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Home , Agdestis, Alluaudia, Alluaudiopsis, Caryophyllaceae, Claytonia, Claytonia megarhiza

Caryophyllids What are caryophyllids? Diversity and • First of the core we will examine: Caryophyllids, of Caryophyllids Rosids, Asterids core eudicots

. . . carnations, cacti, chenopods . . . • =

• 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 = • 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? only - “” if present appear to be of staminal origin

saltbush- 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

glasswort- Amaranthaceae chickweed- spring-beauty-

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 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 - tamarisk Halophytum Halophytaceae

Caryophyllids Caryophyllids Suite of morphological and/or physiological What are caryophyllids? adaptations for life in the arid world - succulence, no • troublesome “weeds” , C4 and CAM photosynthesis, salt excretion

C4 and Crassulacean Acid Metabolism

Tamarix - tamarisk - baby’s-breath

3 Caryophyllids What are caryophyllids? • some, but not all, of the carnivorous plants - low N

Sundews -

New Phytologist Research 847

variable 12 (mean annual precipitation) showed a comparatively Climate occupancy reconstruction results consistent pattern of relatively dry to intermediately wet 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 , the ducted these analyses for visualization and for comparison with phytolaccoids, early-diverging , 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 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

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 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 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. Carnation -Caryophyllaceae some reversals

Sea fig -Aizoaceae Core Caryophyllales

N containing – very different from flavonoids

Core Caryophyllales Core Caryophyllales 2. free-central (or 2. free-central placentation (or ) = “Centrospermae” basal) = “Centrospermae”

. . . generates

capsule pyxis utricle

5 Core Caryophyllales Core Caryophyllales 3. curved embryo in = 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 – usually a cyme herbs, often weedy Note 3 way coronaria - mullein pink split, middle branch is oldest

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 , 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 opening by teeth or valves

*Caryophyllaceae - Carnations *Caryophyllaceae - Carnations Huge family, 87 genera, 2300 species; widespread but - 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 cucubalus armeria bladder campion white campion deptford pink

*Amaranthaceae - *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 Amaranthus - Froelichia - • calyx is persistant around the fruit 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 & invasives

Celosia - cock’s comb Gomphrena - globe amaranth Atriplex - saltbush Salsola - Russian thistle

*‘’ - 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 - Portulaca - purslane purslane

11 *‘Portulacaceae’ - Purslanes *‘Portulacaceae’ - Purslanes

CA 2 CO 5+ A 5+ G (2-5) Portulaca grandiflora rock rose (Argentina) now in family Montiaceae • 5 merous flowers (petals?) • capsule of various types w. NAm alpine spring beauty

Portulaca - pyxis valvate purslane

*‘Portulacaceae’ - Purslanes *Cactaceae - cacti

• New World stem fame flower succulents protected by spines • 100 genera / 1400 species Lewisii bitter-root

Montiaceae

Phemeranthus rugospermus

12 *Cactaceae - cacti *Cactaceae - cacti • fleshy, succulent, P ¥ A ¥ G (4) often epiphytes • no leaves, except • ‘hypanthium’ with many • spines or glochids • ¥ at areoles • epigynous flower

*Cactaceae - cacti *Cactaceae - cacti Rhipsalis P ¥ A ¥ G (4) • great diversity: columnar, prickly pear, barrel, • placentation parietal (rare!) • relationships emerging • fruit prickly pear chollo

saguaro

Pereskia

13 *Cactaceae - cacti *Cactaceae - cacti

• Caribbean “Pereskia” at • Caribbean “Pereskia” at base of family! – now called base of family! – now called Rhodocactus Rhodocactus

• then S. American Pereskia

*Cactaceae - cacti *Cactaceae - cacti

Selenicereus grandiflorus Queen-of-the-night 25 cm diameter flowers!

• Caribbean “Pereskia” at base of family! – now called Rhodocactus

• then S. American Pereskia

• then the rest of American Lophophora williamsonia taxa (mescaline)

14 *Cactaceae - cacti - African ‘cacti’

• upper midwest cacti • Madagascar & east Africa • convergent also with American Opuntia macrorhiza - Fouquieriaceae (ocotillo) plains prickly-pear Opuntia humifusa - Alluaudia Didierea eastern prickly-pear Alluaudiopsis

Alluaudia

* - pokeweed *Phytolaccaceae - pokeweed

Caryophyllid clade • Small family of / of • small petaloid sepals only CA 5 CO 0 A 10 G (5+) tropical and temperate regions • stamens 2X sepals • carpels +/- fused, each with one ovule

Phytolacca americana - pokeweed Phytolacca octandra

15 *Phytolaccaceae - pokeweed *Polygonaceae - smartweeds • CA 5 CO 0 A 10 G (5+) Caryophyllid clade • also look at Polygonaceae - • berry fruits smart weed family • often poisonous • 61 genera / 1,100 species • dyes (poke = puccoon: Algonquian for red/orange dye)

Persicaria amphibia – Rivinia - bloodberry Phytolacca americana - pokeweed water smartweed

*Polygonaceae - smartweeds *Polygonaceae - smartweeds

• herbs (few shrubs) of wet or arid regions • herbs (few shrubs) of wet or arid regions • alternate, simple leaves at swollen nodes • modified as sheath = ocrea

Persicaria hydropiper - water – Persicaria amphibia – pepper water smartweed water smartweed

16 *Polygonaceae - smartweeds *Polygonaceae - smartweeds

• flowers congested P 3+3 (5) A 3X G (3) • and Persicaria • basically 3 merous, tepals petaloid () largest genera • one-seeded, three-angled achene

Persicaria - smartweed

Persicaria amphibia – water smartweed - curly doc Rumex orbicularis - water doc

*Polygonaceae - smartweeds

• horticulturally important

Rheum rhabarbarum Garden esculentum locally adventive

17