Boraginales Phylogeny Poster Families and genera with plesio-/apomorphic and (non)diagnostic traits

herbs, shrubs P 5-merous, style gynobasic fr nutlet (eremocarps) ovary eremocarpic, locules 4 Boraginaceae SEE Boraginaceae Phylogeny Poster ovules 1/locule, seeds 4/fr ovules ≤ 4

placentation apical Boraginales I Wellstediaceae fr flattened capsule, dehiscence septicidal SW Africa and SW Arabian peninsula/Horn of Africa Wellstedia 6

usu. herbaceous usu. stigma single-branched/capitate habit spiny Codonaceae K lobes 10–12, C lobes 10–12; nectary chambers + seed transfer cells absent SW Africa Codon 2 lvs deeply lobed to 1-2 pinnate taproot C scales present, reduced or absent locules 2 Boraginaceae seeds 1-many Phacelia c. 200 plant base bulb-like from tubers Wellstediaceae lvs entire C scales absent sometimes v-shaped folds present on C tube Codonaceae seeds 2-many Romanzoffia 5 Hydrophyllaceae lvs opposite C scales absent Namaceae seeds 1-4 Draperia 1 Heliotropiaceae

annuals Lennoaceae seeds 5-15 Eucrypta II* Ehretiaceae annuals flw pendent, C becoming paper-like C scales absent Hoplestigmataceae seeds 6-15 elaiosomes Emmenanthe 1 Coldeniaceae annuals G bicarpellate Cordiaceae C scales present as reduced pairs at base of each filament fr capsule, loculicidal seeds 4-8 Ellisia 1

annuals Hydrophyllaceae seeds 5-15 Eucrypta I* annuals flw generally with appendages (reflexed) between sepals herbaceous C scales present or reduced to pairs of glands bordering each filament flw petals aestivation contorted seeds 1-8 Pholistoma 3 ovary 1-locular placentae enlarged, lining or filling ovary lvs pinnatifid to pinnate fr capsule appearing 2- or 5-celled roots fibrous C scales linear, pair bordering but free from filaments Americas (mainly SW N America) ovary 1-locular seeds 1-3 Hydrophyllum 8 annuals flw generally with appendages (reflexed) between sepals C scales present as pairs at base of each filament, sometimes reduced seeds 1-20 Nemophila 11

K lobes divided to base, very unequal, outer 3 cordate, inner 2 linear, strongly accrescent in fruit C scales linear, paired, free from filaments seeds 4-8 Tricardia 1

flw solitary on elongate peduncles C scales absent seeds many Hesperochiron 2 acaulescent, short pleiocorm C scales semicircular, paired, free from filament base seeds 2-4 Boraginales II Howellanthus 1 stamens sub-/unequal in length, sub-/unequally inserted on C tube, testa transfer cells + staminal appendages present or fr capsules filament bases dilated membranaceous or Nama c. 50 semi-cartilaginous, dehiscing loculicidally by 2 valves stinging hairs stamens equal, equally inserted on C tube S to C America Wigandia 6 Namaceae evergreen shrubs stamens (un)equal in length, (un)equally inserted on C tube ovary 1-/2-locular fr capsules cartilaginous, staminal appendages reduced or obsolete Eriodictyon c. 10 placentae narrow, partially or dehiscing completely dividing ovary loculicidally/ septicidally fr capsule appearing 2-celled by 4 valves subshrub flw petals aestivation imbricate densely glandular, sticky, ill-scented America seeds striate 1 (mainly southwestern N America) Turricula**

shrubs trichomes secreting strongly scented resin mericarpids apically winged NW Argentina Ixorhea 1

lianas or subscandent shrubs anthers with pubescent or papillose apex, connectives protracted Heliotropiaceae fr usually drupes S & C America, Caribbean Myriopus c. 25 annuals/shrubs or lianas (Tournefortia) style-stigma complex + usually Kranz chlorenchyma (C4) anthers with pubescent or papillose apex, pyrrolizidine alkaloids connectives protracted Euploca c. 100

anthers apically glabrous, connectives not protracted worldwide subtrop. to temperate zones (few in AUS) Heliotropium c. 300

flw 8-merous stamens in two series, thecae basally spreading Lennoa 1 Lennoaceae

flw 4–10-merous holorhizoparasites stamens in 1 series, thecae basally parallel 3 achlorophyllous Pholisma lvs reduced to cataphylls G ≥ 5 fr dehiscence circumscissile trees or shrubs SW N America K aestivation imbricate fr with 1–4 pyrenes Ehretia c. 40 perennials mostly evergreen small shrubs infl flw bisexual involucre persistent in fr ebracteose NW Argentina thyrsoids, Cortesia 1 partly reduced shrubs to 1 flw flw usu. blue endocarp anthers connate, buzz pollination multilayered AUS c. 20 ovules ≤ 4 Ehretiaceae Halgania shrubs or herbs (mainly prostrate) flw bisexual mostly shrubs or trees fr schizocarps separating into four (or fewer) dry nutlets America (mainly southwestern N America, Galapagos Isl.) pan(sub)tropical Tiquilia c. 28

trees or shrubs (partly lianescent) K aestivation valvate fr with 4 triangular pyrenes Neotropics and E Africa Bourreria 50

Hartmut H. Hilger, Prof. Dr. shrubs (partly lianescent), axillary spines Theodor C. H. Cole, Dipl. Biol. flw unisexual C America, Caribbean 9 Dahlem Centre of Plant Sciences (DCPS) trees/shrubs Rochefortia Institute of Biology – Botany Freie Universität Berlin Altensteinstr. 6, D-14195 Berlin, Germany trees, dioecious C America & northern S America Lepidocordia 2 • concept of order following the recommendations of the Boraginales Working Group trees, deciduous; buds perulate • hypothetical tree based on molecular phylogenetic data (2018) K connate, irregularly splitting, C 11-15, connate, opening irregularly, • characters taken from Kubitzki, ed. (2016) FGVP XIV and Luebert et al. (2016) A 20-35, in 3 irregular series • branch lengths deliberate, not expressing actual time scale Hoplestigmataceae fr drupe, endocarp undivided, 4-seeded • the characters listed do not necessarily apply to all members of the respective clade W Africa Hoplestigma 2 • position of various characters on the tree uncertain • Ehretiaceae: topology still generally uncertain and under major reinvestigation (M. Gottschling, pers. comm.) • Hilsenbergia included in Bourreria procumbent herbaceous annuals • Nama in Namaceae polyphyletic adventitious roots + • Tournefortia and Argusia included in Heliotropium Coldeniaceae flw tetramerous trop. Asia ? (widespread throughout Tropics) 1 * Hydrophyllaceae: unpublished topology according to G. Walden (pers. comm.) Coldenia ** Turricula uncertain according to G. Walden (pers. comm.) stigmatic branches 4 References endocarp undivided Baldwin et al. (2012) The Jepson Manual, 2nd edn. Ferguson (1998) Syst. Bot. 23(3): 253-268 Cordiaceae usually enclosed in inflated calyx, partly winged Gottschling et al. (2014) Mol. Phylogen. Evol. 72: 1-6 cotyledons plicate Cordia (incl. Varronia) c. 350 Hilger and Diane (2003) Bot. Jahrb. Syst. 125(1): 19-51 Kubitzki, ed. (2016) FGVP XIV, Springer Luebert et al. (2016) Taxon 65(3): 502-522 Walden and Patterson (2012) Madroño 59(4): 211-222 Walden (2015) UC Berkeley Electronic Theses and Dissertations, http://escholarship.org/uc/item/8tq9z0kb

see also: Stevens PF (2018) APweb – www.mobot.org/MOBOT/research/APweb * Thanks to Marc Gottschling and Genevieve Walden for valuable comments

Boraginaceae Angiosperm Phylogeny Phylogeny Poster Poster © The Authors, 2018 (CC-BY) © The