Distribution of areole areas Distribution of areole eccentricities Disentangling ecological and phylogenetic signals in leaf venation patterns Aextoxicon punctatum ( Aextoxicaceae )
NCLC #: 2932 , Phot. #: 10 D e n s i t y D e n s i t y Walton Green
In the past, however, it has proved difficult to measure the venation even of modern leaves completely, or to characterize the relationships between specific aspects of a vein network and their associated functions or phylogenetic significance. Most earlier work, therefore, has either been qualitative or focussed on single characters. We automated the following measurements of each areole
Using photographs of modern cleared leaves and image analysis software, we have succeeded in measuring a number on photographs of cleared leaves of 120 species. Sagittaria sp. of attributes of the vein network of more than 100 leaves, representing most major angiosperm groups and a few ferns. ( Alismataceae ) The measurements are repeatable and correspond to intuitive and descriptive notions of vein organization, like areole NCLC #: 797 , Phot. #: 14 D e n s i t y D e n s i t y size and shape. We have observed significant relationships between venation attributes and ecological variables like growth form. Attributes of venation are also not randomly distributed phylogenetically. This supports the contention that leaf venation patterns encode ecological and historical signals, and gives promise that multivariate quantitative data will See also: Unsatisfactory Images help disentangle them. Leaf rank in angiosperms: phylogenetic and ecological Sagittaria sp. ( Alismataceae ) variation in leaf vein organization. NCLC #: 797 , Phot. #: 15 Little, Stefan A., Green, W. , D e n s i t y D e n s i t y Hickey, L. J., Wilf, P.
Session: 67 Systematics Section Location: Wasatch B/Cliff Lodge - Level C Date: Wednesday, July 29th, 2009 Time: 2:45 PM Periomphale gaultherioides Number: 67008 ( Alseuosmiaceae ) Abstract ID:880 NCLC #: 8418 , Phot. #: 16
Area D e n s i t y D e n s i t y The allometry of leaf Convex Area Eccentricity vascular networks. Solidity Price, Charles, Green, W., Wing, S. L., Weitz, J. Perimeter Equivalent diameter Periomphale gaultherioides Session: 62 Evolutionary Developmental Major and minor axes of best fit ellipse ( Alseuosmiaceae ) Biology (Evo-Devo) Mean distance to a vein NCLC #: 8418 , Phot. #: 17 Location: Cottonwood A/Snowbird Center D e n s i t y D e n s i t y Date: Wednesday, July 29th, 2009 Time: 10:30 AM Number: 62002 Abstract ID:329 1 Satisfactory Images 2 Examples of some different images... 3 ...with their associated areole measurements. 4
black = mode; red = median; blue = mean black = mode; red = median; blue = mean black = mode; red = median; blue = mean
Pittosporum glabratum Gastonia floribunda Anacardiaceae Viburnum pauciflorum Baccharis pedunculata Archibaccharis subsessilis 4.5 5.5 6.5 7.5 0.65 0.75 0.85 0.95 Philactis zinnioides Periomphale gaultherioides Ilex dipyrena Lundia spruceana Louteridium mexicanum Fraxinus chinensis 1 0 Goetzea elegans Cordia dichotoma
Hoya camphorifolia 9 Rauvolfia balansae Motandra guineensis Alstonia congensis logArea.MODE Emmotum acuminatum 8 Gomphandra quadrifida Arctostaphylos nevadensis Pyrola acerifolia Moneses uniflora 7 Clethra mexicana Actinidia latifolia
Mastixia caudatifolia 6 Iresine interrupta Achatocarpus mexicanus
Tetrocera akara 7 . 5 Schoepfia jasminodora Vatica mindanensis Parashorea plicata Bixa excelsa Bombax aquaticum
Acer argutum 6 . 5 Rhus succedanea Nothopegia castaneaefolia Metopium brownei logPerim.MODE Heeria argentea Cotinus americanus Comocladia acuminata 5 . Buchanania sessilifolia Astronium fraxinifolium Anacardium pumilum Bursera gummifera
Jarilla heterophylla 4 . 5 Akania hillii Salix floridana Salix humilis Rinorea lanceolata Caraipa punctulata Caryocara barbinerve Ochna wallichii Gomphia angustifolia Brackenridgea nitida
Brunellia conocladifolia 0 . 9 Cunonia macrophylla Taxon sampling of 77 (in red) of 481 families in Cheiloclinium gleasonianum Eccentricity.MODE Campylostemon angolense Alnus glabrata Carya ovata Haston et al. (2007) Taxon 56(1):7–12. Quercus arizonica Amelanchier asiatica
Coussapoa villosa 0 . 8 Artocarpus kemando Celtis formosana Uraria clarksei Boxplots by habit Calliandra canescens Kibessia acuminatissima for 120 angiosperm species Axinaea macrophylla 0 . 9 5 Stachyurus praecox Austrobaileya scandens Vitis balanseana Ribes lacustre Clematis pitcheri Itea nutans Smilax acutifolia
Cercidiphyllum japonicum 0 . 8 5 Miliusa longipes Aextoxicon punctatum 1 0 Styloceras laurifolium Solidity.MODE Periomphale gaultherioides Trochodendron aralioides Platanus occidentalis Meliosma obtusifolia Styloceras laurifolium Clematis pitcheri 0 . 7 5 Platytinospora buchholzii Hypserpa neo-caledonica Euptelea francheti
9 Phragmites communis
Smilax acutifolia 0 . 6 5 Amborella trichopoda Dioscorea floribunda Philactis zinnioides Sagittaria sp. Montrichardia arborescens
Beilschmiedia alata 9 Apollonias canariensis Sassafras albidum Calycanthus occidentalis 8
8 Piptostigma preussii
Oxandra lanceolata 7 Dioscorea floribunda Miliusa longipes Melodorum latifolium MeanDistance.MODE 6 Goetzea elegans Ephedranthus guianensis Archibaccharis subsessilis Ribes lacustre l o g ( m d a r e ) Sagittaria sp. Duguetia quitarensis Disepalum longipes 5 Cymbopetalum costaricensis
Bocageopsis multiflora 4 7 Bocagea laurifolia Emmotum acuminatum Asteranthe asterias Artabotrys rufus 3 Artabotrys thomsonii Anaxagorea prinoides Annona muricata Afroguatteria bequaertii Michelia figo 6 7 8 9 10 0.80 0.90 3 4 5 6 7 8 9 Parashorea plicata Magnolia grandiflora 6 Aristolochia taliscana Uraria clarksei Drimys lanceolata Canella winteriana Afrolicania elaeosperma Chloranthus glabra Unknown/Indet. Herbaceaous Woody Austrobaileya scandens Nuphar microphyllum Amborella trichopoda The within-family spread is a significant fraction of the overall Woodwardia japonica Onoclea sensibilis spread for all variables measured. There do not seem to be The seven herbaceous species we were able to sample do not obvious phylogenetic patterns evident above the level have areoles significantly larger than woody species. 500 2000 5000 20000 0.5 0.6 0.7 0.8 0.9 1.0 0.5 0.6 0.7 0.8 0.9 1.0 of family. 5 Areole area in pixels 6 Areole eccentricity Areole solidity 7 8
Akania hillii Sagittaria sp. Periomphale gaultherioides Amborella trichopoda Iresine interrupta ( Akaniaceae ) ( Alismataceae ) ( Alseuosmiaceae ) ( Amborellaceae ) ( Amaranthaceae )
Comparison of areole size with leaf rank D e n s i t y D e n s i t y D e n s i t y D e n s i t y red woody; black herbaceaous NCLC #: 8308 , Phot. #: 13 NCLC #: 797 , Phot. #: 15 NCLC #: 8418 , Phot. #: 17 NCLC #: 7864 , Phot. #: 19 NCLC #: 14981B , Phot. #: 21 r2= 0.237 , slope= -1.23 , intercept= 20.42 , p = 0.000180 , n= 49 4.5 5.5 6.5 7.5 0.65 0.75 0.85 0.95
Carya
Quercus 1 0 Anacardium pumilum Astronium fraxinifolium Buchanania sessilifolia Cotinus americanus Metopium brownei Artocarpus Platanus ( Anacardaceae ) ( Anacardaceae ) ( Anacardaceae ) ( Anacardaceae ) ( Anacardaceae ) 9 4 r 0 D e n s i t y D e n s i t y D e n s i t y D e n s i t y Ochna NCLC #: 4201 , Phot. #: 23 NCLC #: 4203 , Phot. #: 25 NCLC #: 12856 , Phot. #: 27 NCLC #: 4305 , Phot. #: 31 NCLC #: 4221 , Phot. #: 35 logArea.MODE 8 Leaf ranks provide a Vatica MeliosmAaextoxicon Akania 7 B r u n e l i a Aristolochia Celtis Acer Viburnum subjective, semi- Clethra Axinaea
6 Anac ardium Nothopegia castaneaefolia Rhus succedanea Afroguatteria bequaertii Anaxagorea prinoides Annona muricata Alnus Ch Itea eiloclin ( Anacardaceae ) ( Anacardaceae ) ( Annonaceae ) ( Annonaceae ) ( Annonaceae ) quantitative method ium Rauvolfia Gomphandra Bombax Afrolicania D e n s i t y D e n s i t y D e n s i t y D e n s i t y 7 . 5 Rinorea Vitis NCLC #: 15466a , Phot. #: 37 NCLC #: 12947 , Phot. #: 39 NCLC #: 4469 , Phot. #: 41 NCLC #: 9500 , Phot. #: 43 NCLC #: 19 , Phot. #: 45 Rhus Fraxinus of sampling more Metopium M a s t i x
Buchanania A r c t o s a p h y l 6 . 5 3 r 0 logPerim.MODE genera than we have Cordia Cunonia Salix Ilex Amelanchier Bursera Annonaceae Pittosporum 5 . yet been able to Artabotrys rufus Artabotrys thomsonii Asteranthe asterias Bocagea laurifolia Bocageopsis multiflora Schoepfia Baccharis Iresine ( Annonaceae ) ( Annonaceae ) ( Annonaceae ) ( Annonaceae ) ( Annonaceae ) 4.5 5.5 6.5 7.5 0.65 0.75 0.85 0.95 D e n s i t y D e n s i t y D e n s i t y D e n s i t y Ribes NCLC #: 5460 , Phot. #: 47 NCLC #: 5462 , Phot. #: 49 NCLC #: 3452 , Phot. #: 51 NCLC #: 4470 , Phot. #: 53 NCLC #: 2443 , Phot. #: 55
4 . 5 measure explicitly. Styloceras L e a f r n k Hickey, L. J. 1977. Stratigraphy and Paleobotany of the Magnolia 1 0 9
Golden Valley Formation (Early Tertiary) of Western North Cymbopetalum costaricensis Duguetia quitarensis Ephedranthus guianensis Melodorum latifolium Miliusa longipes logArea.MODE Cymbopetalum 8
0 . 9 ( Annonaceae ) ( Annonaceae ) ( Annonaceae ) ( Annonaceae ) ( Annonaceae ) 2 r 0 Eccentricity.MODE D e n s i t y D e n s i t y D e n s i t y D e n s i t y Dakota GSA Memoirs 150. Achatocarpus NCLC #: 2446 , Phot. #: 57 NCLC #: 9508a , Phot. #: 61 NCLC #: 9511 , Phot. #: 63 NCLC #: 2450 , Phot. #: 65 NCLC #: 7855 , Phot. #: 67 7 6 0 . 8 Boxplots of leaf rank by habit for 1620 genera Drought tolerance and leaf rank for 1620 genera 7 . 5
Oxandra lanceolata Piptostigma preussii Motandra guineensis Rauvolfia balansae Platanus occidentalis
0 . 9 5 ( Annonaceae ) ( Annonaceae ) ( Apocynaceae ) ( Apocynaceae ) ( Platanaceae ) D e n s i t y D e n s i t y D e n s i t y D e n s i t y NCLC #: 2892 , Phot. #: 69 NCLC #: 3922c , Phot. #: 71 NCLC #: 5099 , Phot. #: 75 NCLC #: 7388 , Phot. #: 77 NCLC #: 732 , Phot. #: 79 6 . 5 logPerim.MODE 0 . 8 5 Drimys
Solidity.MODE 5 . 1 r 0 4 r 0 0 . 7 5
Sassafras albidum Arctostaphylos nevadensis Carya ovata Brackenridgea nitida Ilex dipyrena 4 . 5 6.5 7.0 7.5 8.0 8.5 9.0 9.5 ( Lauaceae ) ( Ericaceae ) ( Juglandaceae ) ( Ochnaceae ) ( Aquifoliaceae ) D e n s i t y D e n s i t y D e n s i t y D e n s i t y 0 . 6 5 NCLC #: 771A , Phot. #: 85 NCLC #: 714 , Phot. #: 88 NCLC #: 6794 , Phot. #: 91 NCLC #: 2473 , Phot. #: 96 NCLC #: 11913 , Phot. #: 99
3 r 0 Log modal areole area in pixels 9 L e a f R n k 8 Eccentricity.MODE 0 . 9 7 Montrichardia arborescens Gastonia floribunda Aristolochia taliscana Austrobaileya scandens Alnus glabrata
MeanDistance.MODE 6 ( Araceae ) ( Araliaceae ) ( Aristolochiaceae ) ( Austrobaileyaceae ) ( Betulaceae ) 2 r 0 There is a strong correlation between measured areole area D e n s i t y D e n s i t y D e n s i t y D e n s i t y 5 NCLC #: NA , Phot. #: 101 NCLC #: 6087 , Phot. #: 103 NCLC #: 14922 , Phot. #: 105 NCLC #: 12191 , Phot. #: 109 NCLC #: 12183 , Phot. #: 112 0 . 8 4 and leaf rank. We therefore conclude that the lack of significant 3
ecological or phylogenetic signal in our measurements to date 0 . 9 5
6 7 8 9 10 0.80 0.90 3 4 5 6 7 8 9 1 r 0 Lundia spruceana Bixa excelsa Bombax aquaticum Cordia dichotoma Brunellia conocladifolia
is probably a result of insufficient taxon sampling. The final 0 . 8 5 parasite aquatic herb shrub woody tree vine Non-xeric Xeric ( Bignoniaceae ) ( Bixaceae ) ( Bombacaceae ) ( Boraginaceae ) ( Brunelliaceae ) tile shows some of our areole size measurements superimposed NCLC #: 218 , Phot. #:D e n s i t y 114 NCLC #: 3233 , Phot. #D e n s i t y : 116 NCLC #: 2526 , Phot. #D e n s i t y : 118 NCLC #: 220 , Phot. #:D e n s i t y 120 NCLC #: 6505 , Phot. #: 122 Solidity.MODE Herbaceous species do not clearly cluster when measured by 0 . 7 5 on the images from which they were obtained. This shows how
any of the variables considered. Genera with different growth forms and drought tolerances 0 . 6 5 the mode and spread of the measured size distribution Bursera gummifera Styloceras laurifolium Calycanthus occidentalis Canella winteriana Viburnum pauciflorum ( Burseraceae ) ( Buxaceae ) ( Calycanthaceae ) ( Canellaceae ) ( Caprifoliaceae ) have significantly different leaf ranks. 9 NCLC #: 1720 , Phot. #D e n s i t y : 124 NCLC #: 3203b , Phot. #D e n s i t y : 126 NCLC #: 2317 , Phot. #D e n s i t y : 128 NCLC #: 1159a , Phot. #D e n s i t y : 130 NCLC #: 821 , Phot. #: 132 9 10 successfully capture a gestalt impression of areole sizes. 11 8 7
MeanDistance.MODE 6 5 4 3
6 7 8 9 10 0.80 0.90 3 4 5 6 7 8 9
13 14 15 16