Caryophyllids What are caryophyllids? Diversity and Evolution • First of the core eudicots we will examine: Caryophyllids, of Caryophyllids Rosids, Asterids core eudicots . carnations, cacti, chenopods . • = order Caryophyllales • APG III in 2009 places caryophyllids as sister lineage to the asterids – but probably dates back to 110-100 mya Caryophyllids Caryophyllids What are caryophyllids? What are caryophyllids? • 34 families & 11,155 species = • Exhibit unusual adaptations to “stressful” 6% of eudicot diversity environments — desert or arid regions: high salt, low water, xerophytes • Unusual (!) group of families core eudicots not all previously thought to form a natural order • Share one character? sepals only - “petals” if present appear to be of staminal origin saltbush- Amaranthaceae cacti- Cactaceae 1 Caryophyllids Caryophyllids What are caryophyllids? What are caryophyllids? • Exhibit unusual adaptations to “stressful” • Exhibit unusual adaptations to “stressful” environments — salt marshes, halophytes environments — alpine, tundra, cushion plants glasswort- Amaranthaceae chickweed- Caryophyllaceae spring-beauty- Montiaceae Caryophyllids Caryophyllids What are caryophyllids? What are caryophyllids? • “new” (unplaced) members to the group • “new” (unplaced) members to the group include desert families include desert families Simmondsia chinensis jojoba Frankenia laevis Simmondsiaceae Frankeniaceae Sonoran Desert endemic Canary Islands 2 Caryophyllids Caryophyllids What are caryophyllids? What are caryophyllids? • “new” (unplaced) members to the group • “new” (unplaced) members to the group include desert families include desert families Tamarix - tamarisk Halophytum Tamaricaceae Halophytaceae Caryophyllids Caryophyllids Suite of morphological and/or physiological What are caryophyllids? adaptations for life in the arid world - succulence, no • troublesome “weeds” leaves, C4 and CAM photosynthesis, salt excretion C4 and Crassulacean Acid Metabolism Tamarix - tamarisk Gypsophila - baby’s-breath 3 Caryophyllids What are caryophyllids? • some, but not all, of the carnivorous plants - low N Sundews - Droseraceae New Phytologist Research 847 variable 12 (mean annual precipitation) showed a comparatively Climate occupancy reconstruction results consistent pattern of relatively dry to intermediately wet clades We performed climate occupancy ancestral reconstruction anal- throughout the group. Indeed, only a few clades inhabiting wet yses on the phylogeny of 2843 taxa that included taxa with at ecosystems (in this case, the wet tropics) exist in the Caryophyl- least three sampled geographic coordinates (Figs 1–3). We con- lales, specifically small groups within the carnivorous clade, the ducted these analyses for visualization and for comparison with phytolaccoids, early-diverging Polygonaceae, and other small diversification and WGD results (see later). Results for individ- groups throughout the Caryophyllales. The principal compo- ual bioclimatic variables and principal components can be nent loadings are presented in Figs 2 and S5. Principal compo- found in Figs S2–S4. Bioclimatic variable 1 (mean annual tem- nent 1 (PCA1) showed significant differentiation throughout perature, Fig. 1) showed that there are several strong phyloge- the Caryophyllales, as, for example, early-diverging Polygo- • why this incredible diversity – ecology, physiology, habit, color? netic patterns of clades with preferences for colder or warmerNepenthesnaceae vs the– restAsian of Polygonaceae, pitcher early plant diverging Caryophyl- regions. For example, Polygonaceae, Caryophyllaceae, and laceae vs the rest of Caryophyllaceae, phytolaccoids vs Montiaceae each are dominated by taxa with preferences for Aizoaceae, and Portulacineae + relatives vs Cactaceae, to name a cold environments, although each also contains early-diverging few. These results generally reflect the extensive ecological diver- taxa with preferences for warm environments. By contrast, taxa sification throughout the group. They also reflect significant inhabiting warm environments predominate in Cactaceae, diversification in the temperate regions of the world, especially Amaranthaceae, Aizoaceae, the carnivorous clade (Droseraceae, within the Caryophyllaceae and Polygonaceae, contrasted with Drosophyllaceae, Nepenthaceae, Ancistrocladaceae, Dioncophyl- extensive diversification in the succulent lineages (especially laceae), and the phytolaccoid clade (Nyctaginaceae, Phytolac- Aizoaceae and Cactaceae) found in relatively dry and warm caceae, Petiveriaceae, Sarcobataceae, and Agdestis). Bioclimatic environments. Amaranthaceae 5,x v Caryophyllids Polygonaceae u t s Caryophyllid clade ff • examine all carnivorous plants r w 6 later – Halloween lecture! cc ee Plumbaginaceae bb [check out botanical Halloween Caryophyllaceae aa ii costumes for Extra Credit] hh z Carnivores 7 8 13 10 9 y 11,ff 12,gg Bioclim 1 • also look at Polygonaceae - Diversification shift (MEDUSA ML) d p e smart weed family Diversification shift (MEDUSA set) a 2 Putative duplication b • focus on “core Caryophyllales” f 4 1,n o j 3 m h l Phytolaccoids Core Caryophyllales g k Portulacineae i Aizoaceae Smith et al. 2018 Angiosperm Phylogeny Website Fig. 1 Chronogram of the Caryophyllales with putative whole genome duplications mapped along with identified diversification shifts. Diversification analyses were performed on the maximum likelihood tree as well as the bootstrap tree set and those shifts that were identified in both groups are shown. www.mobot.org/MOBOT/Research/A The• brancheswhy are this colored basedincredible on Bioclim variable 1diversity (mean annual temperature). – ecology, The numbers next physiology, to duplications correspond tohabit, the numbers incolor? Table 3 and the letters next to diversification shifts correspond to Table 4. Pweb/welcome.html • whole genome duplications & diversification shifts? Ó 2017 The Authors New Phytologist (2018) 217: 836–854 New Phytologist Ó 2017 New Phytologist Trust www.newphytologist.com 4 Core Caryophyllales Core Caryophyllales Caryophyllid clade • defined by 5 features but with 1. betalains Carnation -Caryophyllaceae some reversals anthocyanin Sea fig -Aizoaceae betalain Core Caryophyllales N containing – very different from flavonoids Core Caryophyllales Core Caryophyllales 2. free-central placentation (or 2. free-central placentation (or basal) = “Centrospermae” basal) = “Centrospermae” . generates capsule fruits capsule pyxis utricle 5 Core Caryophyllales Core Caryophyllales 3. curved embryo in ovule = 4. pollen shed in trinucleate stage campylotropous vs. most common 2-nucleate campylotropous ovule micropyle Core Caryophyllales Core Caryophyllales 5. sieve tube plastids with crystal Caryophyllid clade • problematic for family proteins surrounded by protein circumscription / recognition filaments • examine 3 groups phloem Core Caryophyllales 6 *Caryophyllaceae - Carnations *Caryophyllaceae - Carnations Caryophyllid clade • carnation or pink family - • dichasium inflorescence – usually a cyme herbs, often weedy Note 3 way Lychnis coronaria - mullein pink split, middle branch is oldest flower Minuartia - sandwort *Caryophyllaceae - Carnations *Caryophyllaceae - Carnations • dichasium inflorescence – usually a cyme CA 5, (5) CO 5 A 5,10 G (2-5) • leaves opposite, swollen nodes • 5 merous flowers, calyx fused +/- • corolla not fused, often lobed (staminal origin?) 7 *Caryophyllaceae - Carnations *Caryophyllaceae - Carnations CA 5, (5) CO 5 A 5,10 G (2-5) CA 5, (5) CO 5 A 5,10 G (2-5) • anthers of 1-2 whorls • free-central or axile placentation • 1 pistil of 2-5 carpels • capsule fruit opening by teeth or valves *Caryophyllaceae - Carnations *Caryophyllaceae - Carnations Huge family, 87 genera, 2300 species; widespread but Cerastium - mouse-ear chickweed characteristic of temperate and warm temperate regions of the Northern Hemisphere. Minuartia michauxii sand rockwort 8 *Caryophyllaceae - Carnations *Caryophyllaceae - Carnations Saponaria officinalis - bouncing bet, soapwort Silene cucubalus Silene vulgaris Dianthus armeria bladder campion white campion deptford pink *Amaranthaceae - amaranths *Amaranthaceae - amaranths • herbs, often halophytes or weeds, worldwide • flowers small, bracted, congested, CA 5 CO 0 A 5 G (2-3) • 174 genera & 2,050 species lacking corolla • includes chenopods (old Chenopodiaceae) • bisexual or unisexual, monoecious or dioecious Chenopodium - lamb’s quarter Salicornia - glasswort Chenopodium - lamb’s quarter 9 *Amaranthaceae - amaranths *Amaranthaceae - amaranths • fruit is 1-seeded circumscissle capsule • native, weedy, and horticultural species (utricle) or basal seeded achene Amaranthus - Froelichia - • calyx is persistant around the fruit amaranth cottonseed utricle Spinacea - spinach Chenopodium *Amaranthaceae - amaranths *Amaranthaceae - amaranths • native, weedy, and horticultural species • native, weedy, and horticultural species Cycloloma atriplicifolium – winged pigweed Beta vulgaris - beet Bassia (Kochia) - summer cypress 10 *Amaranthaceae - amaranths *Amaranthaceae - amaranths • native, weedy, and horticultural species • desert specialists & tumbleweed invasives Celosia - cock’s comb Gomphrena - globe amaranth Atriplex - saltbush Salsola - Russian thistle *‘Portulacaceae’ - Purslanes *‘Portulacaceae’ - Purslanes Caryophyllid clade • belongs to a succulent group of • herbs, succulents families • world-wide, especially western N. • family boundaries obscure - e.g. America, 50 genera, 500 species Montiaceae = spring beauties Portulaca