Laticifer starch grain morphology and laticifer evolution in Euphorbia (Euphorbiaceae)

The micromorphology of starch grains in the nonarticulated laticifer was examined in 200 taxa of succulent and nonsucculent Euphorbia. Rod, spindle, osteoid, discoid and intermediate shapes were found in latex. These unusual starch grain shapes are assumed derivatives from more rounded grains present in other angiospermous cells. Unbranched, rod–shaped grains were present in latex of annual and perennial herbaceous taxa. Spindle, osteoid, and discoid shapes, often branched, occurred in xerophytic, succulent taxa. Leafy taxa in several subgenera possessed rod–shaped grains. Taxa of the primitive subgenus, Esula , possessed the shortest rod–shaped grains derived from rounded grains common in parenchyma. Length of rod–shaped grains increased through herbaceous subgenera and culminated in Poinsettia where grain length may be extremely great. Dwarf or shrubby succulents with thickened roots have osteoid grains. Shrubby or arborescent coralline and cactiform species, specialized taxa, possessed large branched grains. Grain morphology of Euphorbia was compared with that from latex of other genera of the Euphorbiaceae. These in–depth analyses of laticifer plastids demonstrates the applicability of starch grain morphology for interpreting the progressive evolution of the nonarticulated laticifer as a cell type within the genus Euphorbia.     

Functional andromonoecy in Euphorbia (Euphorbiaceae)

The occurrence of cyathia containing staminate flowers but lacking a pistillate flower was studied in 17 species of Euphorbia. Male cyathia were found in the majority of species studied (88.2%) giving functional andromonoecy. In the male cyathia, the pistillate flower is generally totally absent, but sometimes a vestigial pistillate flower with a non-functional ovary is present. The proportion of male cyathia varied at both the population and species level. The position of male cyathia within the inflorescence showed a constant pattern among species: the proportion of male cyathia decreased from the first to the last levels of the pleiochasia. In general, perennial species had significantly higher proportions of male cyathia than annual species (mean 20 and 2.3%, respectively). In annual species there was a trend for production of male cyathia only in the first level of the inflorescence, whereas in perennials production up to the fourth level of the inflorescence was usual. Functional andromonoecy is common in Euphorbia and represents a new sex segregation in the genus. The selective forces causing this secondary sex segregation in Euphorbia, such as improved pollination or increased outcrossing, are discussed.     

Isolation and partial characterization of latex lectins from three species of the genus Euphorbia (Euphorbiaceae)

Three N-acetylgalactosamine-specific lectins were isolated from the latices of Euphorbia calcina L., Euphorbia dalberi L. and Euphorbia sp. (an undetermined species) by affinity chromatography on fetuin-agarose. They are all glycoproteins [about 12.5% (w/w) carbohydrate] of M, around 140 000 and appear to be tetrameric molecules composed of different subunits. All three lectins have similar amino acid (with high contents of asparagine/aspartic acid, giycine and leucine) and carbohydrate (with glucosamine, mannose.) fucose and xylose) compositions. In addition. they are closely related serologically.     

Two new species of Euphorbia subsp. Euphorbia (Euphorbiaceae) from east and northeast Somalia

Two small species of succulent, spiny Euphorbia are described as new, both with limited distributions, E. ammophila from the coastal plain of eastern Somalia, and E. densispina from the mountain ranges of the northeast.     

Phylogenetics and evolution of phyllotaxy in the Solomon's seal genus Polygonatum (Asparagaceae: Polygonateae)

Polygonatum is the largest and most complex genus in tribe Polygonateae, comprising approximately 57 species widely distributed in the warm temperate, subtropical and boreal zones of the Northern Hemisphere. However, phylogenetic relationships in the genus remain poorly understood. The objectives of this study were to reconstruct the phylogenetic relationships of the genus using four plastid markers, and to examine the evolution of leaf arrangement in Polygonatum in the phylogenetic context of its closely related taxa. Thirty Polygonatum species were sampled to infer phylogenetic relationships using maximum‐likelihood and Bayesian analyses. The evolution of leaf arrangements was reconstructed using Bayesian, parsimony and likelihood methods. The phylogenetic analyses supported the current generic delimitation of Polygonatum, with Heteropolygonatum recognized as a distinct genus. Three major lineages in Polygonatum were well supported, largely correlated with geographical distribution and the most recent classification at the sectional level. However, our results did not support the currently recognized series, especially the two large series Verticillata and Alternifolia. Bayesian analyses support the alternate‐leaf arrangement as the ancestral state for Polygonatum, but parsimony and maximum‐likelihood analyses suggest an equivocal state for crown Polygonatum. Leaf arrangement was found to be evolutionarily labile. A new nomenclatural combination was made: P olygonatum section S ibirica (L.I.Abramova) Y.Meng, comb. nov. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176 , 435–451.     

The origins and evolution of the genus Myosotis L. (Boraginaceae)

Although morphologically well defined, the phylogeny and taxonomy of Myosotis has been uncertain. In particular it has been unclear whether the genus had a Northern Hemisphere or Australasian origin. However, separate analyses of the ITS and the 3' region of matK, as well as a combined analysis of ITS, 3' matK, the psbA-trnA spacer, and 3' ndhF regions indicate that several distinct lineages exist within Myosotis and strongly support a Northern Hemisphere origin for the genus. Further, the observed transoceanic distributions and levels of genetic divergence between lineages indicate that long distance dispersal has been important for establishing the current geographic range expansion of Myosotis. Our molecular data also suggest that the diversification of Australasian Myosotis has occurred since the late Tertiary and is largely due to radiation within and from New Zealand. This inference is consistent with the findings of recent phylogenetic studies on other New Zealand alpine genera. Our results highlight the important role played by late Tertiary and Quaternary climate change in explaining current floristic diversity. The genetic relationships reported here also suggest that the current infrageneric taxonomy of Myosotis does not fully reflect the evolution of the genus.     

New pair-spined species of Euphorbia (Euphorbiaceae) from Somalia

Eighteen new species of Euphorbia subgenus Euphorbia are described and their relationships discussed. All are endemic to Somalia, with limited distributions in the northern Regions. One, E. phillipsioides , has been identified mistakenly for some time as E. phillipsiae , which is shown to be conspecific with E. golisana. E. xylacantha , considered by N. E. Brown to be synonymous with E. monacantha , is reinstated as a distinct but variable species.     

Phylogenetics and reticulate evolution in Pistacia (Anacardiaceae)

The systematic position and intrageneric relationships of the economically important Pistacia species (Anacardiaceae) are controversial. The phylogeny of Pistacia was assessed using five data sets: sequences of nuclear ribosomal ITS, the third intron of the nuclear nitrate reductase gene (NIA-i3), and the plastid ndhF, trnL-F and trnC-trnD. Significant discordance was detected among ITS, NIA-i3, and the combined plastid DNA data sets. ITS, NIA-i3, and the combined plastid data sets were analyzed separately using Bayesian and parsimony methods. Both the ITS and the NIA-i3 data sets resolved the relationships among Pistacia species well; however, these two data sets had significant discordance. The ITS phylogeny best reflects the evolutionary relationships among Pistacia species. Lineage sorting of the NIA-i3 alleles may explain the conflicts between the NIA-i3 and the ITS data sets. The combined analysis of three plastid DNA data sets resolved Pistacia species into three major clades, within which only a few subclades were supported. Pistacia was shown to be monophyletic in all three analyses. The previous intrageneric classification was largely inconsistent with the molecular data. Some Pistacia species appear not to be genealogical species, and evidence for reticulate evolution is presented. Pistacia saportae was shown to be a hybrid with P. lentiscus (maternal) and P. terebinthus (paternal) as the parental taxa.     

Preliminary results on the ichthyocidal properties of Euphorbia ingens (Euphorbiaceae)

Euphorbia ingens belongs to the family Euphorbiaceae, which includes a variety of latex-producing plants, the majority of them having ichthyocidal properties (Coates-Palgrave 2000). Tests to determine the degree of virulence of E. ingens latex used Cyprinus carpio and Xenopus laevis as the main test organisms. Fish and frogs were exposed to various concentrations of latex to determine the concentration lethal only to the fish. The same concentrations were then used to determine the stability of the latex, by exposing fish in tanks with and without biological filtration. The breakdown period varied, with it breaking down quicker with biological filtration than without it. Once the stability of the latex within a system was determined, crabs, aquatic snails, frogs and fish were all simultaneously tested in a tank at the same concentration as before. Within 12 hours all the fish and half the frogs died, whereas the crabs and snails appeared not to suffer any detrimental effects. The poison degraded and became harmless to fish within 48 hours. Thus, the latex of Euphorbia ingens is a temporarily effective ichthyocide when applied in the correct concentrations.     

Chromosome and breeding system evolution of the genus Mercurialis (Euphorbiaceae): implications of ITS molecular phylogeny

 The internal transcribed spacers (ITS1 and ITS2) of nuclear ribosomal DNA were amplified and sequenced from 19 samples representing all species of the genus Mercurialis and two outgroup species, Ricinus communis and Acalypha hispida. The length of ITS1 in the ingroups ranged from 223 to 246 bp and ITS2 from 210 to 218 bp. Sequence divergence between pairs of species ranged from 1.15% to 25.88% among the ingroup species in the combined data of ITS1 and ITS2. Heuristic phylogenetic analyses using Fitch parsimony on the combined data of ITS1 and ITS2 with gaps treated as missing generated 45 equally parsimonious trees. The strict consensus tree was principally concordant with morphological classification. Within the genus, the ITS sequences recognised two main infrageneric clades: the M. perennis complex including three Eurasian stoloniferous species (M.␣leiocarpa, M. ovata and M. perennis) and the western Mediterranean group including eight both annual and perennial species. Of the western Mediterranean clade, the annual and perennial species grouped respectively into two different groups, and the annual life form is revealed as a synapomorphic character derived from perennial, whereas in the Eurasian clade ITS phylogeny suggested M. leiocarpa as basal clade sister to M.␣perennis and M. ovata. ITS phylogeny failed to resolve the relationships among the different cytotypes of M. ovata and M. perennis. ITS phylogeny also suggested rapid karyotypic evolution for the genus. The karyotypic divergence among the perennial species of western Mediterranean region did not corroborate the nucleotide sequence divergence among the species. Optimisation of chromosome numbers onto the ITS phylogeny suggested x=8 to be the ancestral basic chromosome number of the genus. ITS phylogeny confirmed that the androdioecy of M. ambigua is derived from dioecy. The nucleotide heterozygosity and additivity in ITS sequences clearly confirm the interspecific hybridisation in the genus Mercurialis. Received December 22, 2001; accepted May 21, 2002?Published online: November 14, 2002 Address of the authors: Martin Kr?henbühl, Yong-Ming Yuan (correspondence) and Philippe Küpfer, Institut de Botanique, Laboratoire de botanique évolutive, Université de Neuchatel, Emile-Argand 11, CH-2007 Neuchatel, Suisse. (e-mail: yong-ming.yuan@unine.ch)     

New herbaceous and woody species of Euphorbia (Euphorbiaceae) from Somalia

A small prostrate herb, Euphorbia salsicola , a woody shrublet with curious tubers, E. scatorhiza , and a small tree with decorative bracts, E. thulinii , are described as new species of Euphorbia. Within subgenus Lyciopsis the morphological variability of E. cuneata in Somalia is discussed and a new subspecies cretacea is described. E. gypsophila , a closely related but distinct species found only on gypsum is also described as new, together with E. dunensis , a shrublet from the coastal dunes and E. tripartita , another shrub in the same subgenus related to E. matabelensis. As far as present records show, all the new taxa are endemic in Somalia.     

The occurrence of C(2) photosynthesis in Euphorbia subgenus Chamaesyce (Euphorbiaceae)

This study investigated whether Euphorbia subgenus Chamaesyce subsection Acutae contains C(3)-C(4) intermediate species utilizing C(2) photosynthesis, the process where photorespired CO(2) is concentrated into bundle sheath cells. Euphorbia species in subgenus Chamaesyce are generally C(4), but three species in subsection Acutae (E. acuta, E. angusta, and E. johnstonii) have C(3) isotopic ratios. Phylogenetically, subsection Acutae branches between basal C(3) clades within Euphorbia and the C(4) clade in subgenus Chamaesyce. Euphorbia angusta is C(3), as indicated by a photosynthetic CO(2) compensation point (Г) of 69 μmol mol(-1) at 30 °C, a lack of Kranz anatomy, and the occurrence of glycine decarboxylase in mesophyll tissues. Euphorbia acuta utilizes C(2) photosynthesis, as indicated by a Г of 33 μmol mol(-1) at 30 °C, Kranz-like anatomy with mitochondria restricted to the centripetal (inner) wall of the bundle sheath cells, and localization of glycine decarboxlyase to bundle sheath mitochondria. Low activities of PEP carboxylase, NADP malic enzyme, and NAD malic enzyme demonstrated no C(4) cycle activity occurs in E. acuta thereby classifying it as a Type I C(3)-C(4) intermediate. Kranz-like anatomy in E. johnstonii indicates it also utilizes C(2) photosynthesis. Given the phylogenetically intermediate position of E. acuta and E. johnstonii, these results support the hypothesis that C(2) photosynthesis is an evolutionary intermediate condition between C(3) and C(4) photosynthesis.     

Explosive seed dispersal in two perennial Mediterranean Euphorbia species (Euphorbiaceae)

The distance of explosive dispersal, its pattern in time, and the relative importance of autochory have been studied in two diplochorous species: Euphorbia boetica and E. nicaeensis. The seeds of E. boetica released by explosive dispersal reached a median distance of 156 cm and a maximum of almost 8 m, while the distances reached by the seeds of E. nicaeensis were lower: a median of 132 cm and a maximum of 5 m. The differences in explosive dispersal distance between species seem to depend on both seed mass and caruncle retention. The seeds of both species present a caruncle, but in E. boetica this is tiny, and in most cases is shed during the explosion of the capsules. The distances reached by the seeds of these species, dispersed just by capsule explosion, were similar to or greater than the distances to which ants disperse seeds in the Mediterranean sclerophyllous vegetation. Diplochorous plants may maximize either the distance of primary dispersal or that of secondary dispersal. Given that the seeds of E. boetica, that lose their caruncles, are not gathered by myrmecochorous ants, the results suggest that E. boetica maximizes its primary dispersal distance, whereas E. nicaeensis favors its secondary dispersal.     

Chromosome evolution in the genus Ophioglossum L.

Cytological observations on eleven species of Ophioglossum revealed low gametic ( n ) chromosome numbers of 30, 34 and 60 in populations of O.eliminatum , contrasting with an earlier report of n = 90 in the same species. The rest of the species is based on n =120.Cytologically studied species of Ophioglossum exhibit a range of chromosome numbers from n = 30 in O.eliminatum to n =720 in O.reticulatum. The weighted highest common factor (HGF) from all the reported chromosome numbers in twelve species was found to be 30. This number is proposed as the palaeobasic chromosome number for the genuS. Reported chromosome numbers which are not multiples of 30 were subjected to sequential analysis, yielding three distinct ultimate base numbers, 4, 5 and 6, which can produce n = 30 in seven different ways. The neobasic number, n= 120, appears to have arisen through various combinations and permutations of these, theoretically 2401 routes; only a relatively few of these routes exist today, suggesting that extreme selection has been exerted against the majority, and further suggesting that Ophioglossum represents an evolutionary dead end through repeated cycles of polyploidy and is possibly at the verge of extinction. The stoichiometric model of evolution, which derives the various chromosome numbers possessed by the twelve species from the basic and ultimate basic chromosome numbers, is used to explain chromosomal evolution in the genus.     

Phylogenetics of Puya (Bromeliaceae): Placement, major lineages, and evolution of Chilean species

Puya (Bromeliaceae), a large genus of terrestrial bromeliads found throughout a range of elevations in the Andes and central Chile, is of great systematic, evolutionary, and biogeographical interest. This first molecular phylogenetic study of Puya and related bromeliads employs matK, trnS-trnG, rps16, and PHYC sequences. Chloroplast DNA, nuclear DNA, and combined DNA data all place Puya closest to subfamily Bromelioideae. Nuclear and combined data support Puya as monophyletic, and the two subgenera are nonmonophyletic. All data indicate that the Chilean species of Puya are early diverging within the genus, consistent with Chilean genera as the first-diverging members of subfamily Bromelioideae. Central Chile is identified as a key region for understanding the biogeographical history of Bromeliaceae, as is true with other South American plant groups. A complicated history involving early chloroplast capture and later secondary hybridization and/or introgression is seen in Chilean lineages. These events help explain the occurrence of sterile inflorescence tips, floral color and shape, and leaf indument. The ecological radiation of Puya appears coincident with the final, recent rise of the Andes and subsequent high-elevation habitat diversification. Additionally, geographical distribution, rather than moisture or elevational adaptations, correlates to species relationships. Evolution of CAM photosynthesis has occurred multiple times.     

Seed production, predation and dispersal in the Mediterranean myrmecochore Euphorbia characias (Euphorbiaceae)

Seed production of a population of 193 reproductive individuals of Euphorbia characias from the north-east Iberian peninsula was estimated at 155 000 seeds per year The dehiscence period lasts for four weeks The timing of ballistic dispersal is uniformly distributed throughout the day Seeds remain on the surface a mean time (± SE) of 52.3 ± 6.7 mm Several and species take the seeds in varying proportions Messor barbarus (0 225), a granivorous ant which eats the seeds, Aphaenogaster senilis (0 313), Pheidolc pallidula (0 296) and Tapinoma nigerrimum (0 162) which take the seeds to their nests but do not eat them Half a million ant foraging trips per day indicate a very strong short term influence of ants on fate of Euphorbia seeds Ants as a group have a dual role of predation and dispersal of Euphorbia characias seeds The characteristics of this Euphorbia (sclerophylly, diplochory, firmness of elaiosome, absence of subsidiary myrmecochorous features) better fit the syndrome described for Australian myrmecochores than the one proposed for European and North American myrmecochores