scale characters) in 23 taxa of Lacertidae. In some taxa, such as Acanthodactylus opheodurus , morphologies of O and R lizards differed significantly. In these, usually O individuals were morphologically typical (modal), while R individuals were extreme; in others, the contrary occurred. Indices of pattern similarity detected fewer differences between O and R lizards than those based on absolute values. We developed unbiased estimates of classification rates of discriminant analysis. The order of inclusion of characters in the discriminant functions of males and females of the same species differed more than the order for the same sex in other species. Some morphological differences between O and R lizards seem adaptive or linked by pleiotropy to adaptive traits. Congenital morphological differences between O and R lizards are probably more frequent than detected because we show that age and geographical heterogeneity of our samples are confounding factors to O–R differences. R-morphologies might reflect microevolution in favour of paedotypic forms, possibly possessing greater regenerating capacities than relatively more peratypic forms.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 78 , 307–324.