Abstract
We investigated fruit and seed morphology in 48 taxa of 22 genera in the tribe Rhinantheae s.l. and related genera of Orobanchaceae. Fruit and seed morphology are heterogeneous across the studied taxa. The most useful features for this study are the indumentum of the capsule, seed number per capsule, seed coat ornamentation, and secondary structure of the seed coat. On the basis of seed coat features, five major types are recognized, viz. reticulate, cristate-winged, sulcate, psilate and irregularly striate. In addition, phenetic analysis of 19 characters produced an unweighted pair-group method using arithmetic averages tree and displayed highly heterogeneity at infra- and inter-genera. The classification of Rhinantheae s.l. is discussed and keys to the genera considered in this work, based exclusively on fruit or seed characters, are provided.
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Acknowledgments
The authors thank the herbaria at the Kunming Institute of Botany (KUN), and Institute of Botany (PE), the Chinese Academy of Sciences, and California Academy of Sciences for allowing us to collect study materials. We are grateful to X. K. Fan for SEM technical help, H.T. Li and A. Shah for data analyses, and R. Turkington and I. Taylor for reading earlier drafts of this manuscript. This study is supported by Grants from the National Natural Science Foundation of China (No. 30970201, No. 30570115), National Key Basic Research Program of China(No. 2014CB954100), and Key Research Program of the Chinese Academy of Sciences (No. KJZD-EW-L07).
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Appendices
Appendix 1
See Table 2.
Appendix 2
See Table 3.
Appendix 3: Descriptions of characters and character states
Capsule:
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(1)
Shape. Two character states are described: ovoid (0) and oblong (1). Some capsules are narrowly or broadly ovoid, these are included in ovoid (0) as a qualitative description.
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(2)
Color. Two character states are described: yellow (0) and brown (1).
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(3)
Surface. Three character states are described: glabrous (0), eglandular hairs (1), and aculeate (2). Glabrous indicates no extra structures on the surface. Eglandular hairs are the most characteristic trait, and the hairs usually display an enlarged base. Aculeate indicates triangular protruberances covered by claviform tips.
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(4)
Type of indumentum. Three character states are described: pilose (0), strigose (1), and branched hairs (2). Pilose indicates long and erect hairs. Strigose indicates short and inclined hairs. Branched hair indicates several hairs on a single stem.
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(5)
Texture. Two character states are described: coriaceous (0) and papyraceous (1). Coriaceous is defined as tough and pliable, papyraceous as soft and fragile.
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(6)
Apex. Three character states are described: acuminate (0), cuspidate (1), and emarginate (2). Acuminate indicates a sharp point at the apex. Cuspidate indicates a narrowing to a point. Emarginated indicates a notch at the apex.
Seed:
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(7)
Size. Four character states are described: dust-like (0), small (1), medium (2), and large (3). These character states are divided based on the scatter of the mean seed length.
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(8)
Number. Three character states are described: several (0), many (1), and numerous (2). Several are defined as fewer than 10 seeds per capsule. Numerous is defined as above 100 seeds. Many are defined as between 10 and 100 seeds.
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(9)
Lateral shape. Two character states are described: concave (0) and flattened (1). Concave indicates depressed on the seed surface to varying degrees. Flattened displays a smooth plane on the surface.
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(10)
Wing. Two character states are described: unwinged (0) and winged (1). The seed has a membranous protrusion along its longitude (1) or not (0).
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(11)
Wing number. Two character states are described: one (0) and several (1).
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(12)
Surface projections. Three character states are described: smooth (0), spinulose (1), and tuberculate (2). Smooth indicates no projections on the seed surface. Spinulose indicates tiny and protruding pinpoints. Tuberculate indicates small rounded protuberances.
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(13)
Testa. Two character states are described: tightly-fitting (0) and loosely-fitting (1). The testa of the seed is separated from the inner structure when mature (1) or not (0).
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(14)
Primary ornamentation. Five character states are described: reticulate (0), cristate-winged (1), sulcate (2), psilate (3), and irregularly striate (4). Reticulate indicates a network-like pattern. Cristate-winged indicates several protrusions along longitude of epidermal cells. Sulcate indicates paralleled lineation through the seed surface. Psilate indicates smooth surface. Irregularly striate indicates short and disorderly lines on the surface.
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(15)
Adjacent ridges of reticulum region. Two character states are described: trough (0) and ridge (1). For reticulate seeds, there may be a slight sunken region (0) or raised region (1) between the adjacent ridges of the reticulum.
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(16)
Reticulum lines. Three character states are described: obscure (0), normal (1), and deep (2). Obscure indicates blurred lines in the reticulum. Normal indicates clear lines to the reticulum. Deep indicates strongly uplifted lines in the reticulum.
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(17)
Reticulum line height. Two character states are described: uniform (0) and uneven (1). The lines of the reticulum are lengthened equally (0) or not (1).
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(18)
Mesh shape. Three character states are described: narrowly or shortly rectangular (0), isodiametrically polygonal (1), and irregularly polygonal (2).
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(19)
Secondary ornamentation. Four character states are described: smooth (0), reticulate (1), lineate (2) poroid-alveolate (3), and hook-like (4). Frequently, the seeds are reinforced with secondary thickening of various types on the epidermal cell wall.
Appendix 4
See Table 4.
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Dong, LN., Wang, H., Wortley, A.H. et al. Fruit and seed morphology in some representative genera of tribe Rhinantheae sensu lato (Orobanchaceae) and related taxa. Plant Syst Evol 301, 479–500 (2015). https://doi.org/10.1007/s00606-014-1087-8
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DOI: https://doi.org/10.1007/s00606-014-1087-8