CATRINA (2008), 3 (1): 1 -10 PROCEEDINGS OF THE SECOND INTERNATIONAL CONFERENCEON THE ROLEOF GENETICSAND © 2008BY THE EGYPTIAN SOCIETYFOR ENVIRONMENTAL SCIENCES BIOTECHNOLOGY IN CONSERVATIONOF NATURAL RESOURCES , ISMAILIA , EGYPT , JULY 9-10, 2007 Systematic Reviewofthe Papaveracea Adans.andStatusofSomeGenera Wafaa K. Taia Alexandria University, Faculty of Science, Botany Department, Alexandria-Egypt ABSTRACT This work deals with the systematic relations of family Papaveraceae according to the new data derivedfrom molecular analysis. Anoverall view has beengiventoelucidate the taxonomic positionof the family withinthe basal eudicots. Characteristic features of all the group has beengivenwithclear view of the general features of the family andits taxonomic divisions. Complete list of the genera, as listedby Royal Botanical Gardens inKew has beenaddedwithsummary of the recent phylogenetic relationships of the family and its genera. Leaf and fruit morphological variations of 14 species belonging to Papaveraceae s.s., 7 species belonging to Fumariaceae s.s. and two species from Hypecoaceae s.s. have been studied to assess the new classification of the Papaveraceae s.l. with complete description of their morphological variations and their position according to the new classificationof the family. Keywords: Eudicots, Magnoliopsida, Papaveraceae, Phylogeny, Ranunculales, Taxonomy. Taxonomic Status of the Papaveraceae s.l. Ranunculaceae , Papaveraceae, Fumariaceae, The status of family Papaveraceae s.l. has been faced Berberidaceae, Menispermaceae , and Aristolochiaceae . with numerous opinions. Before the extensive ribosomal The family is closely related to the Fumariaceae which and DNA sequences researches concerning the often included within it (Norris, 1941, Fohne, 1962, phylogeny of the taxonomic taxa, the family has been Kolbe, 1978, Behnke and Barthlott, 1983 Chase et al., considered by Wettstein (1935) as one of the 1993; Taia and Sheha, 2003). Meanwhile, the family Rhoeadales families. Later, Melchior (1964), Tamura has close affinity to members of Ranunculales with the (1974), Benson (1979) and Tutin et al. (1993) renamed only differences that gynoecium is paracarpous and the the order Rhoeadales by Papaverales s.l. with four presence of secretory idioblasts or laticifers in the suborders; Papaverinae, Capparinae, Tovarianae and Papaveraceae s.l., and both families are considered as Moringanae . The family was found to be closely related primitive families. Parker (1982) separated the to the Brassicaceae , Capparidaceae , Resedaceae , Papaveraceae as an order Papaverales while the Tovariaceae and Moringaceae (Blagowestschenski, Ranunculaceae as order Ranunculales and both orders 1955, Hegnauer, 1961 and Gershenzon and Mabry, under subclass Magnoliidae , class Magnoliopsida . This 1983). The position of these families in the same order class refers to a small group containing Papaveraceae, is due to flower characters, such as regular and Ranunculaceae and Berberidaceae as sister group to hypogynous flowers, anthers arranged in several whorls Monocots. Based on the recent work on molecular and the carpels are syncarpous and possess two to many analysis, done by Hoot et al. (1999), the Papaveraceae parietal ovules. Norris (1941) considered the s.l. is considered as one of the families belonging to Papaveraceae s.l. closely related to both the order Ranunculales and they up graded the order Fumariaceae and Brassicaceae and they all evolved Ranunculales from Magnoliidae to the base of the from the same origin according to torus anatomy and Eudicots, which is in close association with the nectary characteristics. Monocots. Family Papaveraceae Adans. nom.cons. s.l . which is a north temperate, mostly herbaceous family consisting Phylogenetic Opinions within the Family of 23 genera and about 250 species (Heywood, 1993), Cronquist (1981) considered the Ranunculales as one Newly recorded species raises the number of genera to of the woody magnoliids because their flowers have free 35 as compared with the list given by Royal Botanical parts that are sometimes spirally arranged. Cronquist Gardens, in Kew, 2006. The family has three synonyms; (1981) suggested that the connection to the woody Chelidoniaceae Martinov., Echscholziaceae Ser. and magnoliids was via Illiciaceae (ANITA grade; Qui et Platystemonaceae (Rchb.ex Spatch) Lilja. According to al. , 1999) due to the presence of triaperturate pollen Cronquist System (1981), family Papaveraceae s.l. is grains in both groups. Spichiger and Savolainen (1997) classified under subclass Magnoliidae together with the pointed to the similarities between the Ranunculales and Magnoliaceae , Nymphaeaceae and Ranunculaceae . Papaverales with the monocots as they share many This subclass is characterized by well developed features such as imperfect vessels; inaperturate or flowers with separated perianth or calyx and corolla, the uniaperturate pollen grains (or derived types). Recent stamens are numerous and gynoecium is apocarpous. In works based on molecular analysis considered the Papaveraceae gynoecium is paracarpous. According to Ranunculales as a monophyletic group (Fig. 1) (Chase South West Virginia flora, the Magnoliideae comprises et al. , 1993, Drinnan et al., 1994, Hoot and Crane 1995, nine families; Magnoliaceae , Annonaceae, Lauraceae, Soltis et al. , 1997, Kallersjo et al. , 1998, ____________________________________________ * Corresponding author: [email protected] Systematic review of the Papaveracea and status of some genera General Features of the Family The family has plants in different varieties of life forms and morphological characters. They are herbs or sub-shrubs, shrubs, or even small trees ( Dendromegon rigida) , annuals, biennials, or perennials. Plants have tap roots or rhizomes with leafy or naked erect or spreading stems. Leaves are basal and/or cauline, alternate to opposite or whorled, simple ex-stipulate petiolate or sessile with entire blade or lobed in pinnate, subpalmate, or palmate orders of lobes. In Argemone the leaves are dissected with spiny margins. The leaves are mostly glabrous, except few species covered with ramified or glandular hairs with smooth walls. Stomata are either diacytic or anomocytic with isodiametric or elongated epidermal cells (Heywood, 1993). The family has variety of flower arrangements, forms and colors, but all are bracteate, radially symmetric, pedicellate or sessile; receptacle sometimes expanded and forming cup or ring beneath calyx (in Eschscholzia, Figure (1) : Phylogeny of the Ranumculales , showingrelative Meconella and Platystemon) . The flowers are either positions of the major families within this order (Tree solitary or arranged in groups which are either terminal adapted from Hoot et al., 1999). Numbers above lines or axillary. The inflorescences are either cymose or indicate the number of nucleotide changes supportingeach racemose, umbelliform or corymbiform. Flowers are branch. Number below the branches is the percentage of sometimes perigynous; sepals two or three, ob-ovate, times that the branch was recovered in 1000 bootstrap distinct or connate always caducous. Petals are distinct, replications. colored and ob-ovate, usually four or more, sometimes absent. Stamens are numerous in many whorls, Hoot et al. , 1999). Hilu et al. (2003), Kim et al. (2004) sometimes 4-15 in Meconella and Canbya , with bi- and Worberg et al. (2006) supported the close locular anthers (Kiger, 1996). Pollen grains appear association of the Papaveraceae s.l. with the spheroidal or sub prolate, medium sized, usually with Fumariaceae then with the Eupteleaceae and the three tricolpate apertures. In some species such as Papaver families are in close relation with the rest of the argemone, Argemone mexicana, and Roemeria hybrida , Ranunculales families. pollen grains are polyporate (Taia and Sheha, 2003). Despite the relations between the Papaveraceae s.l. Pistil is 1, 2 to many (22) united carpels with one or genera are still obscure, due to the lack of fossils two locules, sometimes multilocular by placental records for that group and their geographical intrusion, placenta two or more in parietal position. distribution. Kadereit et al. (1994) found that the genera Style is usually one or absent with sessile stigma. Papaver L., Meconopsis Vig., Stylomecon G.Taylor and Stigma lobes are 2 to many in circular disc or radiating Roemeria Medik. Belonging to Papaveraceae s.str. ones. Fruits are capsules dehiscent by pores, valves or subfamily Papaveroideae are morphologically similar. dissociating and breaking transversely into one seeded In the mean time Schwarzbach & Kadereit (1995) segments (only in Platystemon ). Seeds are always many reported that these genera form a monophyletic clade on small, sometimes arillate or carunculate with different the basis of molecular analysis. Kadereit et al. (1997), colors varies from white to black or different shades of on the basis of molecular data concluded that (1) Asian brown. Meconopsis is not a monophyletic group, but paraphyletic in relation to Papaver s.l. including Classification of the Family M.cambrica , Stylomecon and Roemeria ; (2) both The Papaveraceae s.l. is subdivided into three Roemeria and Stylomecon are nested with Papaver s.l., subfamilies by Hoot et al., 1997; Pteridophylloideae, Roemeria is sister group to Papaver sect. Argemonidium Papaveroideae , and Fumarioideae. Ernst (1962), Layka and Stylomecon to papaver californicum ; and (3) the (1976), Heslop-Harison and Shivana