A Review on Phytochemical Analysis and Ethnobotanical Uses of Haloxylon Stocksii
Total Page:16
File Type:pdf, Size:1020Kb
Load more
Recommended publications
-
Seed Ecology Iii
SEED ECOLOGY III The Third International Society for Seed Science Meeting on Seeds and the Environment “Seeds and Change” Conference Proceedings June 20 to June 24, 2010 Salt Lake City, Utah, USA Editors: R. Pendleton, S. Meyer, B. Schultz Proceedings of the Seed Ecology III Conference Preface Extended abstracts included in this proceedings will be made available online. Enquiries and requests for hardcopies of this volume should be sent to: Dr. Rosemary Pendleton USFS Rocky Mountain Research Station Albuquerque Forestry Sciences Laboratory 333 Broadway SE Suite 115 Albuquerque, New Mexico, USA 87102-3497 The extended abstracts in this proceedings were edited for clarity. Seed Ecology III logo designed by Bitsy Schultz. i June 2010, Salt Lake City, Utah Proceedings of the Seed Ecology III Conference Table of Contents Germination Ecology of Dry Sandy Grassland Species along a pH-Gradient Simulated by Different Aluminium Concentrations.....................................................................................................................1 M Abedi, M Bartelheimer, Ralph Krall and Peter Poschlod Induction and Release of Secondary Dormancy under Field Conditions in Bromus tectorum.......................2 PS Allen, SE Meyer, and K Foote Seedling Production for Purposes of Biodiversity Restoration in the Brazilian Cerrado Region Can Be Greatly Enhanced by Seed Pretreatments Derived from Seed Technology......................................................4 S Anese, GCM Soares, ACB Matos, DAB Pinto, EAA da Silva, and HWM Hilhorst -
Genetic Diversity and Population Structure of Haloxylon Salicornicum Moq
RESEARCH ARTICLE Genetic diversity and population structure of Haloxylon salicornicum moq. in Kuwait by ISSR markers 1 1 1 1 2 Fadila Al SalameenID *, Nazima HabibiID , Vinod Kumar , Sami Al Amad , Jamal Dashti , Lina Talebi3, Bashayer Al Doaij1 1 Biotechnology Program, Environment and Life Sciences Research Centre, Kuwait Institute for Scientific Research, Shuwaikh, Kuwait, 2 Desert, Agriculture & Ecosystems Program, Environment and Life Sciences a1111111111 Research Centre, Kuwait Institute for Scientific Research, Shuwaikh, Kuwait, 3 Environment Pollution and a1111111111 Climate Change Program, Environment and Life Sciences Research Centre, Kuwait Institute for Scientific a1111111111 Research, Shuwaikh, Kuwait a1111111111 * [email protected] a1111111111 Abstract OPEN ACCESS Haloxylon salicornicum moq. Bunge ex Boiss (Rimth) is one of the native plants of Kuwait, extensively depleting through the anthropogenic activities. It is important to conserve Halox- Citation: Al Salameen F, Habibi N, Kumar V, Al Amad S, Dashti J, Talebi L, et al. (2018) Genetic ylon community in Kuwait as it can tolerate extreme adverse conditions of drought and salin- diversity and population structure of Haloxylon ity to be potentially used in the desert and urban revegetation and greenery national salicornicum moq. in Kuwait by ISSR markers. programs. Therefore, a set of 16 inter simple sequence repeat (ISSR) markers were used to PLoS ONE 13(11): e0207369. https://doi.org/ 10.1371/journal.pone.0207369 assess genetic diversity and population structure of 108 genotypes from six locations in Kuwait. The ISSR primers produced 195 unambiguous and reproducible bands out of which Editor: Ruslan Kalendar, University of Helsinki, FINLAND 167 bands were polymorphic (86.5%) with a mean PIC value of 0.31. -
1276-1284, 2008 © 2008, Insinet Publication
Journal of Applied Sciences Research, 4(10): 1276-1284, 2008 © 2008, INSInet Publication Assessment of Selected Species along Al-Alamein-Alexandria International Desert Road, Egypt A.A. Morsy, A.M. Youssef, H.A.M. Mosallam and A.M. Hashem Department of Botany, Faculty of Science, Ain Shams Univ., Cairo, Egypt Abstract: The present study aim at identifying the ecophysiological behavior of ten xerophytic species in relation to different habitat and stress conditions during winter and summer along Al-Alamein-Alexandria international desert road. Physical and chemical changes in soil along the different examined sites were reflected in changing plant community types. The selected plant species comprise: Haloxylon salicornicum, Anabasis articulata, Zygophyllum decumbens, Agathophora alopecroids, Cornulaca monocantha, Artemisia monosperma, Echinops spinosus, Thymelia hirsuta, Deverra triradiata and Noaea mucronata. Mechanical analysis revealed that soil associated with the studied plants is consisted of different fractions; coarse and fine sand being the dominant fractions for all samples. The contents of succulence, photosynthetic pigments, carbohydrates, nitrogenous compounds, proline and minerals were analyzed and the results showed that, although the studied species belonging to one ecological group; xerophytes, there were wide difference in their metabolism, indicating the wide range of adjustment mechanism behaved by such plants under comparable habitat conditions. In contrast to organic solutes, inorganic ions (in some of the selected -
On the Winter and Spring Aspect of the Macrolepidoptera Fauna of Jordan with Remarks on the Biology of Some Species
© Münchner Ent. Ges., download www.biologiezentrum.at Mitt. Münch. Ent. Ges. 102 65-93 München, 15.10.2012 ISSN 0340-4943 On the winter and spring aspect of the Macrolepidoptera fauna of Jordan with remarks on the biology of some species (Lepidoptera: Tineoidea, Cossoidea, Bombycoidea, Papilionoidea, Geometroidea, Noctuoidea) Dirk STADIE and Lutz LEHMANN († 1) Abstract The results of two lepidopterological expeditions to Jordan from December 20 to December 30, 2008 (provinces of Irbid, Karak, Tafila und Ma’an) and March 19 to March 25, 2009 (provinces: Zarqa, Tafila, Irbid and Balqa) are presented. A total of 166 species of Macrolepidoptera could be recorded, among them 21 species new for Jordan. Desertobia heloxylonia lawrencei subsp. n. is described. The female genitalia of Odontopera jordanaria (STAUDINGER, 1898) comb. n., are figured for the first time. The larvae of Odontopera jordanaria STAUDINGER, 1898, Minucia wiskotti (PÜNGELER, 1902), Allophyes benedictina (STAUDINGER, 1892), Agrochola pauli (STAUDINGER, 1892), Agrochola scabra (STAUDINGER, 1892), Agrochola staudingeri RONKAY, 1984, Conistra acutula (STAUDINGER, 1892), Polymixis apora (STAUDINGER, 1898) and Polymixis juditha (STAUDINGER, 1898) are described and figured. Introduction Our knowledge of the winter aspect of the macrolepidoptera fauna of the southwestern Palaearctic region is still lower compared to other seasons. It has been shown during expeditions that in countries with subtropical- mediterranean climate an interesting macrolepidoptera fauna with numerous poorly known or even new species could be ascertained, in the eremic as well as in the steppe and mediterranean zone. The focus of this communication is on the Noctuidae family, because Noctuid species dominate the species composition in winter (e.g. -
A Synopsis of the Family Chenopodiaceae in India
Pleione 6(2): 273 - 297. 2012. ISSN: 0973-9467 © East Himalayan Society for Spermatophyte Taxonomy A synopsis of the Family Chenopodiaceae in India T. K. Paul Botanical Survey of India, Central National Herbarium, Howrah-711103, India E- mail: [email protected] Received revised 07.12.2012; Accepted 11.12.2012 Abstract The present paper presents a concise account of Chenopodiaceae in India. In all 19 genera with 50 species, 1 subspecies, 3 varieties have been recognized and another 2 genera and 14 species are cultivated or introduced. The genera and species are arranged in alphabetical order. Within the enumeration Key to genera and species, correct nomenclature, reference to type materials wherever available, phenology and distribution also have been added. Key words: India, Chenopodiaceae, Synopsis, comb. et stat. nov. INTRODUCTION The plants of Chenopodiaceae Ventenat, commonly known as ‘Goosefoot’ family, are mostly grow as weed and some are food plants like spinach, chard, beets, sugar beet and quinoa. The family is placed in the order Caryophyllales by Cronquist (1981), Takhtajan (1969) and Dahlgren (1975). Hutchinson (1959) and Thorne (1968, 1992) included the family in the order Chenopodiales, Ulbrich in Engler & Prantl (1934) in the order Centrospermae and Bentham & Hooker (1880) in the series Curvembryeae. Bentham & Hooker (1880) divided the family into two series, cyclobeae and spirolobeae. Cyclobeae is characterized by annular embryo, albumen copious whereas in spirolobeae the embryo is spiral and albumen scanty or absent. Williams & Ford-Lloyd (1974) recognised three subfamilies: Chenopodieae (embryo cyclical, operculum absent, endosperm absent, ovary superior), Salsoleae (embryo spiral, operculum absent, endosperm absent, ovary superior), Beteae (embryo cyclical, operculum present in fruit, endosperm present, ovary semi-inferior). -
WOOD ANATOMY of CHENOPODIACEAE (AMARANTHACEAE S
IAWA Journal, Vol. 33 (2), 2012: 205–232 WOOD ANATOMY OF CHENOPODIACEAE (AMARANTHACEAE s. l.) Heike Heklau1, Peter Gasson2, Fritz Schweingruber3 and Pieter Baas4 SUMMARY The wood anatomy of the Chenopodiaceae is distinctive and fairly uni- form. The secondary xylem is characterised by relatively narrow vessels (<100 µm) with mostly minute pits (<4 µm), and extremely narrow ves- sels (<10 µm intergrading with vascular tracheids in addition to “normal” vessels), short vessel elements (<270 µm), successive cambia, included phloem, thick-walled or very thick-walled fibres, which are short (<470 µm), and abundant calcium oxalate crystals. Rays are mainly observed in the tribes Atripliceae, Beteae, Camphorosmeae, Chenopodieae, Hab- litzieae and Salsoleae, while many Chenopodiaceae are rayless. The Chenopodiaceae differ from the more tropical and subtropical Amaran- thaceae s.str. especially in their shorter libriform fibres and narrower vessels. Contrary to the accepted view that the subfamily Polycnemoideae lacks anomalous thickening, we found irregular successive cambia and included phloem. They are limited to long-lived roots and stem borne roots of perennials (Nitrophila mohavensis) and to a hemicryptophyte (Polycnemum fontanesii). The Chenopodiaceae often grow in extreme habitats, and this is reflected by their wood anatomy. Among the annual species, halophytes have narrower vessels than xeric species of steppes and prairies, and than species of nitrophile ruderal sites. Key words: Chenopodiaceae, Amaranthaceae s.l., included phloem, suc- cessive cambia, anomalous secondary thickening, vessel diameter, vessel element length, ecological adaptations, xerophytes, halophytes. INTRODUCTION The Chenopodiaceae in the order Caryophyllales include annual or perennial herbs, sub- shrubs, shrubs, small trees (Haloxylon ammodendron, Suaeda monoica) and climbers (Hablitzia, Holmbergia). -
The C4 Plant Lineages of Planet Earth
Journal of Experimental Botany, Vol. 62, No. 9, pp. 3155–3169, 2011 doi:10.1093/jxb/err048 Advance Access publication 16 March, 2011 REVIEW PAPER The C4 plant lineages of planet Earth Rowan F. Sage1,*, Pascal-Antoine Christin2 and Erika J. Edwards2 1 Department of Ecology and Evolutionary Biology, The University of Toronto, 25 Willcocks Street, Toronto, Ontario M5S3B2 Canada 2 Department of Ecology and Evolutionary Biology, Brown University, 80 Waterman St., Providence, RI 02912, USA * To whom correspondence should be addressed. E-mail: [email protected] Received 30 November 2010; Revised 1 February 2011; Accepted 2 February 2011 Abstract Using isotopic screens, phylogenetic assessments, and 45 years of physiological data, it is now possible to identify most of the evolutionary lineages expressing the C4 photosynthetic pathway. Here, 62 recognizable lineages of C4 photosynthesis are listed. Thirty-six lineages (60%) occur in the eudicots. Monocots account for 26 lineages, with a Downloaded from minimum of 18 lineages being present in the grass family and six in the sedge family. Species exhibiting the C3–C4 intermediate type of photosynthesis correspond to 21 lineages. Of these, 9 are not immediately associated with any C4 lineage, indicating that they did not share common C3–C4 ancestors with C4 species and are instead an independent line. The geographic centre of origin for 47 of the lineages could be estimated. These centres tend to jxb.oxfordjournals.org cluster in areas corresponding to what are now arid to semi-arid regions of southwestern North America, south- central South America, central Asia, northeastern and southern Africa, and inland Australia. -
Terrestrial Environment of Abu Dhabi Emirate, United Arab Emirates
of Abu Dhabi Emirate, United Arab Emirates TERRESTRIAL ENVIRONMENTS OF ABU DHABI EMIRATE, UNITED ARAB EMIRATES of Abu Dhabi Emirate, United Arab Emirates TERRESTRIAL ENVIRONMENTS OF ABU DHABI EMIRATE, UNITED ARAB EMIRATES Page . IV TERRESTRIAL ENVIRONMENTS OF ABU DHABI EMIRATE, UNITED ARAB EMIRATES H. H. Sheikh Khalifa bin Zayed Al Nahyan President of the United Arab Emirates Page . V TERRESTRIAL ENVIRONMENTS OF ABU DHABI EMIRATE, UNITED ARAB EMIRATES Page . VI TERRESTRIAL ENVIRONMENTS OF ABU DHABI EMIRATE, UNITED ARAB EMIRATES H. H. Sheikh Mohammed bin Zayed Al Nahyan Crown Prince of Abu Dhabi, Deputy Supreme Commander of the UAE Armed Forces Page . VII TERRESTRIAL ENVIRONMENTS OF ABU DHABI EMIRATE, UNITED ARAB EMIRATES Page . VIII TERRESTRIAL ENVIRONMENTS OF ABU DHABI EMIRATE, UNITED ARAB EMIRATES H. H. Sheikh Hamdan bin Zayed Al Nahyan Deputy Prime Minister, Chairman, EAD Page . IX TERRESTRIAL ENVIRONMENTS OF ABU DHABI EMIRATE, UNITED ARAB EMIRATES · اﻟﺒﻴﺎﻧﺎت · ﺑﺸﻜﻞ ﻋﺎم، ﺗﻢ إﻋﺪاد اﻷوراق اﻟﻘﻄﺎﻋﻴﺔ اﻷﺻﻠﻴﺔ ﺑﺸﻜﻞ ﺟﺪﻳﺪ ﻗﺪم ﻓﻴﻬﺎ · اﻷدوات واﻷﺳﺎﻟﻴﺐ ﻣﺠﻤﻮﻋﺔ ﻗﻴﻤﺔ ﻣﻦ اﳌﻌﻠﻮﻣﺎت · اﻟﺘﻮﻋﻴﺔ · مل ﺗﺼﻞ ﻣﺸﺎرﻛﺔ اﻟﴩﻛﺎء واﻟﺠﻬﺎت اﳌﻌﻨﻴني إﱃ اﻟﺤﺪ اﳌﺨﻄﻂ ﻟﻪ · ﺑﻨﺎء اﻟﻘﺪرات · ﺗﻢ أﻋﺪاد اﻷوراق اﻟﻘﻄﺎﻋﻴﺔ ﺑﺪون دﻋﻢ ﻛﺎﰲ ﻣﻦ اﻟﻬﻴﺌﺔ أو اﻟﴩﻛﺎء واﻟﺠﻬﺎت ·اﻟﺴﻴﺎﺳﺔ اﳌﻌﻨﻴني، وﺑﺎﻟﺘﺎﱄ، ﻛﺎن ﻋﲆ ﻣﺆﻟﻒ اﻟﻮرﻗﺔ اﻟﻘﻄﺎﻋﻴﺔ ﺗﺤﻤﻞ ﻋﺐء إﻋﺪاد ورﻗﺔ اﻷوراق اﻟﻘﻄﺎﻋﻴﺔ ﻫﺬا اﻟﻘﻄﺎع ﰲ وﻗﺖ زﻣﻨﻲ ﻣﺤﺪود ﻧﻮﻋﺎ ﻣﺎ · ﰲ ﺑﻌﺾ اﻟﺤﺎﻻت ﻛﺎﻧﺖ اﻟﺒﻴﺎﻧﺎت اﳌﺴﺘﺨﺪﻣﺔ ﻗﺪميﺔ ﻧﺴﺒﻴﺎ ﺧﻼل اﻟﺴﻨﻮات اﳌﺎﺿﻴﺔ ﻗﺎﻣﺖ ﻣﺨﺘﻠﻒ اﻟﻘﻄﺎﻋﺎت اﳌﻌﻨﻴﺔ ﺑﺸﺆون اﻟﺒﻴﺌﺔ ﺑﺘﺠﻤﻴﻊ ﻛﻢ ﻣﻦ اﳌﻌﻠﻮﻣﺎت اﳌﺘﻨﻮﻋﺔ ﺑﻌﺪة ﺻﻮر ﺗﺼﻒ ﻣﺎ ﻫﻮ ﻣﻌﺮوف ﻋﻦ اﻟﺒﻴﺌﺔ ﰲ إﻣﺎرة · مل ﻳﺘﻢ إﺿﻔﺎء اﻟﻄﺎﺑﻊ اﳌﺆﺳﴘ ﻋﲆ ﻋﻤﻠﻴﺔ ﺟﻤﻊ اﻟﺒﻴﺎﻧﺎت وﺗﺒﺎدﻟﻬﺎ أﺑﻮﻇﺒﻲ ودوﻟﺔ اﻹﻣﺎرات اﻟﻌﺮﺑﻴﺔ اﳌﺘﺤﺪة واﻟﺨﻠﻴﺞ اﻟﻌﺮيب. -
AL SOQEER, A. – ABDALLA, W. E. : Epidermal Micro-Morphological Study on Stems of Members
El Ghazali et al.: epidermal micro-morphology of chenopodiaceae - 623 - EPIDERMAL MICRO-MORPHOLOGICAL STUDY ON STEMS OF MEMBERS OF THE FAMILY CHENOPODIACEAE EL GHAZALI, G. E. B.1* – AL SOQEER, A.2 – ABDALLA, W. E.1 1Faculty of Science and Arts at Al Rass, Qassim University, Saudi Arabia (fax: +9661633351600) 2Faculty of Agriculture and Veterinary Medicine, Qassim University, Saudi Arabia *Corresponding author e-mail: [email protected] (phone+966502492660) (Received 25th May 2016; accepted 25th Aug 2016) Abstract. Stems of 14 species belonging to 10 genera of the family Chenopodiaceae were examined under scanning electron microscope (SEM). Micro-morphological characters such as sculpture of epicuticular wax, trichomes and surface relief were investigated. These characters were clumped into five groups. These groups do not coincide solely with the five tribes of the species studied. The study presented epicuticular wax sculptures and surface reliefs not previously reported for the family Chenopodiaceae. Keywords: scanning electron microscope, epicuticular wax sculpture, trichome, surface relief, semi-arid Introduction Epidermal characters as revealed by scanning electron microscopes (SEM) exhibit a variety of micro-morphological features (Barthlott, 1990), influenced by the environment (Sattler and Rutishauser, 1997; Koch and Ensikat, 2008), and act as a barrier against mechanical damages, insects, excessive light and loss of water (Bird and Gray, 2003; Rudall, 2007; Mauseth, 2008). These epidermal characters can reflect the relationships between habitat and phytogenetics of the plants (Liu, 2006). Chenopodiaceae Vent., commonly known as Goosefoot family, is cosmopolitan, especially diverse in arid, semi-arid, saline and hypersaline ecosystems (Kuhn et al., 1993; Hedge et al., 1997). Members of the family Chenopodiaceae are represented in the flora of Saudi Arabia by 74 species belonging to 23 genera which grow in many regions (Chaudhary, 1999), with Qassim Region accommodating a considerable number (Al-Turki, 1997). -
Conservation Condition of Haloxylon Salicornicum (Moq.) Bunge Ex Boiss
Int.J.Curr.Microbiol.App.Sci (2014) 3(10) 310-325 ISSN: 2319-7706 Volume 3 Number 10 (2014) pp. 310-325 http://www.ijcmas.com Original Research Article Conservation condition of Haloxylon salicornicum (Moq.) Bunge ex Boiss. in degraded desert habitats of northern Kuwait Raafat Hassan ABD El-WAHAB1,2*, Ahmad Rashed Al-RASHED2, and Yousef Al-HAMAD2 1Botany Department, Faculty of Science, Suez Canal University, Ismailia 41522, Egypt 2Science Department, Faculty of Basic Education, Public Authority for Applied Education and Training, Adailiya 73251, Kuwait *Corresponding author A B S T R A C T The present study aims to analyse the effects of more than 10 years of in situ conservation on vegetation structure and soil conditions of desert habitats dominated by Haloxylon salicornicum. in northeast of Kuwait. Size measurements of Haloxylon shrubs and sediments of nabkas were carried out inside and outside K e y w o r d s Sabah Al-Ahmad Natural Reserve. Edaphic conditions of nabkas and interspaces were analysed. Analysis of variance and t test show significant variations of nabka Conservation, vegetation structures and most of soil conditions between disturbed and non- Human disturbed sites. Differences of soil conditions between nabkas and interspaces are impacts, significant in non-disturbed sites. Species richness, total plant cover, and size Land features of Haloxylon shrubs and nabkas in non-disturbed sites are higher and more degradation, developed than in disturbed sites. Size structure analysis indicates that Haloxylon Plant salicornicum is suffering population decline and should consider as vulnerable diversity, species in Kuwait. Human impacts are the main threats affecting the health and Nabka abundance of nabkas and causing land degradation and species loss. -
(Haloxylon Persicum Bunge) in the Arabian Peninsula
AL-KHALIFAH & SHANAVASKHAN 51 Tropical Ecology 48 (1): 51-60, 2007 ISSN 0564-3295 © International Society for Tropical Ecology www.tropecol.com On the distribution, status and phenology of Ghada ( Haloxylon persicum Bunge) in the Arabian Peninsula NASSER S. AL-KHALIFAH* & A.E. SHANAVASKHAN *Natural Resources and Environmental ResearchInstitute, King Abdulaziz City for Science and Technology, Post Box No.6086, Riyadh 11442, Kingdom of Saudi Arabia Abstract: The distribution and status of Haloxylon persicum Bunge ('Ghada') have been evaluated for the Arabian Peninsula. Analysis of the data using IUCN guidelines showed that the taxon qualifies for the IUCN status 'lower risk' but 'conservation dependent'. Its geographical distribution is from nearer or above the Tropic of Cancer, up to 50 o N latitude. Phenological studies of the natural populations revealed vernal and aestival flowering. Vernal flowers produced viable seeds but aestival flowers were mostly male or very rarely polygamous. Seed germination studies were conducted to probe in to the poor regeneration in natural populations. Results indicated that deep burial of seed and the non-availability of adequate moisture in the germination medium are the main limiting factors for the population expansion. Ghada is a C 4 plant showing unusual behaviour of producing C 3 seedlings. These C 3 seedlings produced during winter help the taxon to perpetuate, even in adverse conditions of the desert. Resumen: La distribución y el estatus de Haloxylon persicum Bunge ('ghada') fueron evaluados en la Península Arábiga. El análisis de los datos usando los lineamientos de la IUCN mostró que este taxón merece el estatus de la IUCN de ‘bajo riesgo’ pero ‘dependiente de medidas de conservación’. -
From Cacti to Carnivores: Improved Phylotranscriptomic Sampling And
Article Type: Special Issue Article RESEARCH ARTICLE INVITED SPECIAL ARTICLE For the Special Issue: Using and Navigating the Plant Tree of Life Short Title: Walker et al.—Phylotranscriptomic analysis of Caryophyllales From cacti to carnivores: Improved phylotranscriptomic sampling and hierarchical homology inference provide further insight into the evolution of Caryophyllales Joseph F. Walker1,13, Ya Yang2, Tao Feng3, Alfonso Timoneda3, Jessica Mikenas4,5, Vera Hutchison4, Caroline Edwards4, Ning Wang1, Sonia Ahluwalia1, Julia Olivieri4,6, Nathanael Walker-Hale7, Lucas C. Majure8, Raúl Puente8, Gudrun Kadereit9,10, Maximilian Lauterbach9,10, Urs Eggli11, Hilda Flores-Olvera12, Helga Ochoterena12, Samuel F. Brockington3, Michael J. Moore,4 and Stephen A. Smith1,13 Manuscript received 13 October 2017; revision accepted 4 January 2018. 1 Department of Ecology & Evolutionary Biology, University of Michigan, 830 North University Avenue, Ann Arbor, MI 48109-1048 USA 2 Department of Plant and Microbial Biology, University of Minnesota-Twin Cities, 1445 Gortner Avenue, St. Paul, MN 55108 USA 3 Department of Plant Sciences, University of Cambridge, Cambridge CB2 3EA, UK 4 Department of Biology, Oberlin College, Science Center K111, 119 Woodland Street, Oberlin, OH 44074-1097 USA 5 Current address: USGS Canyonlands Research Station, Southwest Biological Science Center, 2290 S West Resource Blvd, Moab, UT 84532 USA 6 Institute of Computational and Mathematical Engineering (ICME), Stanford University, 475 Author Manuscript Via Ortega, Suite B060, Stanford, CA, 94305-4042 USA This is the author manuscript accepted for publication and has undergone full peer review but has not been through the copyediting, typesetting, pagination and proofreading process, which may lead to differences between this version and the Version of Record.