Acknowledgements

This study was supported financially by the project 'Studies on the Establishment of SeedBank base for the Asian network' at the Korea National Arboretum (KNA), Pocheon, Korea.


Appendix

Instead, Scrophularia are characterized by perennial herbs without rhizomes, five corolla lobes,

As predicted, three Veronica species (V. arvensis, V. didyma var. lilacina, and V. persica) form an isolated group. Based on the infrageneric classification as done in [47], V. arvensis (with haploid chromosome, n = 8) belongs to subgenus Chamaedrys and V. didyma var. lilacina (as V. polita) and V. persica, both having base chromosome n = 7, belong to Pocilla. V. linariifolia, V. longifolia, V. dahurica, and V. incana, all have the chromosome number n = 17, and the reticulate-striate, cristate group belongs to Pseudolysimachium, whereas V. peregrina (n = 9) belongs to subgenus Beccabunga. The present results confirmed what was reported in previous studies on Veronica (for example, [32, 34]), that seed morphological data can be employed to evaluate the relationships of taxa within the genus and that seed characters can also provide

The SEM investigations reveal that the reticulate surface pattern as a primary sculpture is a confusing case. The pattern is found in the members of Orobanchaceae, Plantaginaceae, and Scrophulariaceae, although quite variations are observed in secondary striation and anticlinal wall pattern. The current study likewise concurs that there is no even single seed features representing all investigated species. Nonetheless, few seed characters somehow nicely characterized the group of particular taxa, for instance, seeds of Orobanchaceae are either elliptical cylindrical or ovoid in shape and of Plantaginaceae are elliptical flattened or ovoid flattened. Correspondingly, reticulate-striate surface sculpture of Veronica, Veronicastrum, and Scrophularia is comparable, although later one is without cristae. The seed coat anatomy indicates that the epidermis and endothelium are informative characters. All the Scrophularia species have indistinct epidermis but distinct endothelium, whereas most of the Veronica has distinct epidermis and endothelium. On the other hand, all the Melampyrum are without both layers, but other

The concept of Scrophulariaceae s.l. has changed considerably since the application of molecular approaches in plant systematics. Studies have shown that the traditional Scrophulariaceae are an unusual assemblage of plants distributed throughout the phylogenetic tree of Lamiales [1, 2, 5–7]. Despite the limited number of taxa investigated, the present seed morphological analysis highlights the high heterogeneity existing within the studied taxa of Scrophulariaceae s.l., which may be observed at a higher taxonomic rank than genus. In particular, primary surface sculpture, anticlinal and periclinal walls of epidermal cell, epidermal cell shape, and seed coat layers are the important seed features observed in this study, and these features can be used to distinguish different groups of taxa in the family. The results of this study therefore suggest that seed coat micromorphology and anatomy

This study was supported financially by the project 'Studies on the Establishment of SeedBank

base for the Asian network' at the Korea National Arboretum (KNA), Pocheon, Korea.

four didynamous stamens, and bifid stigmas.

222 Advances in Seed Biology

additional support to infrageneric discrimination in Veronica.

Plantaginaceae have well-distinct epidermis and endothelium.

have significant taxonomic importance.

Acknowledgements



Plant name

V. kiusiana var. glabrifolia

V. linariifolia

V. longifolia

V. nakaiana V. peregrina

V. persica V. pusanensis

V. pyrethrina V. rotunda var.

V. undulata Veronicastrum

Linaria japonica

Scrophularia

S. grayana S. kakudensis.

S. koraiensis S. takesimensis

Epidermal

Rectangular,

Irregular Rectangular,

Irregular Irregular Irregular Irregular

Convex, smooth to finely folded

Convex, smooth to finely folded

Convex or pitted, smooth to finely folded Shallow, smooth, unevenly thickened

 Polygonal

 elongated

 cell shape

Periclinal wall Concave, finely granulate Concave, finely granulate

Convex, finely folded

Concave, smooth to folded

Anticlinal wall Slightly raised, straight, finely granulated

Slightly raised, folded

Shallow, finely folded

Distinctly raised, straight to wavy, finely

folded, unevenly thickened

Shallow or slightly raised

Shallow

Seed coat Epidermis

 Distinct Distinct

Distinct

Distinct

Indistinct

Indistinct

 Indistinct

Degenerated

 Absent

Degenerated

 Absent

Degenerated

 Absent

 Well represented

 Present

 Well represented

 Present

 Well represented

 Absent

 Well represented

 Absent

Endothelium

 buergeriana

 sibiricum

subintegra

Broad elliptical, flattened

Broad ovoid,

Ovoid, Ovoid, flattened

Ovoid Ovoid Ovoid Ovoid Ovoid

plano-convex

 0.73 0.46 2.03 1.68 10.3 0.71 1.08 0.76 1.13 0.73

1.2 0.74 1.01 0.6

plano-convex

 0.62 0.40

Shape Broad elliptical, flattened

Broad elliptical, flattened

Broad elliptical, flattened

Broad ovoid, flattened

Elliptical, flattened

Ovoid, flattened

Broad elliptical, flattened

Broad elliptical, flattened

Average size (mm)

 1.17 1.01

 1.19 0.85

 1.19 0.99

 0.94 0.74

 0.71 0.41 1.17 0.79

 1.01 0.79

 0.95 0.78

 0.99 0.77

 1.16

 1.4

 1.54

 1.28

 1.73

 1.49

 1.29

 1.23

 1.29

 1.56

 1.61

 1.68

 1.47

 1.45

 1.56 1.64 1.72

Lateral, distinctly protuberant

Basal, distinctly protuberant Lateral, distinctly protuberant

Basal, distinctly protuberant Basal, distinctly protuberant Basal, distinctly protuberant Lateral, distinctly protuberant Lateral, distinctly protuberant Lateral, distinctly protuberant Lateral, distinctly protuberant Lateral, distinctly protuberant

Basal, deep Basal, deep Basal, deep Basal, deep Basal, deep Basal, deep

Length/width

 ratio Hilum

Primary surface sculpture

Reticulate-striate,

Reticulate-striate,

Reticulate-striate,

Reticulate-striate,

Faintly reticulate

Reticulate-verrucate

Colliculate

Reticulate-striate

Reticulate-striate

Reticulate-striate

Reticulate-striate

Reticulate-striate

Reticulate-striate,

 cristate

Reticulate-corrugate

Reticulate-striate,

 cristate

Reticulate-striate,

 cristate

Reticulate-striate,

 cristate

 cristate

 cristate

 cristate

 cristate

224 Advances in Seed Biology


Appendix 1. Seed characters studied for Scrophulariaceae s.l.

Morphological Studies on Seeds of Scrophulariaceae s.l. and Their Systematic Significance http://dx.doi.org/10.5772/intechopen.70572 227


Epidermal

Polygonal, Polygonal, elongated

Polygonal, Polygonal, Polygonal, Polygonal, elongated Polygonal, elongated

Polygonal, Polygonal, elongated

Polygonal, Polygonal, elongated Polygonal, elongated Polygonal, elongated Polygonal, elongated Polygonal, elongated Appendix 1. Seed characters studied for

Scrophulariaceae

 s.l.

Concave, striate

Concave, striate

Concave, striate

Concave, striate

isodiametric

Convex, finely folded

Concave, striate

Concave, striate

isodiametric

Concave, striate

Concave, striate

Concave, striate

elongated/isodiametric

 Concave, striate

isodiametric

Convex with central tubercle, smooth

elongated/isodiametric

 Flat,

smooth/folded

Concave, striate

elongated/isodiametric

 Concave, striate

 cell shape

Periclinal wall

Anticlinal wall Distinctly raised, straight to wavy,

folded, unevenly thickened

Distinctly raised, straight to wavy,

folded, unevenly thickened

Slightly raised, folded, unevenly

thickened

 Slightly raised, folded, unevenly

thickened

Distinctly raised, straight to wavy,

folded, unevenly thickened

Distinctly raised, straight to wavy,

folded, unevenly thickened

Distinctly raised, straight to wavy,

folded, unevenly thickened

Distinctly raised, folded, unevenly

thickened

Distinctly raised, straight to wavy,

folded, unevenly thickened

Shallow, finely folded, straight to wavy

Distinctly raised, straight to sinuous,

unevenly thickened

Distinctly raised, straight to sinuous,

unevenly thickened

Distinctly raised, straight to sinuous,

unevenly thickened

Distinctly raised, straight to sinuous,

unevenly thickened

Distinctly raised, straight to sinuous,

unevenly thickened

 Distinct Distinct

Distinct

Distinct

Distinct

Distinct

Degenerated

 Present

Degenerated

 Present

Degenerated

 Present

Degenerated

 Present

Degenerated

 Present

 Well represented

 Present

Seed coat Epidermis

Distinct

Distinct

Distinct

Indistinct

Distinct

Distinct

Distinct

Distinct

Distinct

 Well represented

 Present

 Well represented

 Present

 Well represented

 Present

 Well represented

 Present

 Well represented

 Present

Degenerated

 Absent

 Well represented

 Present

 Well represented

 Present

 Well represented

 Present

226 Advances in Seed Biology

Endothelium


Appendix 2. Data matrix of the character states used for analysis.
