5/14/17 Week 7; Monday Announcements: Quiz in lab today; Exam in lecture on Friday (review Wed 5:30 pm HCK 320) 3rd and final extra credit exercise. Lecture: Asteridae s.l. Continued Lamiales This group includes members of 2 or more orders in various traditional classifications (e.g., Cronquist - overhead), including the much more narrowly defined Lamiales s.s. and the Scrophulariales. Altogether, there are about 20 family-level clades and ca. 22,000 species. Oleaceae (lab-only family) – Oleaceae has flowers with radial symmetry and is unusual within Lamiales for this fact. However, they diverged within Lamiales very early and help provide evidence that the earliest plants in Lamiales evolution were radially symmetric. Bilateral symmetry, which characterizes most of the Lamiales, is evolutionarily derived within the group. Lamiaceae (Labiatae) - Mint family (258 gen/6970 spp) 1) woody/herbaceous 2) often aromatic – the large subfamily Nepetoideae characterized by aromatic oils instead of iridoids. These include many culinary herbs and ingredients in perfumes, including sage, oregano, marjoram, rosemary, thyme, basil, lavender, etc. 3) stems often square (traditional herbaceous mints) – demo: Lamium album 4) leaves usually opposite and decussate; sometimes whorled; no stipules 5) Flowers - zygomorphic; bilabiate (usually 2 upper lobes/3 lower lobes) - stamens and stigma usually enclosed in upper lip of corolla - stamens 4 (or 2); often didynamous (2 uneven pairs) - gynoecium 2 carpels, each with 2 ovules and a false septum between ovules of each carpel; often with a gynobasic style arising from between 4 mericarps - fruit: schizocarp forming 4 nutlets or a 1-4 seeded drupe Floral formula: X 5, 5, 2 or 4, 2 nutlets or drupe [carpels fused] Until very recently, the Lamiaceae was considered to be an easily defined family (although experts acknowledged some fuzzy distinctions relative to the Verbenaceae), hence the pre-Linnaean name Labiatae. Studies by Cantino, Wagstaff and Olmstead, now indicate that most of the old-world members of the Verbenaceae are intimately related to the Lamiaceae. Only a small part of the traditional Verbenaceae (fortunately, including the type genus Verbena) are now recognized as Verbenaceae. (Overhead) Page 59 5/14/17 “Scrophulariaceae” figwort family (269 gen/5100 spp, but….) (Scrophulariaceae s.s. 54/1800) 1) herbs or shrubs (ours usually herbs) (rarely with square stems, including Scrophularia!) 2) leaves usually alternate (some opposite); no stipules 3) many parasitic species – now segregated into family Orobanchaceae - hemiparasites - still capable of photosynthesis - holoparasites - without chlorophyll and completely dependent on their host 4) Flowers - zygomorphic; often bilabiate - didynamous stamens (2-long; 2-short); occasionally reduced to 2 - fruit a many-seeded capsule (occasionally a berry) Floral formula: X 5, 5, 2 or 4, 2 capsule [carpels fused] Review stamen arrangements: tetradynamous - 4 long +2 short - Brassicaceae monadelphous - filaments fused into a tube; free at end - Malvaceae diadelphous - 9 fused at the filaments, one free - Fabaceae (Papilionoideae) didynamous - 2 long and 2 short – Lamiaceae & Scrophulariaceae This family always has been difficult to characterize by any explicit characters. It seems to have the generalized reproductive morphology of the large order Lamiales, which has ca. 20 families. Recently, research in my lab has determined that the family is polyphyletic. It is now being proposed that several families should be recognized: Scrophulariaceae sensu stricto: Scrophularia and Verbascum are north temperate in distribution, but the rest are mostly southern hemisphere from South Africa and Australia; most species have radial corolla symmetry. Orobanchaceae: the holo- and hemiparasites – world-wide, includes Castilleja and Pedicularis here in the PNW, but also is diverse in Africa, Europe, and Asia. Veronicaceae (Plantaginaceae in Judd et al. textbook): the large majority of the rest of the family, including some additional genera, such as Plantago (wind pollinated), Hippuris and Callitriche (both aquatic plants). This contains most of the North American species, including Penstemon, Veronica, Collinsia, and Plantago. Calceolariaceae: 3 southern hemisphere genera; ca. 300 spp. mostly in Calceolaria Stilbaceae: a small South African family that has been expanded by the inclusion of several scroph genera. Phrymaceae: Mostly the genus Mimulus and a few smaller genera; greatest diversity in western North America, with second area of diversity in Australia. Linderniaceae: About 13 genera and ca. 200 species, distributed in Africa, SE Asia, and North and South America. Mazaceae: 3 genera and ca. 25 species of Asia Rehmanniaceae: 2 genera and 11 species in East Asia Page 60 5/14/17 As with the Verbenaceae, where a smaller portion of the family is associated with the type genus, Verbena, Scrophularia is part of a small portion of the original family. So, most of what we recognize as “Scrophulariaceae” here in the western US, now belong to different families: Orobanchaceae, Phrymaceae, and Veronicaceae (or Plantaginaceae). (Overhead) Page 61 5/14/17 Week 7; Wednesday Announcements: Exam on Friday Review: Two families taken up last time were: Lamiaceae and Scrophulariaceae Which monophyletic? How are they similar? How are they different? Lecture: Asteridae s.l. wrap up. Campanulidae – families with inferior ovaries Caprifoliaceae Honeysuckle family (36 gen/810 spp) – “Cap” in “A MAD Cap Horse” “A” is for Adoxaceae – today, too. This family includes two herbaceous families previously kept separate: Valerianaceae and Dipsacaceae … and excludes two large genera Sambucus and Viburnum (to Adoxaceae) 1) mostly woody (some derived groups are herbaceous; eg, Valerianaceae and Dipsacaceae) 2) lvs opposite; usually simple, with no stipules 3) Flowers - zygomorphic (sometimes weakly so), typically 2 upper and 3 lower lobes, but sometimes bilabiate with 4 upper lobes/1 lower lobe (especially in Lonicera, the honeysuckles) - 2-5 fused carpels, typically with one ovule per carpel and often with one or more ovules aborted; sometimes reduced to a single seeded fruit - ovary inferior Floral formula: X 5, 5, 4-5, 2-5 berry, capsule, drupe, achene [carpels fused; ovary inferior] NOT monophyletic in its traditional circumscription (e.g., Cronquist, 1981), because it was paraphyletic by exclusion of Adoxaceae, Valerianaceae, Dipsacaceae. These families have been realigned by transferring two genera (Sambucus and Viburnum) from Caprifoliaceae to Adoxaceae and including Dipsacaceae and Valerianaceae in Caprifoliaceae (overhead) Adoxaceae (BIOL 317 required family) – includes Sambucus and Viburnum (along with 3 other small genera), two genera traditionally placed in Caprifoliaceae. Page 62 5/14/17 Apiaceae Carrot or parsley family (434 gen/3780 spp) also called Umbelliferae – ‘the umbels’ 1) herbs, including many culinary herbs/spices (dill, caroway, coriander, cumin, fennel, anise), and edible plants (carrot, celery, parsnip, parsley) 2) often aromatic with a variety of non-iridoid 2º compounds (ethereal oils, triterpenoids, sesquiterpene lactones, etc.) 3) lvs. alternate, simple or compound; with sheathing leaf bases 4) Typical inflorescence – Umbel – many flowers emerging from one point subtended by bracts Pseudanthium - Many small flowers together making up a structure that functions as one flower. 5) Flowers - 5-parted - polypetalous - 2 fused carpels, each with one ovule, often splitting apart at maturity, or splitting at the base and staying attached at the tip, schizocarp - ovary inferior Floral formula: * 5, 5, 5, 2 schizocarp [carpels fused; inferior ovary] These plants fit well into the Asteridae, except for the free petals, therefore, they were classified in the Rosidae in Cronquist and other systems However, the developmental study of Erbar shows that the small flowers result from the cessation of development at a stage corresponding to the early stage of flower development in other Asteridae, at which point the lobes of the corolla have developed, but the tube has not. Hence, polypetalous flowers. Early gamopetaly – floral tube forms first in development and then petal lobes Late gamopetaly – petal lobes form first and then connect into a tube Page 63 5/14/17 Asteraceae Sunflower family (1535 gen/23,000 spp) also called Compositae The largest family of dicots with an estimated number of species = 20,000-30,000 1) Sesquiterpene lactones as chemical defense (lacking iridoids) 2) all life forms, but mostly herbs 3) lvs usually simple, without stipules; alt or opp 4) Inflorescence (pseudanthium) a capitulum or ‘head’, surrounded by an involucre, consisting of involucral bracts (remember Fagaceae female flowers?) The alternate name for this family, Compositae, recognizes the ‘composite’ nature of the pseudanthium; each unit made up of many small flowers 5) flowers borne on a flat to conical receptacle; this often with scales or chaff between the flowers 6) Flowers - two kinds: {some have only one or the other, but most have both} Ray florets - usually female, zygomorphic, 3-lobed at tip; - A part of the family (subfamily Cichorioideae) has only ray flowers; these are called ligulate flowers (‘strap-like’) and are 5-lobed at the tip Disk florets - usually hermaphrodite, actinomorphic - calyx reduced to bristles or scales, called pappus; sometimes absent - anthers fused at margins, synantherous androecium - ovary inferior, composed of 2 fused carpels each with one ovule, one ovule