Seed surface characters and their systematic significance in the genus Lathyrus (Leguminosae, Papilionoideae, Vicieae)

Feddes Repertorium 118 (2007) 7–8, 269–285 DOI: 10.1002/fedr.200711139 Weinheim, Dezember 2007

© 2007 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim 0014-8962/07/7-812-269

1 Ain Shams University, Faculty of Education, Biological and Geological Sciences Department, Cairo 2 Alexandria University, Faculty of Science, Botany Department, Alexandria

M. M. ABOU-EL-ENAIN1; M. H. A. LOUFTY2 & A. A. SHEHATA1

Seed surface characters and their systematic significance in the genus Lathyrus (Leguminosae, Papilionoideae, Vicieae)

With 69 Figures and 4 Tables

Summary

Seed morphology of 34 species of Lathyrus was examined by light- (LM) and scanning electron mic-roscopy (SEM). Quantitative and qualitative features were evaluated to identify groups of species using a phenetic analysis. The used characters were that of testa (luster, multicellular sculpture and homogeneity of cell-size); anticlinal walls (level, undulation, thickness and 2ry sculpture); outer periclinal walls (level and 2ry sculpture); hilum (shape and length/ width ratio) and the seed length/width ratio. Nine multi-cellular sculptures and three patterns of secon-dary sculpture of anticlinal walls were recorded. The most common and remarkable pattern was the pro-minent stellate like structures on the angles of the anticlinal walls. Many terms used for describing patterns should be reevaluated within the whole tribe to better define of the character states for the multi-cellular sculpture. The present data were useful to recognize species or groups of species by a unique or combination of character states and had supported the retaining of Lathyrus tingitanus and L. tuberosus in section Lathyrus; the placement of L. neurolobus and L. nissolia in two different monotypic sections; the distinctiveness of the species of sections Aphaca and Pratensis.

Zusammenfassung

Samenschalenmerkmale und ihre systematische Bedeutung in der Gattung Lathyrus (Legumi-nosae, Papilionoideae, Vicieae) Die Samenmorphologie von 34 Lathyrus-Arten wurde mittels Licht- und Elektronenmikroskopie untersucht. Quantitative und qualitative Merkmale wurden gewichtet, um mittels phänetischer Analysen Artengruppen herauszuarbeiten. Berücksichtigt wur-den Merkmale der Testa (Glanz, vielzellige Skulptu-ren, Homogenität der Zellgröße); antiklinale Wände (Niveau, Undulation, Dicke und 2rv Skulptur); äußere periklinale Wände (Niveau und 2rv Skulptur); Hilum (Form und Länge : Breite); Länge : Breite des Sa-mens. Neun vielzellige Skulpturen und drei Muster der sekundären Skulptur der antiklinalen Wände wurden ermittelt. Das verbreitetste und bemerkens-werteste Muster ist die prominente sternähnliche Struktur in den Winkeln der antiklinalen Wände. Zahlreiche der zur Beschreibung der Muster benutz-ten Termini sollten neu bewertet werden, um in der gesamten Tribus den Status der multizellulären Skulpturen besser zu definieren. Die vorliegenden Daten sind geeignet, Arten oder Artengruppen durch ein Merkmal oder eine Kombinatuion von Merk-malsausprägungen zu erkennen. Sie haben den Ver-bleib von Lathyrus tingitanus und L. tuberosus in der Sektion Lathyrus, die Anordnung von L. neurolobus und L. nissolia in zwei verschiedene monotypische Sektionen und die Einordnung der Arten in die Sektionen Aphaca und Pratensis gestützt.

Introduction

The genus Lathyrus L. consists of ca. 160 spe-cies and is distributed throughout the temperate zones of the Northern Hemisphere and extends

into temperate South America and tropical East Africa (KENICER et al. 2005). The main center of diversity is the eastern Mediterranean region, with two secondary centers in North America and temperate areas of South America, respec-

270 Feddes Repert., Weinheim 118 (2007) 7–8

© 2007 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim www.fedrep.de

tively (KUPICHA 1983; ASMUSSEN & LISTON 1998). Most members of Lathyrus are annual or perennial mesophytic herbs with erect or, more usually, climbing and sprawling habit and oc-cur in open wood-lands, forest margins, roadsi-de verges, littoral, alpine, and drought habitats. They include food and fodder crops, ornamen-tals, soil nitrifiers, dune stabilizers, important agricultural weeds, and model organisms for genetic and ecological research (SEIJO & FERNÁNDEZ 2003). The genus is represented in Egyptian wild flora by eight annual species from which L. aphaca L., L. hirsutus L., L. sativus L. and L. marmoratus BOISS. & BLAN-CHE are found with a frequent distribution in Mediterranean and Oases regions, whereas L. setifolius L., L. sphaericus RETZ., L. annuus L., L. gorgonii PARL. are found with a rare to very rare distribution in the Eastern Mediterranean region (TÄCKHOLM 1974; BOULOS 1999). Among the taxonomic treatments of Lathy-rus, KUPICHA’s (1983) sectional delimitation is the most common used account (KENICER et al. 2005). Developing the work of GODRON (1848), BOISSIER (1872), BÄSSLER (1966, 1973, 1981), DAVIS (1970), and CZEFRANOVA (1971), she recognized 13 sections belonging to five subgenera namely: Lathyrostylis, Neu-rolobus, Orobus, Pratensis (subg. Orobus); Lathyrus, Linearicarpus, Notolathyrus, Oro-bus, Viciopsis (subg. Lathyrus) in addition to the monotypic subgenera Aphaca, Clymenum and Nissolia. Difficulties in determination the interspecific relationships have been encoun-tered by the workers in classification of Lathy-rus mainly because of the morphological ho-moplasy and lack of diagnostic characters (BADR et al. 2000; STEELE & WOJCIECHOWSKI 2003). Different criteria have been used to determine the interspecific relationships among the genus Lathyrus e.g. chromosome number (REES & HAZARIKA 1969; YAMAMOTO et al. 1984), karyology (ABOU-EL-ENAIN 1999; SEIJO & FERNÁNDEZ 2003), nuclear DNA (NA-RAYAN 1982; AHMAD & NARAYAN 1994), chemical compounds (ROBESON & HARBORNE 1980; SIMOLA 1986; RANAHABU & HARBORNE 1993) and molecular analysis (ASMUSSEN & LISTON 1998; BADR et al. 2000; KENICER et al. 2005). Although the seed coat characters have been studied by BUTLER (1986) in some Lathy-rus species of medical purposes, little is known

about the ultrastructure of the seed coat surface in most species of the genus. The importance of ultrastructural pattern analysis of the seed coat observed under the scanning electron microscope (SEM) – as a reliable approach for identifying the species and assessing taxonomic relationships – has been well recognized (BARTHLOTT 1981; BOE-SEWINKEL & BOUMAN 1995; KOUL et al. 2000; ARROYO-COSULTCHI et al. 2006; GAMARRA et al. 2007). The present work is dealing with determination of seed coat micro- and macro-morphological characters of 34 Lathyrus spe-cies using light- and scanning electron micros-copy; identifying taxa and establishing their inter-relationship using phenetic methods; comparing the produced data with those of the taxonomic literature in order to identify whe-ther seed characters support or oppose their previous treatment in the genus.

Material and methods

Mature seeds of 34 Lathyrus species were pro-vided by the Institute of Plant Genetics and Crop Plant Research (GAT), Gatersleben, Germany; the International Livestock Center for Africa (ILCA), Addis Ababa, Ethiopia, and the Plant Genetic Resources Conservation Unit of the United States Department of Agriculture (BARC), Georgia, USA (Table 1). Vouchers are deposited at the Herbarium of the Biologi-cal and Geological Sciences Department, Fac-ulty of Education, Ain Shams University (CAIA), Cairo, Egypt.

Scanning electron microscopy (SEM)

2–4 samples per species were washed in water, air-dried, investigated by the stereo-microcope, mounted on brass stubs and coated with a thin layer of gold using JEOL-JFCL 1100E ion sputtering. Coated seeds were examined by light-microscopy and pho-tographed on a JEOL-JSM5300 SEM with an accel-erating voltage of 15 kv at the Electron Microscopic Unit, Faculty of Science, Alexandria University. Close-up views were always taken from the lateral region of the seed. The characters evaluated were that of testa (luster, multicellular sculpture and ho-mogeneity of cell-size); anticlinal walls (level, undu-lation, thickness and 2ry sculpture); outer-periclinal walls (level and 2ry sculpture); hilum-micropyle region or HMR (shape and length/width ratio) and

M. M. ABDOU-EL-ENAIN et al.: Seed surface and systematic value in Lathyrus 271


Table 1 Specimens examined in this study; their sectional delimitation (KUPICHA 1983), sources, location and voucher data

Species Source Location Voucher data

Sect. Aphaca (J.MILL.) DUMORT. 1) L. aphaca L.

GAT

Italy

LAT114/94

Sect. Clymenum (J.MILL.) DC. ex SER. 2) L. articulatus L.

GAT

Morocco

LAT119/93 3) L. clymenum L. GAT Morocco LAT52/90 4) L. ochrus (L.) DC. BARCa Portugal 206373

Sect. Lathyrostylis (GRISEB.) BÄSSLER 5) L. cirrhosus SER.

GAT

Tunisia

LAT17/78 6) L. digitatus (M.BIEB.) FIORI. GAT Ukraine LAT8/89

Sect. Lathyrus L. 7) L. amphicarpos GOUAN.

GATb

Portugal

LAT139/94 8) L. annuus L. GAT Spain LAT152/87 9) L. blepharicarpus BOISS. ILCA Portugal 64985 10) L. cicera L. BARC Spain 208307 11) L. gorgonii HELDR. ex NYMAN GAT Jordan LAT101/84 12) L. hierosolymitanus BOISS. GAT Tunisia LAT142/84 13) L. hirsutus L. GAT Egypt LAT143/93 14) L. latifolius L. GAT Belgium LAT 10/91 15) L. marmoratus BOISS. & BLANCHE ILCAc Syria 65616 16) L. pseudocicera PAMP. ILCA Jordan 65214 17) L. rotundifolius JANKA GAT Armenia LAT 1289 18) L. sativus L. GAT Egypt LAT 52/87 19) L. sylvestris L. GAT Denmark LAT 4l89 20) L. tingitanus L. BARC USA 107839 21) L. tuberosus L. GAT Portugal LAT 11/91

Sect. Linearicarpus KUPICHA 22) L. angulatus L.

GAT

Spain

LAT151/95 23) L. inconspicuus L. GAT Iran LAT164/95 24) L. pannonicus (JACQ.) GARCKE GAT Italy LAT24/87 25) L. sphaericus DE RETZ GAT Jordan LAT34/96

Sect. Neurolobus BÄSSLER 26) L. neurolobus BOISS. & HELDR.

GAT

Iran

LAT19/82

Sect. Nissolia (J. MILL.) DUMORT. 27) L. nissolia L.

GAT

Turkey

LAT68/93

Sect. Orobus (L.) GODR. 28) L. aureus (STEVEN) BORNM.

GAT

Armenia

LAT27/89 29) L. davidii HANCE GAT Austria LAT21/82 30) L. niger (L.) BERNH. GAT Turkey LAT6/89 31) L. vernus (L.) BERNH. GAT Ukraine LAT18/90

Sect. Pratensis BÄSSLER 32) L. laxiflorus (DESF.) KUNTZ

GAT

Austria

LAT20/89 33) L. pratensis L. GAT Austria LAT 22/83 34) L. szowitsii BOISS. ILCA Guatemala 65122

a = United States Department of Agriculture (USDA), Georgia, USA b = Institute of Plant Genetics and Crop Plant Research (IPK), Gatersleben, Germany c = International Livestock Center for Africa gene bank (ILCA), Ethiopia, Addis Ababa



Table 2 Seed characters and character states used in the phenetic analysis

Testa: 1) Luster: glossy (GL), matte (MA), semi-matte (SM). 2) Multicellular sculpture: colliculate – ocellate (CL/OC), colliculate – ruminate (CL/RU), colliculate –

tuberculate (CL/TU), falsifoveate – colliculate (FF/CL), reticulate – colliculate (RT/CL), reticulate – favulariate (RT/FV), reticulate – foveate (RT/FO), reticulate monomorphic (RM), ribbed – ruminate (RB/RU), ruminate (RU).

3) Homogeneity of cell-size: uniform (UN), Irregular (IR).

Anticlinal walls of epidermal cells: 4) Level: highly raised (HR), raised (RA), leveled to sunken (LV/SK), irregular (IR). 5) Undulation: wavy (WA), straight (ST), sinuate (SN), slightly undulate (SU). 6) Thickness: very thick (VT), thick (TK), thin (TN), irregular (IR). 7) Secondary sculpture: stellate (SL), rodlike (RO), buttressed walls (BU).

Periclinal walls of epidermal cells: 8) Level: convex (CO), slightly concave (SC), concave (CN). 9) Secondary sculpture: striate (SI), ribbed to faveolariate (RB/FA), irregular to ribbed (IR/RB), papillate

(PA).

Hilum-micropylar region (HMR): 10) Shape: oblong (OB), ovate (OV), elliptic (EL). 11) Length/width ratio: ≤ 2, 2.1: 2.5, 2.6: 3.5, 3.6: 4.5, 4.6: 5.5, 5.6: 6.5, > 6.5.

Seed: 12) length/width ratio: 1.1 : 1.2, 1.21 : 1.3, 1.31 : 1.4, 1.41 : 1.5, 1.51 : 1.6, ≥ 1.6.

whole seed (length/width ratio). Seed and HMR dimensions were measured from the micrographs and the length/width ratios were calculated by using Microsoft Excel spreadsheet. The seed characters and character states and their abbreviations are given in Table 2. Terminology follows STEARN (1966), BARTHLOTT (1981, 1990).

Data preparat ion

The data editor program NTedit 2.2 (ROHLF 2005) was used for creating the data matrix of computa-tion, whereas the program NTSYS-pc 2.2 (ROHLF 2005) was used in all subsequent analyses. Each of the studied species was considered as operational taxonomic unit (OTU) and Vicia monantha L. was used as out group because of its relevance to Lathy-rus (ASMUSSEN & LISTON 1998). Multistate charac-ters were transformed to two-state characters in coding (SNEATH & SOKAL 1973; CRISCI & LÓPEZ-ARMENGOL 1983) and the raw data matrix was stan-dardized with STAND module.

Phenet ic analysis

Similarity matrix was generated by SIMQUAL module based on Jaccard’s coefficient (J) which is equivalent to Gower’s coefficient when a two-state character matrix is used (SNEATH & SOKAL 1973; ARROYO-COSULTCHI et al. 2006). A phenogram was

constructed by the unweighted pair-group method, arithmetic average (UPGMA). In order to test reli-ability of results, the correlation coefficient (r) value which measures the distortion between the produced phenogram and the relevant similarity matrix (ROHLF & SOKAL 1981) was estimated as follows: The cophenetic (ultramatric) value matrix of the phenogram was computed using COPH module and compared to the related distance matrix using MXCOMP module.

Results

SEM micrographs of epidermal cell shape and multicellular sculpture in the lateral region of mature seeds of the studied species are revealed in Figs. 1–34, whereas those of hilum-micro-pyle region (HMR) are shown in Figs. 35–68. The qualitative and quantitative characters of seed gross morphology and testa appearance are given in Table 3, whereas those of anticli-nal; periclinal walls of testa epidermal cells and HMR are revealed in Table 4. Basic data matrix of the 53 character states of seeds similarity used in the phenetic analysis is given in Ap-pendix 1, whereas the produced UPGMA clus-tering phenogram is illustrated in Fig. 69.



Figs. 1–18 SEM micrographs of epidermal cell shape and multicellular sculpture in the lateral region of mature seeds 1 — L. aphaca; 2 — L. articulatus; 3 — L. clymenum; 4 — L. ochrus — 5 — L. cirrhosus; 6 — L. digitatus; 7 — L. amphicarpos; 8 — L. annuus; 9 — L. blepharicarpus; 10 — L. cicera; 11 — L. gorgonii; 12 — L. hierosolymitanus; 13 — L. hirsutus; 14 — L. latifolius; 15 — L. marmoratus; 16 — L. pseudocicera; 17 — L. rotundifolius; 18 — L. sativus

Scale: 5, 7, 8, 17 = 2.5 µm; the remaining = 1 µm



Figs. 19–34 SEM micrographs of epidermal cell shape and multicellular sculpture in the lateral region of mature seeds 19 — L. sylvestris; 20 — L. tingitanus; 21 — L. tuberosus; 22 — L. angu-latus; 23 — L. inconspicuus; 24 — L. pannonicus; 25 — L. sphaericus; 26 — L. neurolobus; 27 — L. nissolia; 28 — L. aureus; 29 — L. davidii; 30 — L. niger; 31 — L. vernus; 32 — L. laxiflorus; 33 — L. pratensis; 34 – L. szowitsii

Scale: 1 µm



Figs. 35–68 SEM micrographs of the hilum-micropylar region of mature seeds 35 — L. aphaca; 36 — L. articulatus; 37 — L. clymenum; 38 — L. ochrus; 39 — L. cirrhosus; 40 — L. digitatus; 41 — L. amphicarpos; 42 — L. annuus; 43 — L. blepharicarpus; 44 — L. cicera; 45 — L. gorgonii; 46 — L. hierosolymitanus; 47 — L. hirsutus; 48 — L. latifolius; 49 — L. marmoratus; 50 — L. pseudocicera; 51 — L. rotundifolius; 52 — L. sativus; 53 — L. sylvestris; 54 — L. tingitanus; 55 — L. tuberosus; 56 — L. angulatus; 57 — L. inconspicuus; 58 — L. pannonicus; 59 — L. sphaericus; 60 — L. neurolobus; 61k — L. nissolia; 62 — L. aureus; 63 — L. davidii; 64 — L. niger; 65 — L. vernus; 66 — L. laxiflorus; 67 — L. pratensis; 68 — L. szowitsii

Scale: 44, 56, 57 & 67 = 50 µm; 54 and 61 = 250; the remaining = 100 µm



Table 3 Qualitative and quantitative characters of seed gross morphology and testa appearance in the studied species of Lathyrus. For abbreviations, see Table 2

Seed Testa apperance

Size

Species

Shape Colour

Mean Length ± S.E (mm)

Mean Width ± S.E (mm)

L/W Ratio

Luster

Multi- cellular sculpture

Homo- geneity of cell size

1) L. aphaca Oval Brown 3.30 ± 0.04 2.30 ± 0.04 1.44 SM RT/FV IR 2) L. articulatus Oval Brown 5.80 ± 0.08 4.50 ± 0.02 1.28 SM CL/TU IR 3) L. clymenum Oblong Cream with

brown batches 6.20 ± 0.02 5.12 ± 0.01 1.21 SM CL/TU IR

4) L. ochrus Globose Pale brown 6.00 ± 0.01 4.90 ± 0.02 1.23 SM CL/TU IR 5) L. cirrhosus Oval Brown with

dark batches 5.50 ± 0.03 3.50 ± 0.04 1.53 MA RU IR

6) L. digitatus Oval Brown with dark batches

3.90 ± 0.02 2.90 ± 0.02 1.35 MA RU IR

7) L. amphicarpos Oval Pale brown 4.60 ± 0.05 4.00 ± 0.02 1.15 SM RT/FO IR 8) L. annuus Oblong Brown 4.60 ± 0.07 4.15 ± 0.08 1.11 GL RT/FO IR 9) L. blepharicarpus Oblong Cream with

brown batches 5.45 ± 0.05 4.65 ± 0.02 1.17 GL RT/FO IR

10) L. cicera Oval Yellowish brown

5.50 ± 0.07 4.75 ± 0.07 1.16 GL RT/FO IR

11) L. gorgonii Oblong Pale brown 4.80 ± 0.05 4.10 ± 0.02 1.17 SM RT/FO UN 12) L. hierosolymitanus

Oblong Brown 4.60 ± 0.01 3.90 ± 0.02 1.18 MA RT/FO IR

13) L. hirsutus Oval Dark brown 3.35 ± 0.06 3.00 ± 0.04 1.12 GL RT/FO IR 14) L. latifolius Oval Brown 5.70 ± 0.12 4.80 ± 0.15 1.19 GL RT/FO IR 15) L. marmoratus Oblong Cream with

brown batches 5.25 ± 0.02 4.55 ± 0.02 1.15 MA RT/FO IR

16) L. pseudocicera Oblong Cream with brown batches

4.60 ± 0.05 3.83 ± 0.03 1.20 GL RT/FO IR

17) L. rotundifolius Oblong Dark brown 4.60 ± 0.08 3.90 ± 0.07 1.18 SM RT/FO IR 18) L. sativus Globose Cream with

brown batches 6.40 ± 0.05 5.40 ± 0.06 1.19 GL RT/FO IR

19) L. sylvestris Globose Brown 4.00 ± 0.05 3.60 ± 0.04 1.11 GL RT/FO IR 20) L. tingitanus Globose Brown 5.10 ± 0.09 4.60 ± 0.11 1.11 GL RT/FO IR 21) L. tuberosus Oval Dark brown 4.25 ± 0.03 3.60 ± 0.04 1.18 GL RT/FO IR 22) L. angulatus Oblong Dark brown 3.40 ± 0.07 2.60 ± 0.03 1.30 GL RB/RU IR 23) L. inconspicuus Oval Dark brown 3.90 ± 0.04 3.00 ± 0.03 1.30 GL RB/RU IR 24) L. pannonicus Oblong Dark brown 4.25 ± 0.12 2.60 ± 0.10 1.64 GL RB/RU IR 25) L. sphaericus Globose Dark brown 3.50 ± 0.02 2.40 ± 0.02 1.46 GL RB/RU IR 26) L. neurolobus Globose Brown 1.95 ± 0.03 1.20 ± 0.05 1.63 MA CL/RU UN 27) L. nissolia Globose Brown 2.20 ± 0.03 1.80 ± 0.03 1.22 GL FF/CL UN 28) L. aureus Oblong Brown 4.50 ± 0.02 3.50 ± 0.06 1.28 SM RT/CL UN 29) L. davidii Oblong Reddish

brown 3.15 ± 0.03 2.55 ± 0.05 1.24 SM RT/CL UN

30) L. niger Oblong Brown 4.10 ± 0.12 3.20 ± 0.09 1.28 SM RT/CL UN 31) L. vernus Oblong Dark brown 4.00 ± 0.06 2.90 ± 0.04 1.38 SM RT/CL IR 32) L. laxiflorus Oval Brown 3.30 ± 0.06 2.60 ± 0.08 1.27 SM CL/OC UN 33) L. pratensis Globose Brown with

dark batches 2.86 ± 0.02 2.30 ± 0.02 1.24 SM CL/OC UN

34) L. szowitsii Oval Brown with dark batches

6.20 ± 0.02 5.10 ± 0.05 1.22 SM CL/OC UN



Table 4 Qualitative and quantitative characters of anticlinal, periclinal walls of testa epidermal cells and hilum-micropyle region (HMR) in the studied species of Lathyrus. For abbreviations, see Table 2

Anticlinal walls Periclinal walls HMR Species

Level Undu- lation

Thick- ness

2ry Scul- pture

Level 2ry Scul- pture

Shape Mean Length (mm)

Mean Width (mm)

L/W Ratio

1) L. aphaca HR SU IR N CO SI OV 0.4 0.2 2.00 2) L. articulatus IR SU IR SL SC RB/FA EL 0.2 0.1 2.00 3) L. clymenum IR SU IR SL SC RB/FA EL 1.4 0.3 4.70 4) L. ochrus IR SU IR SL SC IR/RB EL 0.6 0.1 3.00 5) L. cirrhosus HR SN TK RO SC RB/FA OV 0.6 0.3 2.00 6) L. digitatus HR SN TK RO SC RB/FA OV 0.4 0.2 2.00 7) L. amphicarpos IR SU IR SL SC RB/FA EL 0.5 0.3 1.70 8) L. annuus IR SU IR SL SC IR/RB OB 1.0 0.3 3.30 9) L. blepharicarpus IR SU IR SL SC IR/RB EL 0.7 0.3 2.30 10) L. cicera IR SU IR SL SC IR/RB OV 0.6 0.3 2.00 11) L. gorgonii IR SU IR SL SC IR/RB EL 0.5 0.2 2.50 12) L. hierosolymitanus IR SU TN SL SC IR/RB EL 0.8 0.4 2.00 13) L. hirsutus IR SU TK SL SC RB/FA EL 0.8 0.4 2.00 14) L. latifolius IR SU TK SL SC IR/RB OB 1.6 0.3 5.30 15) L. marmoratus IR SU IR SL SC IR/RB EL 0.5 0.2 2.50 16) L. pseudocicera IR SU IR SL SC RB/FA OV 0.4 0.3 1.30 17) L. rotundifolius IR ST IR SL SC IR/RB EL 0.3 0.2 1.50 18) L. sativus IR SU IR SL SC RB/FA OB 0.7 0.2 3.50 19) L. sylvestris IR SU IR SL SC IR/RB EL 0.4 0.2 2.00 20) L. tingitanus IR SU IR SL SC IR/RB OB 1.3 0.2 6.50 21) L. tuberosus IR SU IR SL CO IR/RB EL 0.6 0.3 2.00 22) L. angulatus HR ST TK SL SC RB/FA OB 0.9 0.2 4.50 23) L. inconspicuus HR ST TK SL SC RB/FA EL 0.3 0.2 1.50 24) L. pannonicus HR ST TK SL SC RB/FA EL 0.6 0.1 2.00 25) L. sphaericus HR ST TK SL SC RB/FA EL 1.3 0.4 4.30 26) L. neurolobus IR SU IR RO CO IR/RB EL 0.2 0.1 2.00 27) L. nissolia IR SU TK RO SC IR/RB OV 0.6 0.2 3.00 28) L. aureus RA ST TN SL SC IR/RB OB 1.5 0.4 3.75 29) L. davidii RA ST TN BU CO RB/FA OB 1.2 0.4 3.00 30) L. niger RA SN TN BU CO IR/RB OB 0.5 0.2 2.50 31) L. vernus RA ST TN BU CO RB/FA OB 1.5 0.3 5.00 32) L. laxiflorus LV/SK SU IR RO CO SI OB 0.6 0.1 6.00 33) L. pratensis LV/SK SU IR RO CO SI OB 0.5 0.2 2.50 34) L. szowitsii LV/SK ST IR RO CO SI EL 0.5 0.3 1.70

Gross morphology, testa appearance and cell-pattern

Seeds of the studied species are asymmetric and varies in shape from oval, globose to ob-long (Table 3). The seed color is ranging from dark brown with its grades to yellow except for L. cirrhosus, L. digitatus, L. pratensis and L. szowitsii have dark brown batches; L. blepha-ricarpus, L. clymenum, L. marmoratus, L. pseu-docicera and L. sativus have pale brown bat-ches. Size varies from small in L. neurolobus

(1.95 ± 0.03 × 1.20 ± 0.05 mm) and L. nissolia (2.20 ± 0.03 × 1.80 ± 0.03 mm) to very large in L. sativus (6.40 ± 0.05 × 5.40 ± 0.06 mm), but most species have sizes between medium ≈3 × 2 mm and large ≈5 × 4 mm. Their length/ width ratio are ranged from 1.11 in L. annuus, L. sylvestris and L. tingitanus to 1.64 in L. pannonicus. Among the 34 species investi-gated, seed surface is glossy (GL) in 15, semi-matte (SM) in 14 and matte (MA) exclusively in L. cirrhosus, L. digitatus, L. hierosolymita-



Fig. 69 UPGMA phenogram showing clustering of the studied Lathyrus species based on 53 seed character states. Capital letters indicate the large groups, lowercase letters the subgroups. nus, L. marmoratus and L. neurolobus. Nine multicellular sculpture patterns are recognized (Table 3): reticulate-foveate (RT/FO) in 15 species (Figs. 7–21); colliculate-tuberculate (CL/TU) in L. articulatus, L. clymenum and L. ochrus (Figs. 2–4, respectively); falsifove-ate-colliculate (FF/CL) in L. nissolia (Fig. 27); colliculate-ruminate (CL/RU) in L. neurolobus (Fig. 26); ribbed-ruminate (RB/RU) in L. angu-latus, L. inconspicuus, L. pannonicus, L. sphae-ricus (Figs. 22–25, respectively); ruminate (RU) in L. cirrhosus and L. digitatus (Figs. 5, 6, respectively); reticulate-favulariate (RT/ FV) in L. aphaca (Fig. 1); colliculate-ocellate (CL/OC) in L. laxiflorus, L. pratensis, L. szo-witsii (Figs. 32–34, respectively); reticulate-colliculate (RT/CL) in L. aureus, L. davidii, L. niger, L. vernus (Figs. 28–31, respectively). Testa cells size of the studied species are vari-

able from uniform (UN) in L. gorgonii, L. neu-rolobus, L. nissolia, L. aureus, L. davidii, L. ni-ger, L. laxiflorus, L. pratensis and L. szowitsii (Figs. 11, 26–30, 32–34, respectively) to ir-regular (IR) in the remaining species. No no-table variation in size towards the hilum has been observed in all cases.

Anticlinal walls of testa epidermal cells

Anticlinal cell boundaries (Table 4) are highly raised (HR) in L. aphaca, L. cirrhosus, L. digi-tatus, L. angulatus, L. inconspicuus, L. pan-nonicus and L. sphaericus (Figs. 1, 5, 6, 22–25, respectively); leveled to sunken (LV/SK) in L. laxiflorus, L. pratensis and L. szowitsii (Figs. 32–34, respectively); raised (RA) in L. aureus, L. davidii, L. niger and L. vernus (Figs. 28–31, respectively); irregular in the remaining spe-



cies. The walls are thick (TK) in L. cirrhosus, L. digitatus, L. hirsutus, L. latifolius, L. angula-tus, L. inconspicuus, L. pannonicus, L. sphaeri-cus and L. nissolia (Figs. 5, 6, 13, 14, 22–25, 27, respectively); thin (TN) in L. hierosolymitanus, L. aureus, L. davidii, L. niger and L. vernus (Figs. 12, 28–31, respectively); ir-regular (IR) in the remaining species. Undula-tion (SU) of the anticlinal wall is observed in 19 of the 34 species studied; but absent (ST) in L. rotundifolius, L. angulatus, L. inconspicuus, L. pannonicus, L. sphaericus, L. aureus, L. da-vidii, L. vernus and L. szowitsii (Figs. 17, 22, 23, 25, 26, 28, 29, 31, 34, respectively) where-as sinuate (SN) in L. cirrhosus, L. digitatus and L. niger (Figs. 5, 6, 30, respectively). Three patterns of secondary sculpture in anticlinal walls are observed, the first and most remark-able is the prominent stellate like structures on the angles of the anticlinal walls (SL) in 27 of species. The second is buttresses (BU) in L. da-vidii, L. niger and L. vernus (Figs. 29–31, respectively). The third are rodlike structures (RO) protruding from some walls in L. cirrho-sus, L. digitatus, L. neurolobus, L. nissolia, L. laxiflorus, L. pratensis and L. szowitsii (Figs. 5, 6, 26, 27, 32–34, respectively).

Periclinal walls and characters of the HMR

The periclinal walls (Table 4) are flat to slight-ly concave (SC) in the studied species except in L. aphaca, L. tuberosus, L. neurolobus, L. davi-dii, L. niger, L. vernus, L. laxiflorus, L. praten-sis, and L. szowitsii (Figs. 1, 21, 26, 29–34, respectively), where they are concave (CO). Secondary sculpture (microrelief) is irregularly ribbed (IR/RB) in 16 of the species studied (Table 4); ribbed to faveolariate (RB/FA) in 14 species and striate only in L. aphaca, L. laxiflo-rus, L. pratensis and L. szowitsii (Figs. 1, 32–34, respectively). The HMR shape (Fig. 3) is elliptic in 17 species; oblong in 11 species and ovate in L. aphaca, L. cirrhosus, L. digitatus, L. cicera, L. pseudocicera and L. nissolia (Figs. 1, 5, 6, 10, 16, 27, respectively). HMR size varied from 0.2 × 0.1 mm in L. articulatus and L. neurolobus (Figs. 36, 60, respectively) to 1.6 × 0.3 mm in L. latifolius (Fig. 48), whereas their length/width ratio are ranged from 1.3 in L. pseudocicera (Fig. 50) to 6.5 in L. tingitanus (Fig. 54).

Phenetic analysis

The produced phenogram (Fig. 69) revealed that, three clusters A, B and C are separated at the levels of 0.20, 0.27 and 0.28, respectively. From which B and C are united together at the level of 0.21 and clustered with A at the level of 0.11. The first cluster (A) is split into three groups i.e. a1, a2 and a3 from which a1 and a2 are grouped together at the level of 0.27 and cluste-red with a3 at the level of 0.20. The first group (a1) is represented by single phenetic line that joins L. aphaca. The second group (a2) is com-prised L. laxiflorus, L. pratensis and L. szo-witsii. The third (a3) comprises L. aureus, L. niger, L. davidii and L. vernus. The second cluster (B) comprises four groups (b1, b2, b3 and b4) that are distinguished from each other at the levels of 0.56, 0.44 and 0.33, respectively. The first group (b1) consists of the species L. articu-latus, L. clymenum and L. ochrus. The second (b2) is comprised of the species L. amphicarpos, L. hirsutus, L. cicera, L. sylvestris, L. tuberosus, L. pseudocicera, L. rotundifolius, L. blepharicarpus, L. marmoratus, L. gorgonii, L. hierosolymitanus, L. annuus, L. sativus, L. tingitanus and L. latifolius. Each of the third (b3) and the fourth (b4) is represented by unique phenetic line, each of which join a single spe-cies i.e. L. neurolobus and L. nissolia, respecti-vely. The third cluster (C) is split into two groups (c1 and c2) that are distinguished at the levels of 0.63 and 0.85, respectively. The first group (c1) comprises L. cirrhosus and L. digi-tatus, whereas the second (c2) consisted of L. angulatus, L. inconspicuus, L. sphaericus and L. pannonicus. The produced correlation coef-ficient (r) value of 0.94 reveals no significant distortion between such phenogram and its related similarity matrix.

Discussion

The produced phenogram (Fig. 69) based on cluster analysis of the seed characters for the studied species was clearly yielded three major clusters. The first (Fig. 69, A) was distinguished by the semi-matte testa (character no. 3, Appendix 1) and convex periclinal walls (31). L. aphaca was distinguished in the first phene-tic line (i.e. group a1) due to the presence of reticulate-favulariate multicellular sculpture of



anticlinal walls (9). The second group (a2) was comprised L. laxiflorus, L. pratensis and L. szowitsii with colliculate-ocellate multicellu-lar sculpture (4) and slightly sunken (18) anti-clinal walls. The third (a3) was comprised L. aureus, L. niger, L. davidii and L. vernus with reticulate-colliculate (8) multicellular sculpture; slightly raised (17) and thin (26) anticlinal walls. Within group a2, L. szowitsii was distinctive by the absence of undulation of the anticlinal walls (21) and the elliptic hilum shape (40). L. pratensis and L. laxiflorus were characterized from each other by the hilum length/width ratio of 4.6:5.5 (45) and of 5.6:6.5 (46), respectively. Within group a3, L. aureus was characterized by the prominent stellate like secondary sculpture on the angles of the anticlinal walls (28) and the slightly con-cave periclinal walls (32). L. niger was unique by it’s the sinuate anticlinal walls (22) and the hilum length/width ratio of 5.6:6.5 (46). L. davidii was characterized from L. vernus as well as from the other species in the group by the hilum length/width ratio of ≤2 (41). The second cluster (Fig. 69, B) comprised of 20 species that were grouped together due to the irregular testa cell size (Character no. 15, Appendix 1) and each of the irregular level (19); slightly undulate (23); and the stellate structures (28) of the anticlinal walls. The first minor group (b1) was comprised of three spe-cies L. articulatus, L. clymenum and L. ochrus possessing matte testa (2), colliculate-tubercu-late multicellular sculpture of anticlinal walls (6) and seed length/width ratio ranged between 1.21 and 1.3 (49). Within this group, L. ochrus was distinctive by glossy testa (1) and the hi-lum length/ width ratio of 5.6:6.5 (46). L. ar-ticulatus and L. clymenum were characterized from each other by the presence of hilum length/width ratio of 2.1:2.5 (42) and ≤2 (41), respectively. The second minor group (b2) consisted of 15 species with glossy testa (1), reticulate-foveate multicellular sculpture of anticlinal walls (10) and seed length/width ratio ranged between 1.1 and 1.2 (48). Within this group (b2), the species studied were divided into five subgroups. First subgroup (L. amphi-carpos and L. hirsutus) was characterized by the ribbed to faveolariate secondary sculpture of periclinal walls (Character no. 35). The second subgroup (L. cicera, L. sylvestris, L. tu-

berosus and L. pseudocicera) was characterized by the length/width ratio of ≤2 (41). Within this subgroup, L. pseudocicera was similar to the species of the latter group in having the ribbed to faveolariate secondary sculpture of periclinal walls, whereas L. tuberosus was delimited by the convex periclinal walls (31). L. cicera and L. sylvestris were characterized from each other by the presence of ovate sha-ped hilum (39). The third subgroup was repre-sented by a single phenetic line joining L. ro-tundifolius that characterized by the straight anticlinal walls (21). The fourth subgroup (L. blepharicarpus, L. marmoratus, L. gorgonii and L. hierosolymitanus) was distinctive by the elliptic shaped hilum (40). Three species of this subgroup possessed a hilum length/width ratio of 2.1:2.5 (42), whereas L. hierosolymitanus was delimited by the hilum length/width ratio of ≤2 (41). L. blepharicarpus and L. marmora-tus were differentiated from each other by the glossy (1) and matte (2) testa, respectively. The fifth (L. annuus, L. sativus, L. tingitanus and L. latifolius) was delimited by the oblong hilum (38). From this subgroup, L. latifolius was distinctive by the thick anticlinal walls (25). L. tingitanus was delimited due to the hilum length/width ratio of 5.6:6.5 (46), whereas L. annuus and L. sativus were differentiated from each other by the ribbed to faveolariate secondary sculpture of periclinal walls (35) in L. sativus. Each of the third and fourth minor groups (b3 and b4) i.e. L. neurolobus and L. nissolia, was exclusively characterized by falsifoveate-colliculate (7) and colliculate-ruminate (5) multicellular sculpture of anticli-nal walls, respectively. The third cluster (Fig. 69, C) comprises six species that were mainly grouped together by the highly raised (16) and thick (25) anticlinal walls. The first group (c1) included L. cirrhosus and L. digitatus with ruminate multicellular sculpture (13) and sinuate (22) anticlinal walls. Within the group c2, L. angulatus was unique by its elliptic shaped hilum (38) and the hilum length/width ratio of 3.6:4.5 (44). L. pannoni-cus was characterized by the hilum length/ width ratio of 2.1:2.5 (42). The remaining species (L. inconspicuus and L. sphaericus) were characterized from each other by the presence of the seed length/width ratio of 1.41:1.5 (51) in L. sphaericus. The second



group (c2) consisted of L. angulatus, L. incon-spicuus, L. sphaericus and L. pannonicus that were characterized by the ribbed-ruminate multicellular sculpture (12) and absence of undulation (21) of anticlinal walls. The groups a1, a2, a3, b1, b2, b3, b4, c1 and c2 described above are corresponding to the sec-tions Aphaca, Pratensis, Orobus, Clymenum, Lathyrus, Neurolobus, Nissolia, Lathyrostylis and Linearicarpus, respectively, which further supporting delimitation of the studied species in the currently recognized sections as pro-posed by KUPICHA (1983), ASMUSSEN & LIS-TON (1998), KENICER et al. (2005). However, some new groups are established based on the seed similarity.

Sections Aphaca and Pratensis

The monotypic section Aphaca is represented herein by L. aphaca (Fig. 69, group a1), whe-reas section Pratensis (approximately six spe-cies) is represented by L. laxiflorus, L. praten-sis and L. szowitsii (Fig. 69, group a2). Mem-bers of these two sections share distinctive wing-petal architecture and sagittate stipules supplied by an unusual vascular arrangement where the petiole is supplied only by the medi-an traces. Such robust synapomorphies are rare in Lathyrus and prompted KUPICHA (1975, 1983) to suggest that the two sections are close-ly related, a proposal supported by cpDNA, RFLP, seed protein and ITS data (ASMUSSEN & LISTON 1998; BADR et al. 2000; KENICER et al. 2005). Despite this, the present data supports the conclusion of KUPICHA: these sections should be retained as separate because of their morphological distinctiveness. Section Aphaca is distinguished from section Pratensis by its annual habit, large stipules and lack of leaflets. RANABAHU & HARBORNE (1993) also con-cluded similar view based on the variation in structure of flavones, flavonols and proantho-cyanidine in the aerial parts and seeds of the species in the two sections Pratensis and Aphaca. Section Lathyrus

KUPICHA’s (1983) section Lathyrus is repre-sented herein by 15 species that were clustered as a distinctive minor group (Fig. 69, group b2). L. tingitanus and L. tuberosus are suffering from many homoplasious characters (DOGAN et

al. 1992; ASMUSSEN & LISTON 1998), which caused them to be variably placed among the sections of Lathyrus. DOGAN et al. (1992) re-tained L. tingitanus in section Lathyrus and separated L. tuberosus in section Orobus. How-ever, ASMUSSEN & LISTON (1998) rejected the sectional isolation of L. tuberosus proposed by DOGAN et al. (1992) but reported the possibility of separating L. tingitanus in a distinct mono-typic section. BADR et al. (2002) and KENICER et al. (2005) retained both of the species among the section Lathyrus, although they supported ASMUSSEN & LISTON’s (1998) circumscription in the strict sense. The present data is further supporting the retaining each of L. tingitanus and L. tuberosus into section Lathyrus as cir-cumscribed by KENICER et al. (2005) and BADR et al. (2002). DOGAN et al. (1992) placed L. hirsutus in the section Cicercula as defined by DAVIS (1970), and treated L. blepharicarpus, L. ci-cera, L. pseudocicera and L. sativus as belong-ing to section Clymenum. They proposed a new section Gorgonia to accommodate L. gorgonii in addition to L. rotundifolius, which has al-ways been placed in section Lathyrus. The present data contrasts the view of DOGAN et al. (1992) regarding either the establishment of section Gorgonia or the circumscription of section Clymenum, but support the inclusion of these species in KUPICHA’s (1983) section Lathyrus.

Sections Clymenum, Neurolobus and Nissolia

Section Clymenum is represented by L. articu-latus, L. clymenum and L. ochrus (Fig. 69, group b1). L. articulatus is often considered to be synonymous with L. clymenum, and these are similar to L. ochrus, all of which are tradi-tionally placed in section Clymenum on the basis of their phyllodic leaves (KUPICHA 1983). KENICER et al. (2005) confirmed such relation-ship where L. ochrus appeared as sister to L. clymenum in a combined analysis of trnL-F and ITS. The present data is in accordance with this view. KUPICHA (1983) placed L. neu-rolobus and L. nissolia in monotypic sections Neurolobus and Nissolia, respectively based on morphology and habit and suggested that they both occupy isolated positions within the ge-nus. KENICER et al. (2005) reported that, they



are collectively sister group to the rest of Lathyrus and confirmed the close relationship between them. BADR et al. (2000) considered L. neurolopus as a relict species because of it is morphologically distinguished by being a per-ennial and resembles annual Lathyrus species in being autogamous (KUPICHA 1983). Regard-ing this species, as a monotypic section as proposed by BÄSSLER (1966) or its assignment to another section could not be confirmed by evidence from cpDNA (ASMUSSEN & LISTON 1998). The present data supports the placement of L. neurolobus and L. nissolia (Fig. 69, groups a3 and a4, respectively) as monotypic sections as proposed by KUPICHA (1983).

Sections Orobus, Lathyrostylis and Linearicarpus

Section Orobus is the largest and most diverse section in the genus with wide range of mor-phological variation (ASMUSSEN & LISTON 1998). All of its species are perennials with multijugate leaves, purple; many flowered inflorescences; broad deltoid lower calyx teeth; semi-sagittate, foliacious stipules; aristate ra-chises; and three parallel primary leaflet veins. However, most of these characters are also common in approximately 20 erect perennial species constitutes BÄSSLER’s (1966) proposed section Lathyrostylis. This situation made it difficult for CZEFRANOVA (1971) to separate many members from section Lathyrostylis and prompted him with many other authors to treat these species as part of a broader section Oro-bus. BÄSSLER (1966) claimed the two sections to be distinct, describing section Platystylis, which later renamed Lathyrostylis (BÄSSLER 1971, 1981). Most of the subsequent studies agreed with this circumscription (KUPICHA 1983; BADR et al. 2000; KENICER et al. 2005). The seed data presented here are indicating the assignment of Lathyrostylis and Orobus as de-finitely separate sections, which is further sup-porting such interpretations. On the other hand, KENICER et al. (2005) proposed a close rela-tionship between L. sphaericus and L. angula-tus of KUPICHA’s (1983) section Linearicarpus to the section Lathyrostylis. They invoked the questioned monophyly of the section Lineari-carpus established by DOGAN et al. (1992), ASMUSSEN & LISTON (1998) and BADR et al. (2002). In the present study, the species of

section Linearicarpus i.e. L. angulatus, L. in-conspicuus, L. sphaericus and L. pannonicus are clearly distinct from those of section Lathy-rostylis i.e. L. cirrhosus and L. digitatus (Fig. 69, groups c2 and c1, respectively) which dis-agree with the hypothesis of KENICER et al. (2005). However, further phylogenetic data are needed for these difficult sections before any firm systematic decisions can be made. In conclusion, SEM helped to identify twelve characters and allowed to recognize nine multi-cellular sculptures patterns of anti-clinal walls: reticulate-foveate (RT/FO) is sha-red with the fifteen species of section Lathyrus; colliculate-tuberculate (CL/TU) in section Cly-menum (L. articulatus, L. clymenum, L. ochrus); colliculate-ruminate (CL/RU) in section Neuro-lobus (L. neurolobus); falsifoveate-colliculate (FF/CL) in section Nissolia (L. nissolia); rumi-nate (RU) in the species of the section Lathy-rostylis (L. cirrhosus and L. digitatus); ribbed-ruminate (RB/RU) in sections Linearicarpus (L. angulatus, L. inconspicuus, L. pannonicus, L. sphaericus); reticulate-favulariate (RT/FV) in section Aphaca (L. aphaca); reticulate-colli-culate (RT/CL) in section Orobus (L. aureus, L. davidii, L. niger); colliculate-ocellate (CL/ OC) in section Pratensis (L. laxiflorus, L. pra-tensis and L. szowitsii). However, many terms used for describing patterns should be reevalua-ted within the whole tribe to better define of the character states for the multicellular sculpture. Three patterns of secondary sculpture in anticlinal walls are observed, the most remarkable is the prominent stellate like structures on the angles of the anticlinal walls and were observed in 27 species. Seed characters described here are useful to recognize species or groups of species by a unique or combination of character states. The present data further supported each of the retaining of L. tingitanus and L. tubero-sus in section Lathyrus; the placement L. neu-rolobus and L. nissolia in two different mono-typic sections; the distinctiveness of the species of sections Aphaca and Pratensis. On the other hand, the present data did not support each of the separation of L. gorgonii in section Gorgo-nia; inclusion of L. blepharicarpus, L. cicera, L. pseudocicera and L. sativus in section Cly-menum; placing of L. angulatus and L. sphaericus in the section Lathyrostylis. Grouping of the 34 species included in this study in



nine sections is generally in accordance with their previous circumscription as proposed by KUPICHA (1983), ASMUSSEN & LISTON (1998) and KENICER et al. (2005), so that supports the validity of using the seed scanning criteria as valid taxonomic evidence in the genus Lathy-rus.

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Addresses of the authors:

Dr. Maged Mahmoud Abou-El-Enain , Corres- pondent author, Dr. Mohammed Hesham A. Lout fy , Ain Shams University, Faculty of Educa-tion, Biological and Geological Sciences Depart-ment, Roxy, Heliopolis, P.C.11757, Cairo, Egypt;

Dr. Azza Ahmed Shehata , Alexandria University, Faculty of Science, Botany Department, Alexandria, Egypt. Correspondent author: e-mail: [email protected] Manuscript received: July 4th, 2007.

Appendix 1

Basic data matrix of the 53 character states used in the phenetic analysis of Lathyrus seeds similarity. Num-bers correspond to the species in Table 1, plus the outgroup (35)

Character/Species 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 2 3 3 3 3 3 3 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5

Luster 1. glossy 2. matte 3. semi-matte

Multicellular sculpture: 4. colliculate – ocellate 5. colliculate – ruminate 6. colliculate – tuberculate 7. falsifoveate – colliculate 8. reticulate – colliculate 9. reticulate – favulariate 10. reticulate – foveate 11. reticulate monomorphic 12. ribbed – ruminate 13. ruminate

Homogeneity of cell-size: 14. uniform 15 . irregular

(GL) (MA) (SM)

(CL/OC) (CL/RU) (CL/TU) (FF/CL) (RT/CL) (RT/FV) (RT/FO) (RM) (RB/RU) (RU)

(UN) (IR)

0 0 0 1 0 0 0 1 1 1 0 0 1 1 0 1 0 1 1 1 1 1 1 1 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 1 1 1 0 0 0 1 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 1 0 0 0 0



Character/Species 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 2 3 3 3 3 3 3 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5

Anticlinal walls: Level: 16. highly raised 17. raised 18. leveled to sunken 19. irregular

Undulation: 20. wavy 21. straight 22. sinuate 23. slightly undulate

Thickness: 24. very thick 25. thick 26. thin 27. irregular

Secondary sculpture: 28. stellate structures 29. rodlike structures 30. buttressed walls

Periclinal walls: Level: 31. convex 32. slightly concave 33. concave

Secondary sculpture: 34. striate 35. ribbed to faveolariate 36. irregular to ribbed 37. papillate

Hilum-micropylar region (HMR): Shape: 38. oblong 39. ovate 40. elliptic

Length/width ratio: 41. ≤2 42. 2.1 : 2.5 43. 2.6 : 3.5 44. 3.6 : 4.5 45. 4.6 : 5.5 46. 5.6 : 6.5 47. >6.5

Seed length/width ratio: 48. 1.1 : 1.2 49. 1.21 : 1.3 50. 1.31 : 1.4 51. 1.41 : 1.5 52. 1.51 : 1.6 53. ≥1.6

(HR) (RA) (LV/SK) (IR)

(WA) (ST) (SN) (SU)

(VT) (TK) (TN) (IR)

(SL) (RO) (BU)

(CO) (SC) (CN)

(SI) (RB/FA) (IR/RB) (PA)

(OB) (OV)

(EL)

1 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 1 1 0 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 1 1 1 0 0 1 1 0 1 0 0 1 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 0 0 0 0 1 1 0 0 0 0 1 1 0 0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 1 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 1 1 1 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 0 0 0 0 1 1 1 1 0 0 1 1 1 1 1 0 0 0 1 1 1 1 1 1 1 0 0 0 0 1 0 0 0 0 0 1 1 1 0

0 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0

1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 1 1 1 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1

1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 1 1 0 1 1 1 0 0 0 0 0 1 0 0 1 0 1 0 0 0 1 1 1 1 0 0 0 1 0 1 0 0 0 0 0 0 0 1 0 0 0 1 1 1 1 1 0 1 1 0 1 0 1 1 1 0 0 0 0 1 1 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1

0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 1 0 1 0 1 0 0 0 0 0 1 1 1 1 1 1 0 0 1 0 0 0 1 1 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 1 1 0 0 1 0 1 0 1 1 1 0 1 0 1 0 1 0 1 0 1 1 1 1 0 0 0 0 0 0 0 1 1

0 1 0 0 1 1 1 0 0 1 0 1 1 0 0 1 1 0 1 0 1 0 0 0 0 1 1 0 1 0 0 0 0 1 0 0 0 1 0 0 0 0 0 1 0 1 0 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1

0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 1 1 1 1 0 1 1 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0

Documents

Seed surface characters and their systematic significance in the genus Lathyrus (Leguminosae, Papilionoideae, Vicieae)