text:
Plant a shrub or tree; true roots +, origin endogeneous, root cap +, apex multicellular; endodermis +; shoot apical meristem multicellular; lateral meristems +, cork cambium producing cork abaxially, vascular cambium producing phloem abaxially and xylem adaxially; lamina with mean venation density 1.8 mm/mm2 (to 5 mm/mm2).
Plant woody, evergreen; nicotinic acid metabolised to trigonelline, (cyanogenesis via tyrosine pathway); primary cell walls rich in xyloglucans and/or glucomannans, 25-30% pectin [Type I walls]; lignins derived from (some) sinapyl and particularly coniferyl alcohols, thus containing p-hydroxyphenyl and guaiacyl lignin units [so no Maüle reaction]; root xylem exarch, cork cambium deep seated; arbuscular mycorrhizae +; shoot apical meristem interface specific plasmodesmatal network; stem with vascular tissue around central pith [eustele], vascular bundles with interfascicular tissue, ectophloic, endodermis 0, xylem endarch; wood homoxylous, tracheids and rays alone, tracheid/tracheid pits circular, bordered; mature sieve tube/cell lacking functioning nucleus, sieve tube plastids with starch grains; phloem fibres +; stem cork cambium superficial; branches exogenous; leaves with single trace from vascular sympodium ["nodes 1:1"]; vascular bundles collateral [stem: phloem external; leaf: phloem abaxial]; stomata morphology?, pore opening in response to leaf hydration active, control by abscisic acid, metabolic regulation of water use efficiency, etc.; leaves with petiole and lamina, spiral, development basipetal, blade simple; axillary buds +, not associated with all leaves; prophylls two, lateral; plant heterosporous, sporangia borne on sporophylls; microsporophylls aggregated in indeterminate cones/strobili; true pollen +, grains mono[ana]sulcate, exine and intine homogeneous; ovules unitegmic, parietal tissue 2+ cells across, megaspore tetrad tetrahedral, only one megaspore develops, megasporangium indehiscent; male gametophyte development first endo- then exosporic, tube developing from distal end of grain, to ca 2 mm from receptive surface to egg, gametes two, developing after pollination, with cell walls, flagellae numerous; ovules increasing considerably in size between pollination and fertilization, female gametophyte endosporic, initially syncytial, walls then surrounding individual nuclei; seeds "large" [ca 8 mm3], but not much bigger than ovule, with morphological dormancy; embryo cellular ab initio, endoscopic, plane of first cleavage of zygote transverse, suspensor +, short-minute, embryo straight, shoot and root at opposite ends [allorrhizic], white, cotyledons 2; plastid transmission maternal; ycf2 gene in inverted repeat, two copies of LEAFY gene, PHY gene duplications [three - [BP [A/N + C/O]] - copies], nrDNA with 5.8S and 5S rDNA in separate clusters; mitochondrial nad1 intron 2 and coxIIi3 intron and trans-spliced introns present.
Lignans, O-methyl flavonols, dihydroflavonols, triterpenoid oleanane, non-hydrolysable tannins, quercetin and/or kaempferol +, apigenin and/or luteolin scattered, [cyanogenesis in ANITA grade?], S [syringyl] lignin units common [positive Maüle reaction - syringyl:guaiacyl ratio more than 2-2.5:1], and hemicelluloses as xyloglucans; root apical meristem intermediate-open; root vascular tissue oligarch [di- to pentarch], lateral roots arise opposite or immediately to the side of [when diarch] xylem poles; origin of epidermis with no clear pattern [probably from inner layer of root cap], trichoblasts [differentiated root hair-forming cells] 0, exodermis +; shoot apex with tunica-corpus construction, tunica 2-layered; reaction wood ?, associated gelatinous fibres [g-fibres] with innermost layer of secondary cell wall rich in cellulose and poor in lignin; starch grains simple; primary cell wall mostly with pectic polysaccharides, poor in mannans; tracheid:tracheid [end wall] plates with scalariform pitting, wood parenchyma +; sieve tubes enucleate, sieve plate with pores (0.1-)0.5-10< µm across, cytoplasm with P-proteins, cytoplasm not occluding pores of sieve plate, companion cell and sieve tube from same mother cell; sugar transport in phloem passive; nodes unilacunar [1:?]; stomata brachyparacytic [ends of subsidiary cells level with ends of pore], outer stomatal ledges producing vestibule, reduction in stomatal conductance to increasing CO2 concentration; lamina formed from the primordial leaf apex, margins toothed, development of venation acropetal, secondary veins pinnate, overall growth ± diffuse, venation hierarchical, fine venation reticulate, veins (1.7-)4.1(-5.7) mm/mm2, endings free; most/all leaves with axillary buds; flowers perfect, pedicellate, ± haplomorphic, parts spiral [esp. the A], free, numbers unstable, development in general centripetal; P not sharply differentiated, with a single trace, outer members not enclosing the rest of the bud, often smaller than inner members; A many, filament not sharply distinguished from anther, stout, broad, with a single trace, anther introrse, tetrasporangiate, sporangia in two groups of two [dithecal], ± embedded in the filament, with at least outer secondary parietal cells dividing, each theca dehiscing longitudinally, endothecium +, endothecial cells elongated at right angles to long axis of anther; tapetum glandular, cells binucleate; microspore mother cells in a block, microsporogenesis successive, walls developing by centripetal furrowing; pollen subspherical, tectum continuous or microperforate, ektexine columellate, endexine thin, compact, lamellate only in the apertural regions; nectary 0; G superior, free, several, ascidiate, with postgenital occlusion by secretion, stylulus short, hollow, cavity not lined by distinct epidermal layer, stigma ± decurrent, carinal, dry [not secretory]; ovules few [?1]/carpel, marginal, anatropous, bitegmic, micropyle endostomal, outer integument 2-3 cells across, often largely subdermal in origin, inner integument 2-3 cells across, often dermal in origin, parietal tissue 1-3 cells across [crassinucellate], nucellar cap?; megasporocyte single, hypodermal, megaspore tetrad linear, functional megaspore chalazal, lacking sporopollenin and cuticle; female gametophyte four-celled [one module, nucleus of egg cell sister to one of the polar nuclei]; ovule not increasing in size between pollination and fertilization; pollen binucleate at dispersal, male gametophyte trinucleate, germinating in less than 3 hours, pollination siphonogamous, tube elongated, growing between cells, growth rate 20-20,000 µm/hour, outer wall pectic, inner wall callose, with callose plugs, penetration of ovules via micropyle [porogamous], whole process takes ca 18 hours, distance to first ovule 1.1-2.1 mm; male gametes lacking cell walls, flagellae 0, double fertilization +, ovules aborting unless fertilized; P deciduous in fruit; seed exotestal, becoming much larger than ovule at time of fertilization; endosperm diploid, cellular [micropylar and chalazal domains develop differently, first division oblique, micropylar end initially with a single large cell, divisions uniseriate, chalazal cell smaller, divisions in several planes], copious, oily and/or proteinaceous; embryogenesis cellular; germination hypogeal, seedlings/young plants sympodial; dark reversal Pfr -> Pr; Arabidopsis-type telomeres [(TTTAGGG)n]; 2C genome size 1-8.2 pg [1 pg = 109 base pairs], whole genome duplication, ndhB gene 21 codons enlarged at the 5' end, single copy of LEAFY and RPB2 gene, knox genes extensively duplicated [A1-A4], AP1/FUL gene, paleo AP3 and PI genes [paralogous B-class genes] +, with "DEAER" motif, SEP3/LOFSEP and three copies of the PHY gene, [PHYB [PHYA + PHYC]].
[NYMPHAEALES [AUSTROBAILEYALES [[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]]]]]: wood fibres +; axial parenchyma diffuse or diffuse-in-aggregates; pollen monosulcate [anasulcate], tectum reticulate-perforate [here?]; ?genome duplication; "DEAER" motif in AP3 and PI genes lost, gaps in these genes.
[AUSTROBAILEYALES [[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]]]]: vessel elements with scalariform perforation plates in primary xylem; essential oils in specialized cells [lamina and P ± pellucid-punctate]; tension wood +; tectum reticulate; anther wall with outer secondary parietal cell layer dividing; carpels plicate; nucellar cap + [character lost where in eudicots?]; 12BP [4 amino acids] deletion in P1 gene.
[[CHLORANTHALES + MAGNOLIIDS] [MONOCOTS [CERATOPHYLLALES + EUDICOTS]]] / MESANGIOSPERMAE: benzylisoquinoline alkaloids +; polyacetate derived anthraquinones + [?level]; outer epidermal walls of root elongation zone with cellulose fibrils oriented transverse to root axis; P more or less whorled, 3-merous [possible positiion]; embryo sac bipolar, 8 nucleate, antipodal cells persisting; endosperm triploid; ?germination.
I largely follow Ronse De Craene et al. (2003) on the insertion of floral organs. To add where?: A whorled, carpel fusion by congenital occlusion.
Age. The divergence of mesangiosperms, i.e. the age of this branching point, is estimated at (190-)167(-47) m.y. (95% HPD, Smith et al. 2010, Table S3), while a fossil-based estimate is ca 100 m.y. (Crepet et al. 2004). [Check]
Phylogeny. Relationships between the lineages immediately above the basal pectinations in the main tree, the ANITA grade - Amborellales, Nymphaeales and Austrobaileyales here - have been for some time unclear. The positions of Ceratophyllales and Chloranthaceae have been particularly uncertain. Indeed, Graham et al. (2005) found that the inclusion of Ceratophyllum and Chloranthaceae in analyses could destabilise relationships among other early branching clades, e.g., the position of the monocots became labile.
There has been weak support for an (eu)magnoliid clade being sister to the eudicots (e.g. D. Soltis et al. 2000; P. Soltis et al. 2000; Wu et al. 2007; Qiu & Estabrook 2008 [compatibility analysis]); in the first study Chloranthales, monocots, and Piperales were successively sister to the remaining taxa, but these relationships also had little support. Hilu et al. (2003: a matK analysis alone), suggested that Ceratophyllaceae were sister to eudicots (see also D. Soltis et al. 2000; Borsch et al. 2005: three rapidly evolving genes, 62% jacknife, 96% posterior probabilities; Müller et al. 2006: 96% posterior probability; some analyses in Saarela et al. 2007; Qiu & Estabrook 2008). However, Zanis et al. (2002) analysed 11 genes from all genomes and found some support for the magnoliids (strong support for this clade) as being sister to eudicots, with Chloranthaceae, and [monocots + Ceratophyllaceae] occurring as successively more basal branches; below these were the ANITA grade (see also Graham & Olmstead 2000; Borsch et al. 2003). Support for some of these nodes depended on the method of analysis (see also Borsch et al. 2000; Graham & Olmstead 2000; Hilu et al. 2001; Whitlock et al. 2001; Zanis et al. 2002) or the particular gene studied (see Duvall & Bricker 2002: nuclear 18s; Hilu et al. 2003: matK; Duvall et al. 2006: nuclear 18S in particular causes problems). Parkinson et al. (1999) found some evidence for a [Piperales + eudicot] clade, but support was weak. Davis et al. (2004) found the magnoliids to be sister to monocots, with Chloranthaceae and Ceratophyllaceae successively sister to a clade including the few eudicots in the analysis, but support was not exactly overwhelming (the best supported topology in Duvall et al. 2006 is somewhat similar). There was fairly good support for a clade [monocots + Ceratophyllaceae] in a compartmentalised 6-gene analysis (Zanis et al. 2003; see also Qiu et al. 1999; some analyses in Saarela et al. 2007). Müller et al. (2006) found very poorly supported relationships between Chloranthaceae and monocots, which together linked with the magnoliids. In work by Whitlock et al. (2002) largely similar groupings were recognised, but support was only moderate; the exact position of Chloranthaceae remained unclear. Some analyses suggested the possibility of a sister taxon relationship between Chloranthaceae and eudicots (Borsch et al. 2003), while Hilu et al. (2003) in a matK analysis, suggested that they were sister to monocots. Although the support was weak in parsimony analyses, there were 100% posterior probabilities in Bayesian analyses, furthermore, in the latter case only, [Chloranthacaeae + monocots] were sister to magnoliids, although the probabilities there were low. Qiu et al. (2005; see also Löhne & Borsch 2005) found initial rather strong bootstrap support for an association between monocots and Ceratophyllaceae in a 9-gene analysis being vitiated by the failure to obtain much support in any of the subanalyses and by details of the topology obtained in the 9-gene analysis itself (e.g. Acorus sister to the Alismatales that were included) that are rather improbable. Other studies also show no clear pattern (e.g. Jansen et al. 2006a; Qiu et al. (2006b). Interestingly, Piper has the "monocot" pattern of PHY genes (A, B, C genes only), while Ceratophyllum has PHYE (Mathews et al. 1995). Jansen et al. (2006b) and Hansen et al. (2007) found support for a sister-group relationship between Chloranthales and the magnoliids, although the analyses in Mathews (2006a) preferred an unresolved position for Chloranthales and Ceratophyllales above the Austrobaileyales. Duvall et al. (2006) and Qiu et al. (2010: chloroplast data, support weak) found a relationship between Chloranthales and Ceratophyllales, as did N. Zhang et al. 92012), that clade being successively sister to the magnoliids and monocots. Soltis et al. (2007a) found weak support for [Ceratophyllum + eudicots]; relationships of Chloranthaceae were unclear and there was very weak support for monocots as sister to [magnoliids, Chloranthaceae, eudicots, etc.]. Although relationships between basal angiosperms were not their focus, Xue et al. (2012) found the relationships [monocots [[Ceratophyllaceae + Piperaceae] + eudicots]] in a ML analysis of whole chloroplast dataD. Soltis et al. (2005b) had very reasonably summarized their discussion on relationhips in this area by showing a pentatomy (see also P. Soltis et al. 1999).
Morphology in particular has quite often suggested that more or less herbaceous clades like Piperales, Chloranthaceae, and Nymphaeales might be related to monocots, the pal(a)eoherb hypothesis (see e.g. Donoghue & Doyle 1989; Taylor & Hickey 1992; Endress 2000). Analysis of combined morphological and molecular data found that Piperales were sister to monocots (Doyle & Endress 2000, support weak). Although quite popular in the latter years of the last century, its support comes from homoplasies associated with the adoption of the herbaceous habit. Indeed, Piperales are now firmly (usually!) embedded in the magnoliid clade, Nymphaeales are very near the base of the whole angiosperm clade, etc.
Some resolution may be on the way. Thus Graham et al. (2005) found a rather weakly supported (73% bootstrap) [monocot + eudicot] grouping, but this was weaker when Chloranthaceae and Ceratophyllaceae were included. However, evolution of some floral developmental genes, e.g. in the C and D lineages, are also consistent with such a relationship (Kramer et al. 2004). The positional relationships between members of the androecium and the perianth, the stamens being individually opposite perianth members, and perhaps some other characters (trimery of some floral whorls is fairly widespread within Ranunculales) are also consistent with such relationships. (There is the same perianth member/stamen relationship in magnoliids like Lauraceae.) Variation in how calcium oxalate is synthesized may also be of phylogenetic interest, although sampling is very poor; neither members of the ANITA grade nor magnoliids have been examined. Vacuolar crystal formation associated with membranes and paracrystalline bodies with widely spaced subunits are found in eudicots (crystals seem also to be formed in other ways here), while in monocots there are no membrane complexes and the paracrystalline bodies have closely spaced subunits (Horner & Wagner 1995; Evert 2006). Some patterns of RNA editing may be phylogenetically informative at this level (Logacheva et al. 2008).
Other studies have found this [monocot + eudicot] grouping; Jansen et al. (2006b: 37 whole chloroplast genomes, see also Zhengqiu et al. 2006), Cai et al. (2006: 35 whole chloroplast genomes, neither Chloranthales, Ceratophyllales, nor any member of Austrobaileyales included), and Duvall et al. (2006: four genes, three compartments, nuclear PHYC gene, 18S being excluded; overall, a relationships between magnoliids and monocots was preferred), Mathews (2006a: three PHY genes, 105 taxa), and Hansen et al. (2007: 61 protein-coding chloroplast genes). When sequences of complete chloroplast genomes are analysed, an association between monocots and eudicots is more strongly suggested. Thus Jansen et al. (2007) found good support for a [monocot + eudicot] grouping, a number of alternative topologies being excluded, but support in Moore et al. (2007) was somewhat less strong. Ceratophyllum, with quite a long branch, was sister to eudicots (in some reconstructions, Piper, with a very long branch, was also involved), and Chloranthus was sister to the magnoliids with moderate to strong support (Jansen et al. 2007), a position also found by Saarela et al. (2007), many analyses in Moore et al. (2007, 2010), and Smith et al. (2010).
However, in Moore et al. (2010) there were a variety of other "minority" relationships - Ceratophyllum or [Ceratophyllum + Piper] as sister to monocots, and Chloranthaceae sister to a clade including magnoliids, monocots, and eudicots; the latter pair of relationships with moderate jacknife support were found by Davis et al. (2013) in their comprehensive chloroplast analysis focussing on monocots. Goremykin et al. (2009b) recently found a [Ceratophyllum + magnoliid] clade (Chloranthaceae were not included) after removing 2,500 highly variable positions from the analysis of chloroplast genome data; with the removal of 1,000 positions Ceratophyllum was sister to a [monocot + eudicot] clade. Analyses of morphological data gave a grouping [Ceratophyllum + Chloranthaceae] with quite strong support (Endress & Doyle 2009; see also Huang et al. 2010, weak support using the ycf2 gene; Moore et al. 2011, very weak support). Finally, in the massive parsimony analysis of Goloboff et al. (2009) a set of relationships [Ceratophyllum [[Amborella + Nymphaeales]....[[Chloranthaceae + monocots] eudicots]]] was retrieved, Duarte et al. (2010) found the relationships [monocots [magnoliids + eudicots]], and Tamura et al. (2011: 6 plastid, 7 mitochondrial, and 1 nuclear genes sequenced) found some support for a [Piperales + monocot] clade, as did Barrett and Davis (2011: or [Piperales + Ceratophyllales] the sister group). For another study, see Fiz-Palacios et al. (2011).
Lee et al. (2011) found a clade with monocots sister to all angiosperms minus Nymphaeales and Amborellales, and also a [magnoliid + eudicot] clade, both with strong support. Although this may be a sampling problem (no Austrobaileyales, Chloranthales or Ceratophyllales were included), Lee et al. (2011) initially looked at almost 23,000 sets of orthologues from nuclear genomes of 101 genera of land plants (most taxa represented by ESTs, genes included if represented in as few as 4 taxa).
Nevertheless, pending further studies, the relationships [[Chloranthales + magnoliids] [monocots [Ceratophyllales + eudicots]]] are used as a basis for further discussion, rather different from the relationships suggested in the first six editions of this site and also from the tree in A.P.G. II (2003) and Soltis et al. (2005b). Direct links are provided to the pages where the clades just mentioned are discussed further; this discussion will generally be just above or below where the link takes you: Ceratophyllales, Chloranthales, eudicots, and monocots.
[CHLORANTHALES [[MAGNOLIALES + LAURALES] [CANELLALES + PIPERALES]]]: sesquiterpenes +; seed endotestal.
[[MAGNOLIALES + LAURALES] [CANELLALES + PIPERALES]] / MAGNOLIIDS / MAGNOLIANAE Takhtajan: (neolignans +); vessels solitary and in radial multiples, (with simple perforation plates in primary xylem); (sieve tube plastids with polygonal protein crystals); lamina margins entire; A many, spiral [possible position here], extrorse; ovules with hypostase, nucellar cap +, raphal bundle branches at the chalaza; antipodal cells soon die. Back to Main Tree
4 orders, 20 families, 9900 species.
Age. Crown group magnoliids diverged 134.5-125.7 m.y.a. (Moore et al. 2007), while Bell et al. (2010: note topology) suggested ages of (138-)122(-108) or (130-)125(-121) m.y. depending on the method used. Magallón and Castillo (2009) estimated crown group divergence at ca 201.7-198.2 and 128-127.7 m.y.a.. Other estimates range from (181-)155(-136) m.y. old (with eudicot calibration) to (198-)163(-138) m.y. (without: Smith et al. 2010), to (144-)116(-89) m.y. (N. Zhang et al. 2012) or (155-)149, 137(-131) m.y. (Wikstöm et al. 2001), while Magallón et al. (2013: with temporal constraints) suggested an age of around (167.2-)147.7-142.6(-128.5) m.y. for this clade.
An earlier fossil-based estimate for both stem and crown divergence is ca 98 m.y. (Crepet et al. 2004: magnoliids sister to monocots). For summaries of the fossil history of the group, especially prominent in the Mid Cretaceous and later, see Friis et al. (1997, 2006, 2011).
Evolution. Plant-Animal Interactions. Caterpillars of Papilionidae-Papilioninae butterflies are notably common (almost 33% of the records) on members of this group; they are, however, not known on Myristicaceae, in Laurales they predominate on Lauraceae, and in Piperales on Aristolochiaceae (see Scriber et al. 1995 for references; Zakharov et al. 2004). They are also commonly found on Rutaceae, which have similar alkaloids.
Chemistry, Morphology, etc. Hegnauer (1990) discussed the chemistry of the Polycarpicae, which includes Austrobaileyales and Ranunculales; similar isoflavonoids are found in Magnoliaceae, Lauraceae and Chloranthaceae.
For a convenient summary of a number of features of wood anatomy, see Herendeen et al. (1999b). Magnolioid roots are scattered through the clade, being recorded from a number of woody members of Laurales and Magnoliales. These roots are stout, often ³3 mm across, lack root hairs (?always), and are endomycorrhizal - associated conditions (see Baylis 1975; St John 1980). Taxa with broadly lobed leaf blades are scattered throughout this clade (but not in the small Canellales). See Erbar (1983, inc. Illicium) for carpel morphology and Ronse Decraene and Smets (1992b) for androecial morphology.
Phylogeny. Although the sister group relationship of Piperales with Canellales in particular is at first sight unexpected, the magnoliid clade as a whole and the relationships within the group are turning out to be quite robust. There was not much morphological support for this grouping (Doyle & Endress 2000); features like tectum structure, etc., showing considerable variation (J. A. Doyle 2005). However, molecular support for the clade has been increasing in successive studies (e.g. Qiu et al. 1999, 2000, 2005: support levels depend on the analysis, the node sometimes collapses], 2006b, 2010; Zanis et al. 2002; Jansen et al. 2006b; Zhengqiu et al. 2006; Cai et al. 2006; Müller et al. 2006: support for [Canellales + Piperales] poor; Jansen et al. 2007: little maximum parsimony support; Moore et al. 2007: group not in all analyses; Soltis et al. 2007a; Soltis et al. 2011). Support includes the possession of unique indels (Löhne & Borsch 2005). Soltis et al. (2007a) found weak support for a grouping [Magnoliales + Canellales]; relationships in Bell et al. (2010) are [Piperales [Laurales [Canellales + Magnoliales]]], although they have little support.
[MAGNOLIALES + LAURALES]: cuticle waxes as annularly-ridged rodlets, palmitol the main wax; A whorled; pollen 1-2 nexine foliations, outer member massive, lamellate endexine; carpel cross-zone initiated late; ovules 1(-2)/carpel, basal, erect, apotropous; fruitlets 1-seeded.
Evolution. Divergence & Distribution. Magallón and Castillo (2009) suggest that the two clades diverged ca 198.2 and 127.7 m.y.a. - relaxed and constrained penalized likelihood datings, while Magallón et al. (2013) suggested an age of around 137.2 m.y..
There is considerable variation as to how the androecium is initiated; it would not take much for "flowers with inner staminodes" to be an apomorphy at this level. Seed size is notably large in this clade (Moles et al. 2005a; Sims 2012).
MAGNOLIALES Bromhead Main Tree.
Vessels in multiples; secondary phloem stratified; pith septate [with sclerenchymatous diaphragms]; nodes 3:3; petiole vasculature an arc with an adaxial plate; branching from the current innovation; leaves two-ranked, lamina vernation conduplicate; "bract" sheathing; P whorled; G occluded by fusion and secretion; ovule with outer integument 5-10 cells across, obturator +; seeds medium-sized, testa vascularized, multiplicative; endosperm ?type, irregularly ruminate. - 5 families, 154 genera, 2929 species.
Age. Magallón and Castillo (2009) suggest possible ages for the crown group of around 171.5 and 116.6 m.y., Bell et al. (2010) ages of (96-)76, 69(-50) m.y.; other ages are (119-)113, 108(-102) m.y. (Wikstöm et al. 2001).
Note: Possible apomorphies are in bold. However, the actual level at which many of these features, particularly the more cryptic ones, should be assigned is unclear. This is partly because many characters show considerable homoplasy, in addition, basic information for all too many characters is very incomplete, frequently coming from taxa well embedded in the clade of interest and so making the position of any putative apomorphy uncertain. Then there is the not-so-trivial issue of how ancestral states are reconstructed...
Evolution. Divergence & Distribution. There is extensive discussion on character evolution in Sauquet et al. (2003), and much of the character hierarchy here is based on this paper. However, where characters like extrorse/introrse anther dehiscence and ruminate/non-ruminate testa are placed on the tree depends on how the characters are optimised, or even defined (e.g. ruminate endosperm). There is also some conflict with the positions of characters as they are optimised on a more extensive tree for basal angiosperms, although less detailed for Magnoliales (c.f. Ronse De Craene et al. 2003, also Judd et al. 2003). Doyle and Endress (2000) and Soltis et al. (2005) suggest additional characters for the clade, including reduced fibre pit borders, palisade parenchyma, foliar astrosclereids, and pollen with continuous tectum; Doyle and le Thomas (2012) outline pollen evolution. More characters may well need to be added to the apomorphy scheme for the order.
Chemistry, Morphology, etc. Intra- and interfamilial variation of morphological characters that are frequently used to reconstruct phylogenies is considerable, in particular, Furness et al. (2002) emphasize the variability of microsporogenesis in the order.
The "bracts" with sheathing bases that have been reported for a number of Magnoliales (Endress & Armstrong 2011: condition in Degeneriaceae?), are perhaps a little unexpected since the leaf bases in the clade tend to be quite narrow - except for the stipulate Magnoliaceae. Deroin (2010) noted a tendency for the vasculature of the perianth and/or androecium in some Annonaceae and Magnoliaceae to be pentamerous.
For additional information, see Sugiyama (1976a, b, 1979: esp. considerable variation in cotyledonary nodal morphology), Hiepko (1964b: perianth vasculature), Endress (1977b, 1986a, 1994a: floral morphology), van Heel (1981, 1983: carpel development), Erbar and Leins (1983: floral development), Metcalfe (1987: general anatomy), Taylor and Hickey (1995: general), Ronse Decraene and Smets (1996a: androecium), Kimoto and Tobe (2001: embryology), and Kim et al. (2003, 2004: AP3 and P1 genes: Myristicaceae not sampled).
Phylogeny. Molecular data suggested that Myristicaceae are sister to the rest of the order, but support was only moderate (D. Soltis et al. 2000); the addition of morphological data strengthened that position, and also placed Magnoliaceae as sister to the remaining taxa (Doyle & Endress 2000), although the latter position had only moderate support (c.f. P. Soltis et al. 2000; see also Sauquet et al. 2001). The family pairs [Annonaceae + Eupomatiaceae] and [Degeneriaceae + Himantandraceae] are both well supported (D. Soltis et al. 2000; P. Soltis et al. 2000; Doyle & Endress 2000). Other studies confirm these relationships (Sauquet et al. 2003; Müller et al. 2006), and they are followed here.
Hilu et al. (2003: matK analysis alone, ?sampling) found a somewhat different but poorly supported set of relationships, and Qiu et al. (2010: mitochondrial genes) also recovered a somewhat different topology, Degeneriaceae being sister to the rest of the order minus Myristicaceae, although again support was weak. Soltis et al. (2011) found that Magnoliaceae were sister to the rest of the order, the clade [Degeneriaceae + Myristicaceae] had good support although otherwise support was weak; this unexpected topology was ascribed to signal from rDNA sequences. Finally, Morton (2011: nuclear gene Xdh) found a set of relationships [Himantandraceae [Myristicaceae [Degeneriaceae [the rest]]]].
Includes Annonaceae, Degeneriaceae, Eupomatiaceae, Himantandraceae, Magnoliaceae, Myristicaceae.
Synonymy: Annonales Berchtold & Presl, Degeneriales C. Y Wu et al., Eupomatiales Reveal, Himantandrales Doweld & Shevyryova, Myristicales Berchtold & Presl - Magnoliineae, Myristicineae Chatrou - Annonanae Doweld, Magnolianae Takhtajan - Magnoliidae Takhtajan - Magnolipsida Brongniart - Magnoliid I group (Nandi et al. 1998).
MYRISTICACEAE R. Brown Back to Magnoliales
Tree, branches plagiotropic, pseudowhorled; exudate +, red; isoflavonoids, flavonoids diverse [flavones +], polyketide [acetogenins], (tryptamine) alkaloids +, isoquinoline alkaloids 0; cork also in outer cortex; primary stem ± with continuous cylinder; vessel elements with simple or scalariform(-reticulate) perforations; tannin-containing tubes in the xylem; sieve tubes with non-dispersive protein bodies, (plastids with protein crystalloids and starch); petiole bundles bicollateral; (branched) sclereids or fibres +; (acicular) crystals +; hairs branched, arbuscular to stellate (T-shaped); cuticle waxes as platelets; (leaves spiral), (lamina lobed - some Knema); plants dioecious; flowers small (<1 cm across), polysymmetric, receptacle small; P (2-)3(-5), uniseriate, connate; staminate flowers: A whorled (spiral), 2-40, connate (not), (anthers unithecate); (pollen inaperturate, ulceroid, spiraperturate, ektexine granular), pollen aperture membrane sculpted; pistillode 0; carpellate flowers: staminodes 0; G 1; (stylulus ± long), stigma 2-lobed to peltate; ovule 1/carpel, subbasal, inner integument (3-)7-10 cells across; fruit a follicle, dehiscing abaxially as well, (indehiscent); seed large [³1 cm across], pachychalazal, aril +, micropylar-funicular, vascularized, (aril small); exotesta thick-walled, endotesta palisade, lignified, crystalliferous, tegmen multiplicative or not, exotegmen with fibres or sclerotic or tracheidal cells, chalaza massive, with a lignified counter-palisade; endosperm nuclear, with (starch and) oil; embryo with hypocotyl not developed; n = 20, 22, 25, 26; germination hypogeal.
20[list]/475: Myristica (175), Horsfieldia (100), Knema (95), Virola (60). Pantropical, few mainland Africa (map: from de Wilde 2000 [Indo-Malesia]; Heywood 2007; Trop. Afr. Fl. Pl. Ecol. Distr. 1. 2003).[Photo - Carpellate flower, Fruit].
Age. Crown-group Myristicaceae are estimated to be a mere 21-15 m.y. old (J. A. Doyle et al. 2004).
The discovery of fossil seeds apparently of Myristicaceae from the Eocene (London Clay) does not bear on the crown-group age because they cannot be placed precisely on the tree of the family (J. A. Doyle et al. 2008a).
Floral formula: * P [3]; A [2-40]: * P [3]; G 1.
Evolution. Divergence & Distribution. Crown-group diversification, some 21-15 m.y.a., is very recent indeed given possible stem ages of the family that are over 100 m.y. more (see above), its distribution throughout the humid tropics, and its apparently low dispersability (J. A. Doyle et al. 2004). Sauquet et al. (2003) found that there was a long branch leading to crown Myristicaceae and little molecular divergence between its extant members, which might also suggest recent diversification. In this respect Myristicaceae can be compared with Annonaceae, also pantropical but a little younger; there diversification may have begun mid-Cretaceous or slightly later (Doyle et al. 2004; Scharaschkin & Doyle 2005; esp. Couvreur et al. 2011a; Doyle et al. 2012).
See Chatrou (2003) for possible apomorphies.
Ecology & Physiology. Myristicaceae are disproportionally well represented (11 of the 57 species of the family recorded) of the 227 common trees (d.b.h. at least 10 cm) of the Amazonian forests (ter Steege et al. 2103).
Genes & Genomes. Isozyme duplication (in Myristica) suggests ancient polyploidy (Soltis & Soltis 1990).
Chemistry, Morphology, etc. There are free phloem strands in the center of the midrib bundle. The wood rays are not notably broad.
Inflorescence morphology is a little confusing. Ronse de Craene (2010; see also Armstrong & Tucker 1986) shows a large, abaxial "bracteole" in addition to paired lateral prophyllar/bracteolar-like structures that may (staminate inflorescence) or may not subtend buds. The "bract" immediately below the staminate flower of Myristica fragrans has a very broad base.
There are three traces per perianth lobe (Siddiqui & Wilson 1976). The synandrium of Myristica has a sterile apex, probably axial (Armstrong & Tucker 1986), and judging from the vasculature, the unithecal anthers found in some species of Myristica represent half of a divided bithecal anther (Wilson & Maculans 1967; Armstrong & Tucker 1986); anthers in Knema are tetrasporangiate (Xu & Ronse de Craene 2010a). For the complex vascularisation of the carpel in Myristica, see Wilson and Maculans (1967), and for the vascularization of the ovule in Myristica fragrans, see Dickison (2000). A small structure developing between the two integuments is reported by Nair (1972). Corner (1976) commented on the complexity of the seed coat in Myristicaceae which is as elaborate as that of any other angiosperm. Not all taxa appear to have a multiplicative testa, and some have a multiplicative tegmen; the ruminations may develop best in the pachychalazal part of the seed.
For more information, see Hegnauer (1969, 1990: chemistry), Mauritzon (1939a) and Endress (1973), both seed, Kerster and Baas (1981: anatomy), Kühn and Kubitzki (1993: general), de Wilde (2000: Malesian Myristicaceae), Jiménez-Rojas et al. (2002: growth patterns - Massart's model), Sauquet (2003: androecium), Sauquet and Le Thomas (2003: pollen) and Oginuma et al. (2012: chromosome numbers, a few South East Asian-Malesian taxa only known).
Phylogeny. Relationships within the family are unclear. The African and Madagascan taxa may form a clade, possibly sister to Compsoneura (but perhaps long branch attraction), overall, geography and relationships may be summarized as [[Asia + America] [Madagascar and Africa]]. Within Asian/Malesian Myristicaceae, Knema and Myristica may be sister taxa. Sauquet et al. (2001, 2003), Sauquet (2003) and Sauquet and Le Thomas (2003) suggest that the free stamens (in some species they are numerous and apparently spirally inserted) and small aril of Mauloutchia, apparently plesiomorphic features, are in fact more likely to be derived.
[Magnoliaceae [[Degeneriaceae + Himantandraceae] [Eupomatiaceae + Annonaceae]]]: primary stem with distinct bundles [eustele]; wood with broad rays; flowers solitary, large [.1.5 cm across]; receptacle with cortical vascular system; P = K + C; A many, spiral [possible position here], filaments with three veins, anther thecae separate, embedded in the broad connective, connective prolonged [one position]; tectum imperforate; G spiral; ovule with funicular obturator; ³1 seeds/fruitlet; 10-aa deletion in PI-derived motif in AP3 gene.
Age. This node can be dated to 120-100 m.y.a. (Doyle et al. 2004), also (108-)101, 97(-90) m.y.a. (Wikström et al. 2001), (106-)102(-98) m.y.a. (Su & Saunders 2009), or as recently as ca 71.1 m.y.a. (Magallón et al. 2013).
Chemistry, Morphology, etc. There is more than one kind of cortical vascular system here, and what parts of the flower are supplied by which system varies (Ronse De Craene et al. 2003, see also Deroin 1991, 1999a).
MAGNOLIACEAE Jussieu Back to Magnoliales
(Deciduous); (silicon concentration high), sesquiterpene lactones +; vessel elements with simple and scalariform perforation plates, (walls vestured - Liriodendron); wood fluorescing; (secondary phloem ± stratified, rays broad); (sieve tube plastids with polygonal protein crystalloids and starch - Magnolia); nodes ³6:³6; pith septate; resin cells +; petiole also with medullary bundles (strictly annular - Liriodendron); leaves also spiral, lamina vernation laterally or vertically conduplicate (supervolute-curved), stipules +, sheathing stem, open opposite petiole; (dioecious); flowers terminal or axillary, ± polysymmetric; receptacle much elongated; P 2, 3 + 3, or 3 + many, K and C ± distinguishable, ± whorled; (A introrse - Magnolia), (endothecium biseriate), (filament vein single), tapetal cells multinucleate; pollen grains boat-shaped; G 1-many, fusion complete, but secretory canal +, stigma (terminal), dry, elongate (not); ovules(1-)2-12(-16)/carpel, micropyle exo/bistomal/zig-zag, outer integument 4-10 cells across, inner integument 2-3(-4) cells across, parietal tissue 2-8 cells across; fruit dehiscing abaxially (circumscissile), (indehiscent, winged); sarcotesta +, endotesta lignified [fibres], (tegmen ± tanniniferous), (testa inconspicuous - Liriodendron); endosperm not ruminate, (nuclear - "Magnolia" septentrionalis); n = 19.
2[list]/227: Magnolia (ca 225). The Americas (but not W. North America), and South East Asia to Malesia (map: from Good 1974; Lozana-Contreras 1994).[Photo - Collection.]
Age. Molecular estimates of the crown group age are (54-)36, 33(-17) m.y. (Bell et al. 2010: note topology) or (86-)79, 70(-63) m.y. (Wikström et al. 2001).
Archaeanthus, a Cenomanian fossil of some 98 m.y. in age from North America, may be assignable to Magnoliaceae (Dilcher & Crane 1984); it is placed sister to the family in the constrained morphological analysis of Doyle and Endress (2010). It has follicular fruits that fall off the axis each follicle having 10-18 winged seeds. Romanov and Dilcher (2013: ?outgroup) placed Archaeanthus, along with Liriodendroidea, known from its winged seeds, as sister to Liriodendron, which would call into question the accuracy of the molecular estimates above.
Lesqueria, from about the same period, may also belong around here; it has similar fruits (Crane & Dilcher 1984). The fruits of Protomonimia, in Turonian deposits from Japan ca 91 m.y. old, have several carpels borne in spirals on a concave axis; there is a stigmatic crest. The young seeds have a thick testa, the exotesta being palisade and with sinuous anticlinal cell walls (Nishida & Nishida 1988); they may be of a taxon somewhere near Magnoliaceae, although i.a. the shape of its receptacle is very different.
Evolution. Divergence & Distribution. Liriodendron was widely distributed in the Northern Hemisphere in the early Tertiary (Ferguson et al. 1997). For divergence ages of and within Magnolia, see Azuma et al. 92001) and Nie et al. (2008).
Pollination Biology. Beetles are common pollinators, and the flowers of species like Magnolia ovata, pollinated by dynastid scarab beetles, are thermogenic (Seymour 2001; Gottsberger et al. 2012). Floral scents of the family have been extensively studied (Azuma et al. 1999; Gottsberger et al. 2012).
Genes & Genomes. Both isozyme duplication and stomatal size increase over time suggest ancient polyploidy (Soltis & Soltis 1990; Masterson 1994).
Chemistry, Morphology, etc. Sclerenchymatous diaphragms in the pith of the stem are particularly conspicuous in the Michelia group of Magnolia. Terminal tracheids are commonly silicified, forming distinctive phytoliths (Piperno 2006).
The endexine is lamellate and the columellae fused-granular (Xu & Kirchoff 2008). Nectar is secreted from the exposed surfaces of the carpels in some species of Magnolia. The micropyle is exostomal in the Michelia group of Magnolia. The seed of Magnolia may dangle from the open fruitlet attached by extended annular thickenings of the protoxylem vessels.
Liriodendron: leaves lobed, with palmate secondary veins; petiole bundle strictly annular; fruit a samara, single-seeded; seeds lack a sarcotesta, mesotesta more or less sclerotic; endosperm slight and not ruminate. Magnolia: leaves entire; anthers introrse; ovule lacks a funicular obturator(?); fruit dehiscent. but not adaxial (so the fruit is not a follicle in the strict sense); sarcotesta +, scleroendotesta with crystals and lignified fibrils in the cells; endosperm copious. There is also a simple or tubular pore in the seed coat that marks the passage of the vascular bundle through the sclerotesta (Xu 2003).
Some information is also taken from Hegnauer (1969, 1990: chemistry), Bouman (1977: ovules and seeds of Liriodendron), Nooteboom (1993: general), Yamada et al. (2003b: ovules), Pan et al. (2003) and especially Fu et al. (2012 and references), both embryology, Xu (2006) and Xu and Rudall (2006: floral development), and Xu and Kirchoff (2008: pollen morphology); see Azuma et al. (2000), Ueda et al. (2000) and Wang et al. (2006) for molecular phlogenies, Figlar (2000) for branching patterns, Charlton (1994) and Liao and Xia (2007) for phyllotaxis and vernation.
Phylogeny. Magnolia s. str. is pretty wildly paraphyletic, and section Talauma is sister to the rest of Magnolia (Qiu et al. 1995; Kim et al. 2001a, b). Spport along the backbone of then genus (s.l.) is poor, although there are 11 or more well-supported clades along it (Azuma et al. 2001, 2011; Nie et al. 2008; Kim & Suh 2013).
Classification. It seems best to expand Magnolia to encompass the whole of the current Magnolieae; the family thus includes only two genera and tribes are superfluous; see also Nooteboom (2000) and Kim and Suh (2013) for suggestions about classification. However, Romanov and Dilcher (2013) need an order for the family, while Xia (2012) and Sima and Lu (2012) provide a reclassification of Magnolia in which it is divided into 16 genera placed in two tribes... For a checklist and bibliography (under some old generic names), see Frodin and Govaerts (1996).
Synonymy: Liriodendraceae F. Barkley
[[Degeneriaceae + Himantandraceae] [Eupomatiaceae + Annonaceae]]: flower haplomorphic; anthers valvate, inner staminodes +, tapetum secretory; pollen smooth, exine thin, [0.2-1.3 µm across], nexine foliations?, infratectum granular; style with differentiated transmission tissue; seed ruminations irregular, mesotesta fibrous.
Age. Endressinia brasiliana, from the Brazilian Crato formation of some 113 m.y.a. has i.a. distinctive glandular staminodia. It is placed sister to this clade by Doyle and Endress (2010), largely confirming the position suggested by Mohr and Bernardes-de-Oliviera (2004).
Chemistry, Morphology, etc. Doyle (2007) noted that the venation here is often poorly differentiated and so of low rank (but c.f. many Annonaceae, albeit not "basal" members) - I have not placed this character on the tree. For the inner staminodes that are often so conspicuous in the flowers, and their function in pollination - food for pollinators, attractants - see Endress (1984).
[Degeneriaceae + Himantandraceae]: flowers axillary; nexine foliations absent; outer integument with annular opening; x = 12.
Age. The age of this node is around (59-)42, 38(-22) m.y. (Bell et al. 2010) or (80-)73, 63(-56) m.y. (Wikström et al. 2001), c.f. topology.
DEGENERIACEAE I. W. Bailey & A. C. Smith Back to Magnoliales
Alkaloids 0; cork ?; vessel elements with scalariform perforation plates; sieve tube plastids with polygonal protein crystalloids and starch; nodes 5:5; petiole also with medullary bundles; secretory cells 0; cuticle waxes as platelets; leaves spiral; bracts?; K 3, C many, whorled; pollen boat-shaped; G 1, basally ascidiate, occluded by fusion only, stigmatic crest much elongated; ovules many/carpel, outer integument 7-8 cells across, inner integument ca 3 cells across, funicle long; fruit a follicle; exotestal cells palisade, thin-walled, sarcomesotesta +, endotesta with lignified internal fibrils; suspensor massive, embryo with 3 (4) cotyledons.
1[list]/2. Fiji, Viti Levu (map: original!).
Chemistry, Morphology, etc. Some information is taken from Hegnauer (1973, 1990, as Winteraceae: chemistry) and Kubitzki (1993b: general).
HIMANTANDRACEAE Diels Back to Magnoliales
Polyketide alkaloids only; vessel elements with simple (and scalariform) perforations; sieve elements with non-dispersive protein bodies; trichomes peltate; cuticle wax crystalloids 0; branching?; P or bract + bracteoles enclosing the flower in 2 series, caducous, or 2 connate + 4 connate; "C" 3-23 [staminodial]; pollen scabrate; additional staminodes with glands; G 6-30, (1) 2 pendulous ovules/carpel; fruit ± syncarpous, drupaceous, with several stones; seeds laterally flattened, ?ruminations; testa not multiplicative, endotesta aerenchymatous; endosperm development?; seedling?
1 (Galbulimima)[list]/2. The Celebes, New Guinea and N.E. Australia (map: from Hoogland 1972; Endress 1983). [Photo - Flower, Fruit, Buds, Habit]
Chemistry, Morphology, etc. For the testa, see Doweld and Shevyryova (1997); the ruminations are at best obscure. The perianth may be staminodial in origin; according to Johri et al. (1992) the outer integument is not vascularized. Some other information is taken from Hegnauer (1966, 1989: chemistry) and Endress (1993: general).
[Eupomatiaceae + Annonaceae]: sieve tube plastids also with polygonal protein crystalloids; rays 8-15-seriate; petiole bundles arcuate; trunk leaves spiral; prophyll single, adaxial; lamina vernation conduplicate; inflorescence +; nexine foliations undifferentiated; fruit ± berry-like.
Age. Estimates of the time of divergence of these two families vary considerably: Around (69-)55, 50(-35) m.y. (Bell et al. 2010), (97-)91, 82(-76) m.y. (Wikström et al. 2001), and (110.4-)106.3(-102.0) m.y. (Couvreur et al. 2011a: HPD estimates). Other suggested ages are (110.4-)101.7(-99.4) m.y. (Surveswaran et al. 2010: HPD estimates), (101.5-)98.0(-94.9) m.y. (Su & Saunders 2009), or somewhere between 84.7 and 62.6 m.y. (Erkens et al. 2009: a variety of estimates); see also Pirie and Doyle (2012).
Evolution. Divergence & Distribution. Optimisation of prophyll position on the tree is uncertain. Adaxial prophylls are common in Annonaceae, including Anaxagorea, although some taxa have paired, lateral prophylls (Fries 1911).
EUPOMATIACEAE Orban, nom. cons. Back to Magnoliales
Also rhizomatous, with xylopodium or root tubers; pith not septate; vessel elements with scalariform perforation plates; secondary phloem stratification?; sieve tubes with non-dispersive protein bodies [check], plastids also with protein rods; nodes (5-)7(-11):(5-)7(-11); secretory cells +; (stomata anomocytic); inflorescence fasciculate; receptacle concave, calyptra +, thick, deciduous, with sclereids, phyllotaxis spiral; P 0; A introrse, staminodes 15+, petal-like, connate basally, with many glands; pollen with encircling equatorial sulcus; G many, ± connate, ascidiate, occluded by fusion only, placentation sublaminar, stigma flat, papillate; ovules 2-11/carpel; fruit a "berry"; testa ?not vascularized, exotestal cells with thickened unlignified walls, endotesta unlignified, exotegmic cells cuboid, slightly lignified, endotegmic cells enlarged, crushed; n = 10; germination epigeal/phanerocotylar.
1[list]/3. New Guinea and E. Australia (map: from Hoogland 1972; Endress 1983). [Photos - Flower, Flower]
Chemistry, Morphology, etc. The main axes are mixed, initially being orthotropic and bearing spirally-arranged leaves, later becoming more or less plagiotropic and with 2-ranked leaves. There are about three two-ranked bracts on the pedicel; the calyptra itself is often interpreted as being a modified, amplexicaul bract (e.g. Endress 2003b; esp. Kim et al. 2005b). There is no evidence of isozyme duplication (Soltis & Soltis 1990).
Some information is taken from Hegnauer (1966, 1989: chemistry), Endress (1983, 1993: general) and Rix and Endress (2007: an easy account).
ANNONACEAE Jussieu Back to Magnoliales
Vessel elements with simple perforation plates; parenchyma in bands joining rays; lateral traces leaving stele well before central trace; sieve tube plastids also with protein filaments; epidermal cells with single crystals [?level]; (inflorescence leaf-opposed); pedicels articulated [level?]; flowers 3-merous, polysymmetric; P very thick, 3, ± calycine, valvate, + 3 + 3, ± corolline, open to valvate; A whorled, filaments with a single vein; G plicate; funicle short; monocarp abscission at the base; endotestal plug +, mesotesta with several layers of longitudinal and internally of transverse fibres, aerenchyma internal; vascular bundle in antiraphe.
129/2220 [list] - four groups below. Largely tropical.
Age. Crown group diversification may have occurred (90.4-)89.5(-89) m.y.a. (Su & Saunders 2009: calibration on Futabanthus), 82-57 m.y.a. (J. A. Doyle et al. 2004), or 84 m.y.a. (Scharaschkin & Doyle 2005; check); see Couvreur et al. (2011a) for additonal estimates.
Futabanthus, based on fossil flowers from the late Cretaceous (Early Coniacian, ca 89 m.y.a.) of Japan, can perhaps be assigned to crown-group Annonaceae (Takahashi et al. 2008b).
1. Anaxagoreoideae Chatrou, Pirie, Erkens & Couvreur
Ray cells with druses; uniseriate hairs terminate in a rounded cell, other hairs ± stellate; trunk leaves 2-ranked; (petiole vascular tissue ± annular; with adaxial bicollateral plate; small arcuate bundles variously arranged); stomata allelocytic; anther connective tongue-like, inner staminode secretions 0; pollen with granular infratectum; orbicules +; G 3, receptacular vascular system 0; ovules 2/carpel; fruit an explosively dehiscent follicle; seed asymmetric; endotesta aerenchymatous, tegmen alone involved in ruminations, with oil globules; n = 8.
1/21. Tropical America, Sri Lanka to West Malesia, also Halamahera and Ceram (map: from Maas & Westra 1984, 1985). [Photo - Flower, Fruit, Fruit.]
Age. Divergence within Anaxagorea may have begun ca 44 m.y.a. (Scharaschkin & Doyle 2005).
[Ambavioideae [Malmeoideae + Annonoideae]]: ray cells lacking druses; (outer staminodes +), inner staminodes 0; ovules with outer integument 4-6 cells across, inner integument 2-4 cells across, nucellar cap to 5 cells across; fruit berrylets, (monocarp abscission at the apex); seed symmetric, with perichalazal ring; amyloid [xyloglucans] in endosperm +. (map: from Aubréville 1974a; Trop. Afr. Fl. Pl. Ecol. Distr. 1. 2003.)
Age. This node has been dated to (84.4-)74.7(-63.6) m.y.a. (Su & Saunders 2009), (76.6-)68, 64.5(-53.0) m.y. (Erkens et al. 2009), and 70-65 m.y.a. (Couvreur et al. 2011a).
2. Ambavioideae Chatrou, Pirie, Erkens & Couvreur
Anther connective ± tongue-like to peltate-truncate; orbicules ?; carpels 3-8, stipe articulated, (extragynoecial compitum + - Cananga), ovules (1)2-16/carpel, middle integument + (0), 4-6 cells across; (seed arillate), (endosperm ruminations lamelliform); n = 7, 8.
9/50: Drepananthus (26). Tropical, inc. Madagascar.
Age. Crown-group Ambavioideae have been dated to (80.9-)57.0(-52.3) m.y.a. (Surveswaran et al. 2010) and (78-)69.4(-60.2) m.y. (Couvreur et al. 2011).
[Malmeoideae + Annonoideae]: (polyacetylenes +); neolignans 0?; petioles short; anther connective peltate-truncate; (pollen inaperturate), tectum retiperforate, infratectum columellate, outer foliation of nexine massive; ovules 1-many/carpel,
Age. This node has been dated to (69-)66.7-56.6(-54.3) m.y. (Richardson et al. 2004) and (78.1-)67.3(-55.2) m.y. (Su & Saunders 2009).
3. Malmeoideae Chatrou, Pirie, Erkens & Couvreur / The Short Branch Clade
(Pollen globose, ± spiny, disulcate); orbicules + (0); (tegmen with oil globules); endosperm ruminations spiciform (lamelliform); (mesotesta with transverse fibres only - Polyalthia).
Polyalthia (100), Monoon (56), Pseuduvaria (50), Unoniopsis (45). Lowland tropics.
Age. The age of crown-group Malmeoideae has been dated to (66.1-)62.5-53.1(-49.5) m.y. (Richardson et al. 2004) and (55.1-)39.8(-26.7) m.y. (Su & Saunders 2009).
4. Annonoideae Rafinesque / The Long Branch Clade
(Lianes), (deciduous); (C 3); (anthers locellate), (outer staminodes +); pollen (in tetrads or polyads), (inaperturate), (exine foliations numerous), (orbicules +); (G connate); (middle integument + - Artabotrys); (fruit follicle/dehiscing abaxially); endosperm ruminations lamelliform (irregular); n = 7-9.
Guatteria (280), Annona (120-175, soursop, sweetsop: inc. Rollinia), Xylopia (100-160), Uvaria (110-150), Artabotrys (100), Goniothalamus (50-120), Duguetia (70-95), Fissistigma (60), Monanthotaxis (55), Friesodielsia (50-60). Predominantly lowland tropics, rarely temperate.
Age. Crown-group Annonoideae may be (62.5-)60.2-51.1(48.7) (Richardson et al. 2004) or (70.5-)59.6(-48.1) m.y. old (Su & Saunders 2009).
To be assigned to the above two subfamilies.
acetogenins [antimicrobial] +; terminal cell of uniseriate hairs pointed (hairs stellate); (primary stem with continuous cylinder); cambium storied; (sieve tube plastids also with protein fibres); petiole bundle also annular; branched sclereids or fibres common; secretory cells +; (prophyll single, adaxial); inflorescence cymose (leaf opposed), or flowers axillary; flowers A (³3), (latrorse, introrse), tapetal cells amoeboid, multinucleate; (microsporogenesis successive); (pollen with circular sulcus), (trinucleate); G (3-)many (syncarpous; parietal placentation), when 3 opposite outer P, stigma capitate to U-shaped, wet, extragynoecial compitum common; (nucellar cap 0); (sarcotesta +), endotesta crystalliferous, (with thin walled, longitudinal fibres); germination epigeal/crypto- or phanerocotylar.
Synonymy: Hornschuchiaceae J. Agardh, Monodoraceae J. Agardh [where?]
Floral formula - Anaxagoreoideae: * K 3; C 3 + 3; A many, + severalo; G 3.
Everything Else: * K 3; C 3 + 3; A many; G (1-)many.
Evolution. Divergence & Distribution. The biogeography and diversification of the family has been much studied (see also Doyle et al. 2004; Richardson et al. 2004; Su & Saunders 2009; Surveswaran et al. 2010). Erkens et al. (2012b; ?program) found little in the way of changes in diversification rates in the family, speciation behaving as if it were a random branching process, and so there was little need to invoke key innovations.
In any event, it is clear from the above dates that much diversification happened after the break-up of the continents and is Tertiary in age. Couvreur et al. (2011a) contrast the diversification in Annonoideae and Malmeoideae. The former began to diversify ca 66 m.y.a. and the latter much later, ca 33 m.y.a.; the former has about twice as many species as the latter, even if rather paradoxically the latter may have a higher diversification rate - once it got going (see Couvreur et al. 2011a; Thomas et al. 2012, older, but similar disparity, both give other ages). However, although stem age estimates of these subfamilies are in the 70 million year range, crown ages vary widely, Couvreur et al.'s age being the youngest, and Piries and Doyle (2012) the oldest, a crown group estimate of (70-)66(-62) my. Su and Saunders (2009) offer dates for additional splits in the family, focusing on Pseuduvaria, and Thomas et al. (2012) give many dates for divergences within both main subfamilies.
The Old World Uvaria may have originated (38.4-)31.6(-25.1) m.y.a. in Africa, whence it dispersed to other parts of the Old World (Zhou et al. 2012: HPD). The large New World genus Guatteria has undergone much speciation in (Amazonian) South America perhaps 8.8-4.9 m.y. before present, having moved to South America from Central America, perhaps from Africa via Europe (Erkens 2007; Erkens et al. 2007a, b, 2009).
Ecology & Physiology. Annonaceae are important lianes of the tropical South East Asian forests (Gentry 1991), and there have been several origins of this habit within Annonoideae (Meinke et al. 2011). Artabotrys, one of these lianes, has curved hooks on the ultimate branches; these are modified inflorescences. The orthotropic stems may have stout, paired and rather vicious thorns. The other lianes do not have hooks or thorns, and development of the liane habit varies.
The family is notably common in terms of both numbers of species (number three in the list) and individuals in the Amazonian tree flora, but includes only 4 of the 227 species that make up half the stems in the forests there (ter Steege et al. 2013).
Pollination Biology & Seed Dispersal. Saunders (2010: good photographs) summarizes variation in floral morphology in the context of pollination. Flowers in which the stamens are enclosed in some sort of bowl-shaped structure are common, and there are often floral rewards on the inner surfaces of the inner perianth whorl. Pollination of the more or less odoriferous flowers is predominantly - and perhaps ancestrally - by small beetles, but there is also fly pollination (e.g. Su et al. 2008) and pollination by thrips, bees, cockroaches, etc. (Saunders 2012; Gottsberger 2012); a variety of scents is produced (Goodrich 2012 and references). The tough, apically expanded connective that predominates in stamens of flowers of Annonoideae and Malmeoideae may protect the pollen from the depradations of unwanted visitors (Gottsberger 1999, 2012; Silberbauer-Gottsberger et al. 2003). The flowers of some Annonaceae show thermogenesis (Seymour 2001), and some kind of extracarpellary compitum is common (Deroin 1991). The internal staminodes of Anaxagorea cover the gynoecium in the staminate floral phase, apparently preventing selfing (Gottsberger 2012; Saunders 2012). For pollen evolution, see Doyle et al. (2000) and especially Doyle and le Thomas (2012).
Dispersal of fruit is predominantly by mammals and birds. The lianes of tropical South East Asia are notably important food resouces for frugivores because they fruit more or less continually (Leighton " Leighton 1983).
Vegetative Variation. The main axes of Annonaceae are often mixed (?Troll's model), initially being orthotropic and with spirally-arranged leaves, later being more or less plagiotropic with two-ranked leaves (see also Eupomatia). Taxa such as Asimina and Annona have orthotropic axes with 2-ranked leaves. In general, axillary branches depart from the stem slightly to one side of the axil. Branches generally have plagiotropic leaves, and in some Xylopia, etc., the plagiotropic branch systems are frondose and very beautiful.
Chemistry, Morphology, etc. Protoberberidine alkaloids are reported from Annonaceae (Wink 2008). Cork in the roots of Goniothalamus may be superficial (Blunden & Kyi 1974). The wood occasionally fluoresces. Terminal tracheids are fairly commonly silicified (Piperno 2006). For the petiolar anatomy of Anaxagorea, see Jovet-Ast (1942) and Maas and Westra (1984); it is unlikely that there are annular petiolar bundles, although the midrib anatomy can be complex. Tetrameranthus sometimes seems to have opposite leaves; according to George Schatz (pers. comm.) it lacks plagiotropic branches and all branches have spirally-arranged leaves.
There are reports that the pollen grains of both Annona and Asimina may germinate via the proximal pole (Hesse et al. 2009a), although in the former (and some other genera) individual grains of the tetrad may rotate during development (Tsou & Fu 2002, 2006; Lora et al. 2009b). Xylopia, like Anaxagorea also with follicles, has micropylar-arillate seeds (Corner 1976); other genera may also have reduced arils (Svoma 1997). Reports that the micropyle is formed by both integuments need to be confirmed (c.f. Svoma 1998b). Monodora (derived) has a rather thin outer integument, and anatomy of the seed coat of Ambavioideae with a third integument (Zwischenintegument) can be complex (Christmann 1986; see also Perisamy & Swamy 1961).
For an introduction to Annonaceae and much more besides, see the papers in Bot. J. Linnean Soc. 169(1). 2012, including Erkens et al. (2012a) for a comprehensive bibliography; see Kessler (1993) and Bakker (2000a, b) for general information. Also Fries (1919: inflorescence morphology), Jovet-Ast (1942: indumentum and anatomy), Corner (1949: seed anatomy), van der Wyck and Canright (1956: anatomy), Hegnauer (1964, 1989: chemistry), Okada and Ueda (1984: cytology), Morawaetz (1986, 1988: esp. cytology), Deroin (1991b: stigmatic surface and carpel morphology), Steinecke (1993: esp. flower and fruit), Rudall and Furness (1997: tapetum), Svoma (1998a: seeds), van Setten and Koek-Noorman (1992: fruits and seeds), Endress (2011b: ovules), Deroin (1999a: receptacular vascular system), Johnson (2003: leaf insertion), Junikka and Koek-Noorman (2007: bark anatomy), Koek-Noorman and Westra (2012: wood anatomy), van Setten and Koek-Noorman (1986: leaf anatomy), Couvreur et al. (2008a: evolution of syncarpy, etc.), Couvreur et al. (2008b: pollen in the Monodora group), Sun et al. (2008: epidermal anatomy), and Lora et al. (2009: infraspecific variation in number of pollen nuclei in Annona cherimola). For floral (developmental) morphology, see van Heusden (1992), Ronse Decraene and Smets (1990b), Leins and Erbar (1980, 1982, 1996, 2010), Xu and Ronse De Craene (2010b), Endress (1975: carpel and stamen development), and Endress and Armstrong (2011: Anaxagorea).
Phylogeny. See Doyle and le Thomas (1994, 1996) for morphological phylogenies and character evolution, and for a molecular phylogeny of Anaxagorea that is integrated with morphological variation, see Scharaschkin and Doyle (2005, 2006). The position of Unonopsis on the tree has been somewhat erratic, perhaps because of ancient paralogy (Pirie et al. 2007). For relationships in Ambavioideae, see Surveswaran et al. (2010).
Two major clades make up the rest of the family (see also Richardson 2004). Malmeoideae include a polyphyletic Polyalthia (Richardson et al. (2004). This clade is not very speciose, mostly lacking large genera aside from Polyalthia, and has relatively little internal molecular divergence (hence its early name, the short-branch clade: Mols et al. 2008a; Xue et al. 2011). New World members of this clade form a monophyletic group (Pirie et al. 2006). For Pseuduvaria, see Su and Saunders (2006) and Su et al. (2008, 2010), while for the relationships and dimemberment of Polyalthia, see Xue et al. (2011, 2012). Mols et al. (2008b) explore character evolution within Malmeoideae. Annonoideae include the inaperturate pollen clade; here there is more molecular divergence (hence its name, the long branch clade: see also J. A. Doyle et al. 2004; Pirie et al. 2005). Uvaria is polyphyletic (Mols et al. 2004), and Zhou et al. (2009a, esp. b, 2010) suggested that its limits be extended; geography and phylogeny correlate quite well. The limits of genera like Dasymaschalon and Friesodielsia are also unclear (Wang et al. 2012: indepdendent loss and subsequent parallel evolution in taxa that have lost their inner petals). )For Guatteria, see Erkens (2007) and Erkens et al. (2007a, b).
Classification. The classification outlined by Chatrou et al. (2012) is followed here, although they also recognize tribes; their principles of classification are clearly explained. The limits of genera and hence the numbers of their included species in Annonoideae seem particularly uncertain. Zhou et al. (2010) clarify the limits of Uvaria, Xue et al. (2011, 2012) those of Polyalthia, while Surveswaran et al. (2010) discuss generic limits in the ambavioid clade. For a checklist of the New World species of the genus, see Maas et al. (2011); Annonbase should also be consulted.


text:
Tesco Chenille Throw- Stone 125X160Cm
Catalogue Number: 324-8581
- 125x160cm
- Acrylic
- machine washable
Product Details
This chenille throw is made from acrylic, and can be used to add warmth and a touch of colour to your room. This throw comes in an elegant stone finish. This chenille throw is machine washable.
back to topProduct specifications
Variant | |
Colour | Stone |
Key Information | |
Pack Size | 1 |
Type | Throw |
Material | |
Cover Material | Acrylic |
Features | |
Tumble Dry | No |
Machine Washable | Yes - wash at 30°C |
Dimensions | |
Product Height | 160.0 Centimetre |
Product Width | 125.0 Centimetre |
Product Depth | 1.0 Centimetre |
Product Weight | 0.54 Kilogram |
Buy from Tesco
- £10.00
- 10 Clubcard points
- in stock
You might be interested in these items
Customers who bought this also bought
Who are the Sellers at Tesco?
Sellers at Tesco are third party sellers we have invited to sell on Tesco direct. These are companies we've selected to provide you with a wider range of products to choose from in one place. They are a mixture of well-known high street retailers and specialist retailers who operate online. Plus, you earn Clubcard points on all seller orders.
Seller deliveries & last order dates for Christmas
Delivery times and costs vary by seller – further details are shown on the product pages and at checkout.
We know it's important to get your gifts in time for Christmas. The last order date for Christmas delivery varies by seller, please see Delivery information for more details.
For sellers offering Click & Collect, the last order date is 1pm on Friday 20th December.If Click & Collect is available from a seller, it will show on the product page and at checkout.
The delivery and return of seller items is managed by each seller. For further details, or if you have a question about the delivery or return of a seller product, please use the Seller directory to find the seller's 'help' page.

subset: commoncrawl


text:
- Join the France 24 community here
- Log in
Latest update: 18/04/2011
- Freedom of the press - journalists - Libya - media - Morocco - Muammar Gaddafi - NATO - Tunisia - war
Libya: the battle for Misrata
In Libya, the insurgents in Misrata have called for more help from NATO. Now the West is asking itself whether or not to arm the rebels. In Tunisia, residents of Sidi Bouzid say they have hardly seen any change since the fall of Ben Ali. And finally, a wind of liberty has been blowing through Morocco for a month now, and journalists are trying to make the most of it.
React to the article
(1) Reaction
To arm rebles means to arm al-qaeda
by Anonyme - 18/04/2011 - 10:43
it is very danger to arm libya rebles,short feature they will tern against europe and America


text:
Results 1 to 5 of 21
Thread: Strong and unusal boy names?
September 20th, 2013 02:45 PM #1
Strong and unusal boy names?
We just found out we are having a boy! Now I need some help widening my list of names to talk over with my husband. He wants something strong, I want something uncommon. We both like names that are easy to spell/pronounce, but I don't want anything near the top 100. Our last name starts with S, which kind of puts a damper on some of the names we actually both like and agree on, Atlas, Caius, Ramses. I like Rennon, he thinks that it's not quite "strong" enough, he likes Perrin (a character from a book) I think it's no stronger than Rennon!
Any ideas for us??Setra's mom
Baby boy due in February--searching for a name!
September 21st, 2013 10:42 AM #3
Last edited by averelladavina; September 21st, 2013 at 11:17 AM. Reason: can't spell first thing in the morning
Top Girl names:
~Combos I love~
Elizabeth Winona "Elsie" ~ Adelheid Ruby "Addie Rue"
Marilla Delyth "Millie" ~ Casilda Josephine "Cassie Jo"
~Other Firsts~
Hildemara ~ Isannah ~ Magnolia
Top Boy names: (a work in progress)
~Boy combos I'm considering~
Gideon Walker "Gid" ~ Evander Lane "Evan" ~ Malcolm Wesley "Cal" ~ Alaric Marion "Lark"
~Other Possible Firsts~
Mathias ~ Reuben ~ Gilbert ~ Cadmar
September 21st, 2013 11:07 AM #5Sorry for mistakes, I am foreigner.
my girls list
my boys list
If you like unusual and remarkable names, use them!
September 21st, 2013 11:15 AM #7Senior Member
- Join Date
- Sep 2013
- Posts
- 297
September 21st, 2013 12:43 PM #9


text:
Anyone with teenagers knows how hard it is to get them out of bed in time for a 7 a.m. start time. But do you think schools should start later to allow them to get more sleep?
State Superintendent Mike Flanagan thinks so.
He agrees with U.S. Education Secretary Arne Duncan, who has been touting this idea in recent months, most recently yesterday on National Public Radio's "The Diane Rehm Show."
"There's lots of research and common sense that a lot of teens struggle to get up at 6 in the morning to get on the bus or 5:30 in the morning to get on the bus," he said in the interview.
According to a release today from the Michigan Department of Education, Flanagan has been encouraging local school districts to adjust their teaching and learning schedules to accommodate the individual learning needs of the students for the past for years.
The release said he has shared this philosophy on a number of occasions with Secretary Duncan.
"I am encouraged that Secretary Duncan is advocating the change from what has been considered a traditional learning day for students," Flanagan said. "The Secretary understands this and I am hopeful that schools in Michigan will begin to understand the value, and adopt more flexible instructional schedules for their students. It makes sense."
Do you think high schools in Michigan should start later?


text:
Season 1 Ep 10
Season 3 Ep 21
Thursday December 5, 2013
Read more news at 3News
New Zealanders are amongst the happiest people in the world and a solid 3.14% of this jubilation is due to the Old Mout Cider Comedy Gala
TV3 is proud to have supported the first annual Ride to Conquer Cancer which took place over the weekend. The Ride has been declared the largest single fundraising event in New Zealand history!
The BIG DAY OUT wants to send you and four mates to the Big Day Out 2014 - Friday January 17; Western Springs, Auckland.
To make the happiest night of the year even happier, TV3 want to give you the chance to enjoy the celebrations in the VIP Front of House zone at Coca-Cola Christmas in the Park.
Copyright © 2013 MediaWorks TV - All Rights Reserved
Standard Competition/Promotion Terms and Conditions | Website Access Policies | Employment Opportunities | Online Advertising | Complaints


text:
EU hits screen makers with record cartel fine
- Published: 5 Dec 2012 at 19.47
- Online news: World
The European Commission on Wednesday hit seven top TV and computer screen makers, including LG Electronics and Philips, with a record fine of 1.5 billion euros for running decade-long price-fixing cartels.
Cathode ray tubes to be used for TV and computer monitors on a production line in Suwon, South Korea. European Union anti-trust authorities announced Wednesday a record fine for running a cartel, ordering seven companies including LG Electronics and Philips to pay nearly 1.5 billion euros ($1.9 billion) for jacking up the prices for TVs and computer screens for a decade.
"It's the biggest fine for a cartel decision," EU Competition Commissioner Joaquin Almunia said.
This article is older than 60 days, which we reserve for our premium members only.You can subscribe to our premium member subscription, here.


text:
This product has additional required/recommended options. To configure, add to your cart.
Drain Assembly Required
Grab Bar Required
Sonata® one-piece shower module with integral high-dome ceiling, less grab bar kit
Make the most of a compact space with the Sonata corner shower module, featuring an integral high-dome ceiling. Integral ledges provide ample room for shampoo and other shower supplies. This module's one-piece construction allows for easy installation and the recessed nailing flange provides for a smooth, finished look once installed.
- 36"L x 36-1/2"W x 90"H
- Module is pre-drilled for the Sonata accessory kit, consisting of grab bar and shelf kit
Additional Kohler Links
- View Manufacturer Warranty
- Shop All Kohler Products
- Shop All Kohler Sonata Collection Products
Our SKU: Kohler K-1689
This product is listed under the following manufacturer number(s):
- Kohler K-1689-47
Almond - Kohler K-1689-96
Biscuit - Kohler K-1689-95
Ice Grey - Kohler K-1689-G9
Sandbar - Kohler K-1689-0
*Denotes a finish or option that has been discontinued
Product Specifications
Installation Guide | |
Rough-In Guide | |
Specification Sheet | |
Configuration | One Piece |
Country Of Origin | USA |
Depth | 36.5 |
Grab Bars | No |
Height | 90 |
Made in America | Yes |
Material | Acrylic |
Theme | Modern |
Width | 36 |


text: Home History Who Was the First Black Saint? 100 Amazing Facts About the Negro: We know his name, and why a Swiss resort town is named after him. By: Henry Louis Gates Jr. Posted: Oct. 29 2012 12:14 AM 0 Matthias Grunewald (artist). St. Erasmus and St. Maurice. From Halle, Collegiate church. Ca. 1520-24. Painting on wood Editor's Note: For those who are wondering about the retro title of this black history series, please take a moment to learn about historian Joel A. Rogers, author of the 1934 book 100 Amazing Facts About the Negro With Complete Proof, to whom these "amazing facts" are an homage. (The Root) -- Amazing Fact About the Negro No. 3: Who was the first black saint? The jet-setters who frequent the Swiss resort town of St. Moritz are no doubt unaware that it was named after the first black person to be documented as a saint in the Roman Catholic Church. He was known as Saint Maurice (aka St. Mauritius, or St. Moritz). A third-century Roman legionary who was born in Thebes in Upper Egypt, Maurice was martyred in what is today Switzerland for refusing to massacre Christians for the Roman Empire. He was canonized by the early church, long before the Pope reserved the right of the Holy See exclusively to canonize souls in 1634. In the late fourth century Theodore, the bishop of Octudurum, had a vision in which he saw the martyrdom of Maurice and his fellow soldiers at Agaunum, Switzerland. Soon after, the cult of Maurice was established and a church built on the site. Then, in the early sixth century (ca. 515), devotees established an abbey there on land donated by King Sigismund of Burgundy. The abbey is still an active monastery and pilgrimage site, and the tomb of St. Maurice has been excavated. Maurice's feast day is Sept. 22. Maurice was named a saint in the early Middle Ages, and many early depictions were of him as a white man. The first representations of him as a black African appear in the middle of the 13th century, when he is depicted as a black African soldier, in a magnificent stone statue in the Magdeburg Cathedral in central Germany in about 1240 A.D., standing next to the grave of Otto I, Holy Roman Emperor. His cult became important in many places across Germany through the 16th century, and hundreds of images of him stand in European churches. He is the patron saint of soldiers, swordsmiths, armies, infantrymen, weavers and dyers, and is the patron saint of several towns in Europe, including Manresa, Spain, and Piedmont, Italy. The Image of the Black in Western Art Archive in the Du Bois Institute at Harvard University includes 229 representations of Maurice, compiled by Gude Suckale-Redlefsen and published as The Black Saint Maurice in 1988. Most extant images of him are found in central Germany, but seven are in the Czech Republic, six in Denmark, three in Sweden, two each in Norway, Finland, Austria and Poland and one each in France, Estonia and Latvia. Scholars believe that Maurice started being depicted as a dark-skinned African at this time because the Holy Roman Emperor, Frederick II Barbarossa, had ambitions to rule the entire world, just as the Pope claimed spiritual dominion over the entire world. We know that Frederick had black Africans at his court and in his retinue. Magdeburg, where the first image of Maurice as a black African appeared, was on the frontiers of the Empire, and was a region of military expansion. For whatever reasons, though, the black Saint Maurice became the subject of a masterpiece of Western art in Magdeburg and the subject of some of the greatest works of art created during the late Middle Ages and the Renaissance. Page 1 of 2 NEXT PAGE > Powered by Livefyre


text:
The REI Member Dividend
The REI dividend is a unique benefit of membership in the REI co-op. It's comprised of your member refund—typically 10% back on eligible purchases—and/or your REI Visa® card rebates.
Each March, active REI members and REI Visa cardholders who earned a rebate receive a refund in the form of an annual dividend notice.
Only active members receive annual dividend notices. To maintain your active status as an REI member, you need to make net merchandise or shipping purchases of at least $10 per year, unless you joined that year. Member numbers are issued in one name and only that person receives the member refund. Your number can be used by others in your family, but it cannot be transferred to someone else.
Questions? Call us at 1-800-426-4840 for more about your dividend or its redemption.
Item 709207
Specification |
| |||
Best use |
| |||
Material(s) | ||||
Dimensions |
by PowerReviewsPros
Best Uses
Reviewed by 6 customers
Sort by
Displaying reviews 1-6
Best Uses
Comments about Pacsafe 55 Security Web - Small:
I bought this for my 25L backpack. I like that someone will not be able to easily steal my backpack.
However, the gaps (holes) in the mesh are large enough for me to put my hand through and I don't have small hands. So, without much effort someone could empty the contents of my backpack.
I may make my own with stainless cables etc and make the holes about the same size as chicken wire. If Pacsafe did this I would give this thing 5 stars.
Best Uses
Comments about Pacsafe 55 Security Web - Small:
I purchased this item prior to a trip to England during July 2010.
My carry-on for the plane was a camera backpack loaded with fairly expensive gear. Call me paranoid, but knowning my gear was secure, even while on a plane, was a comfort. How easy would it be for someone to steal a lens while you're in the lavtory or asleep? Very without this type of product. The mesh openings seem to be small enough to prevent most valuable from being pulled through.
While traveling through England, the PacSafe continued to put my mind at ease since it provided more security than I ever had before.
Can the cable be cut easily with some tools? Probably. But outside of that, this is a really good solution to ensure your gear cannot be accessed quickly. This creates a good amount of piece of mind.
Oh yeah, no issues getting through security in the US or England even though my backpack was wrapped in the PacSafe.
Well worth the money!
Best Uses
Comments about Pacsafe 55 Security Web - Small:
As a compromise between driving a car & riding a bike/bus, I ride a scooter all year round, barring ice or snow. Riding a motorcycle (even a small one) requires wearing bulky, heavy gear & boots. When I go out to attend social events I either risk body parts & ride "exposed" or I go looking like the Terminator. When I run errands on the weekends I sometimes have to make several trips to offload stuff, wasting time & gas.
Small items I can lock inside the storage compartment under the seat. This handy gadget allows me to lock a change of clothing & shoes, a bag of cat food, groceries, whatever, to the scooter using one of the helmet pegs. Is it perfectly secure? Of course not. A determined thief can cut it off the bike. Although, cutting through stainless steel wire isn't as easy as you might think. Leave stuff unattended on the scooter all night? No. Lock up my riding gear for a couple of hours while I go out for dinner? Yes. I'd trust it to secure my laptop on a train, but I wouldn't trust it to secure a wallet, cell phone, digital camera or other valuables. Use common sense.
The mesh is big, so I put my stuff in a gym bag and put the net around it. Keys, small valuables go in my pocket or purse & come with me. It fits nicely in its holder, doesn't take up much space. I keep it in the storage compartment along with the folded gym bag.
Best Uses
Comments about Pacsafe 55 Security Web - Small:
First of all, I was very impressed at how intuitive this product is to use. And, as it unfolds and packs up very quickly, I was happy to find that I have been highly motivated to use it -- an issue that I had been concerned about before I bought the product about one month ago.
Best of all, I felt I was able to totally relax for the first time in quite awhile during a recent train trip, knowing my laptop and, sometimes, digital SLR/lenses were secure in my daypack while I wandered about the train or went to sleep. Ditto for when I stayed in a couple of hotels. Until now, I hadn't realized how much energy I had been wasting, worrying about whether someone was going to walk off with my stuff. I would actually port my laptop around all day while on vacation because of this fear. No more! I wish I had known about this product earlier.
This PacSafe will definitely come in handy during upcoming stays in budget lodgings, which won't often offer safes either in the room or at the front desk.
I did replace the accompanying keyed padlocks with a combination padlock, though, as I hate fumbling with keys.
Best Uses
Comments about Pacsafe 55 Security Web - Small:
I bought this for an upcoming trip to Costa Rica after seeing some travelers at an airport put in on their checked backpacks. It fit my rolling bag/packpack perfectly and was so easy to use. I want it mostly for security when I'm away from my room or bag. A frequent friend/traveler said he won't leave home without it.
Best Uses
Comments about Pacsafe 55 Security Web - Small:
I bought this "wire net" to protect my laptop in my backpack when I leave it behind in my hotel room (my passport/credit cards/cash I take with me in a money belt). It completely wraps my bag, and protects anything left in the pockets giving me peace of mind when I am out exploring. Two downsides:
1) It comes with a keyed lock. I don't want to keep track of keys (or lose them) when I am traveling so I replaced it with a combination lock.
2) It is a little bit heavy. I am an ultra-light traveller, and I don't usually bring something along of this weight. Still, for the peace of mind it gives me, it is worth it in my opinion.
Displaying reviews 1-6
How are we doing? Give us feedback on this page.


text:
We're sorry!
There aren't any tickets listed right now for CommBank Ashes Series: Australia vs. England at WACA Ground in East Perth, Australia on Friday, December 13, 2013. Sign up for an alert, and we'll let you know when tickets are available!
Note: We will NOT share your email address with any third-party vendors. For more details, see our Privacy Policy.


text:
Shiner Wild Hare Pale Ale - Spoetzl Brewery
Displayed for educational use only; do not reuse. |
Ratings & Reviews |
Sort by: Latest | High | Low | Top Reviewers | Alström Bros |
« first ‹ prev | 1-25 | 26-50 | 51-75 | next › last » |
Ratings: 487 | Reviews: 95 | Show All Ratings:
Reviews by DavidBart:More User Reviews:StonedTrippinColorado 3.61/5 rDev +7.4% look: 3.5 | smell: 3.5 | taste: 3.75 | feel: 3.5 | overall: 3.5 ive done my share of bashing on shiner, we all have, but ive had three solid beers from them lately, so it time for reassessment. yeah, this pale ale isnt zombie dust, but its pretty darn good. clear amber tinted pale beer with a passable white head. the nose is citrusy hops, light and fruity with very little bitter. slightly richer malt profile gives some body. the hops really shine in the flavor, really citrusy, lemons and limes even, its quite a nice surprise. malt supports it with a touch of english sweetness, but the finish is clean, almost like a lager, a delicious combination, highly drinkable, and one of the better shiner brand brews ive gotten into. a six of this on a hot day would be enjoyable work. time to give shine another chance folks. Serving type: bottle 12-16-2013 19:00:32 | More by StonedTrippin UCLABrewN84California 3.29/5 rDev -2.1% look: 3.5 | smell: 3 | taste: 3.25 | feel: 4 | overall: 3.25 Pours a clear honey orange with a foamy khaki head that settles to a partial film on top of the beer. Small streaks of lace form around the glass on the drink down. Smell is of malt, citrus fruit, and medicinal yeast aromas. Taste is much the same with malt, yeast, and earthy citrus zest flavors on the finish. There is a mild amount of hop bitterness on the palate with each sip. This beer has a good level of carbonation with a crisp mouthfeel. Overall, this is an above average beer that is fairly drinkable but the yeast presence in the aroma and flavor somewhat dominate things in this one. Serving type: bottle 12-09-2013 06:29:01 | More by UCLABrewN84 tone77Pennsylvania 3.53/5 rDev +5.1% look: 4 | smell: 3.5 | taste: 3.5 | feel: 3.5 | overall: 3.5 Poured from a brown 12 oz. bottle. Has a dark golden color with a 1/2 inch head. Smell is of hops, a bit of fruit. Taste is well balanced. Hops coming first, mild bitterness, strong malt backbone at the finish. A touch of fruit is there as well. Feels medium bodied in the mouth and overall is a solid beer. Serving type: bottle 12-08-2013 03:12:28 | More by tone77 |
« first ‹ prev | 1-25 | 26-50 | 51-75 | next › last » |
Shiner Wild Hare Pale Ale from Spoetzl Brewery
out of 100
based on 487 user ratings.


text:
"Internet Purchases Only" "Not a Retail Store Stock Item"
This item is viewable at our retail store in LEXINGTON, KY. It may be on display and is available for immediate purchase and pick-up.
click here to see all brands we sell
*Please Note!! Many of our pictures are stock photos provided to us by the manufacturer and do not necessarily represent the actual item being purchased. Please verify this picture accurately reflects the product described by the title and description on this page before you place your order.
Know your distance. Take your shot. Weaver's 8x28mm 1,000-yard laser rangefinder gives you accurate, reliable and user-friendly performance in the field. The laser rangefinder operates on one standard CR2 battery, is IPX3 waterproof, ranges moving targets (in scan mode) and measures distance in feet, yards or meters. Able to range deer-sized animals out to 600 yards and reflective targets to 1000 yards, the Weaver laser rangefinder provides the confidence you need to make your shot.


text:
Title III Part B, Strengthening Historically Black Graduate Institutions Program
| ||||||||||||||||||||||||||||||||||||||||||||
| ||||||||||||||||||||||||||||||||||||||||||||
Program Office: Institutional Service
CFDA Number: 84.031B
Program Type: Discretionary Grants
Also Known As: HBCUs and HBGIs
Program Description
This program provides financial assistance to Historically Black Graduate Institutions(HBGIs) to establish or strengthen their physical plants, financial management, academic resources, and endowment-building capacity. Activities may include:
- Student services; educational equipment acquisition; facility renovation and construction; faculty and staff development; the establishment of a program of teacher education designed to qualify students to teach in public schools;
- The establishment of community outreach programs that will encourage elementary and secondary school students to develop the academic skills and the interest to pursue postsecondary education;
- The acquisition of real property in connection with the construction, renovation, or addition to or improvement of campus facilities;
- Families, especially with regard to student indebtedness and student assistance programs under Title IV; and
- Services necessary for the implementation of projects or activities that are described in the grant application and that are approved, in advance, by the Department, except that not more than two percent of the grant amount may be used for this purpose.
Types of Projects
Funds may be used for the purchase, rental, or lease of scientific or laboratory equipment. Also supported are the construction, maintenance, renovation, and improvement of instruction facilities. Funds support faculty exchanges and the development of academic instruction in disciplines in which black Americans are underrepresented. Projects may support the purchase of library materials as well as tutoring, counseling, and student service programs. Also supported are: funds and administrative management; joint use of facilities; establishment or improvement of development offices; establishment or enhancement of programs of teacher education; establishment of outreach programs; and other activities that a grantee proposes in its application that contribute to carrying out the purposes of the program and are approved by the Secretary as part of the review and acceptance of the grant application.
Additional Information
The Strengthening Historically Black Graduate Institutions Program provides grants to the institutions listed under Eligibility to assist these institutions in establishing and strengthening their endowment funds, academic resources and student services so that they may continue to participate in fulfilling the goal of equality of educational opportunity in graduate education.
Funds may be used for the:
- purchase, rental or lease of scientific or laboratory equipment for educational purposes, including instructional and research purposes;
- construction, maintenance, renovation and improvement in classroom, library, laboratory and other instructional facilities, including purchase or rental of telecommunications technology equipment or services;
- purchase of library books, periodicals, technical and other scientific journals, microfilm, microfiche, and other educational materials, including telecommunications program materials;
- scholarships, fellowships, and other financial assistance for needy graduate and professional students to permit the enrollment of all the students in and completion of the doctoral degree in medicine, dentistry, pharmacy, veterinary medicine, law and the doctorate degree in the physical or natural sciences, engineering, mathematics, or other scientific disciplines in which African Americans are underrepresented;
- establish or improve a development office to strengthen and increase contributions from alumni and the private sector;
- assist in the establishment or maintenance of an institutional endowment to facilitate financial independence pursuant to section 331; and
- funds and administrative management, and the acquisition of equipment, including software, for use in strengthening funds management and management information systems.
Program Map of Participating Institutions -- This Google map shows the 24 institutions eligible to apply for and receive discretionary grants awarded under this section by Institutional Service (IS), U.S. Department of Education. Institutions receiving awards under the Strengthening Historically Black Graduate Institutions (HBGI) Program only were awarded in FY 2007. Institutions awarded grants under both the Historically Black Colleges and Universities (HBCU) and HBGI Programs were awarded their HBGI grant award in FY 2009. HBGI grants, regardless of when awarded, were awarded for a period of five years.


text:
Comings and Goings is a blog written by Theology, Worship and Education Director Charles B. "Chip" Hardwick as he travels throughout the church. God is on the move out and about in the world, working to redeem all things in Jesus Christ. As we join this mission, by the power of the Spirit we see God on the move. This blog contains glimpses of how Chip finds this to be true in his comings and goings.
Can we possibly find God in popular culture? In what ways can books, movies, and TV shows help us to understand who God is?
And what does mid-20th century theologian Karl Barth have to do with all of this?
Check out my recent article in Presbyterians Today to get some insights on these questions.