Comparative Pollen Morphology and Ultrastructure of Mentheae Subtribe
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Available online at www.sciencedirect.com Review of Palaeobotany and Palynology 149 (2008) 174–186 www.elsevier.com/locate/revpalbo Comparative pollen morphology and ultrastructure of Mentheae subtribe Nepetinae (Lamiaceae) ⁎ Hye-Kyoung Moon a, , Stefan Vinckier a,b, Erik Smets a,c, Suzy Huysmans a a Laboratory of Plant Systematics, Institute of Botany and Microbiology, K.U. Leuven, Kasteelpark Arenberg 31, P.O. Box 2437, BE-3001 Leuven, Belgium b Center for Transgene Technology and Gene therapy, Flanders Institute for Biotechnology (VIB-3), K.U. Leuven, campus Gasthuisberg, Herestraat 49, BE-3000 Leuven, Belgium c National Herbarium of The Netherlands, Leiden University Branch, P.O. Box 9514, NL-2300 RA Leiden, The Netherlands Received 6 September 2007; received in revised form 22 November 2007; accepted 1 December 2007 Available online 15 December 2007 Abstract This study provides new pollen data of 52 representative species belonging to all 12 genera in the currently classification of the subtribe Nepetinae, and considers the possible presence of orbicules for the first time. Pollen morphology and ultrastructure were investigated with light, scanning electron and transmission electron microscopy. Nepetinae pollen is small to large (P=16–65 µm, E=17–53 µm), oblate to prolate (P/E=0.7–1.6) in shape and mostly hexacolpate (sometimes octocolpate). The exine stratification in all taxa studied is similar and characterized by unbranched columellae and a continuous, granular endexine. Sexine ornamentation in the Nepetinae is bireticulate, microreticulate or perforate. In perforate and microreticulate pattern a tendency towards a bireticulum could be recognized due to trace of secondary tectal connections. The bireticulate pattern is most common with variations of primary muri and secondary reticulum. In Hymenocrater and Schizonepeta the observed variation of sexine ornamentation is particularly valuable at the generic level. Pollen data support Lophanthus and Nepeta as very closely allied and Lallemantia is clearly distinct from Dracocephalum. The formerly suggested infrageneric relationships within Dracocephalum and Nepeta are only partly corroborated by palynological characters. Orbicules are absent in the Nepetinae. © 2007 Elsevier B.V. All rights reserved. Keywords: Nepetoideae; orbicules; palynology; Nepetinae; sexine ornamentation; ultrastructure 1. Introduction etically informative (Moon et al., in press). The present study aims to further elucidate the palynological diversity within the Pollen of the Lamiaceae has been intensively studied since tribe Mentheae, focusing on the subtribe Nepetinae sensu Erdtman (1945) suggested a division of the family into two Harley et al. (2004). The Nepetinae differs from the two other subfamilies based on aperture number and number of nuclei in subtribes in Mentheae, Salviinae and Menthinae in the length of the mature pollen grains (Lamioideae: tricolpate and bi-nucleate the posterior pair of stamens and the number of calyx nerves pollen, Nepetoideae: hexacolpate and tri-nucleate pollen). (Wagstaff, 1992). Nepetinae sensu Harley et al. (2004) consists However, the tribe Mentheae (Nepetoideae), which includes of 12 genera (ca. 350 species) distributed over large parts of nearly one fourth in all taxa within the Lamiaceae taxa, is still Eurasia and North America (Harley et al., 2004). poorly known from a palynological point of view. In an attempt Nepeta is the largest (ca. 200 sp.) and economically most to fill this knowledge gap, we have presented a detailed pollen important genus in the Nepetinae. It can be subdivided into study of subtribe Salviinae (Mentheae), showing that variations several sections and subsections by habit, leaf morphology, of sexine ornamentation in the pollen characters are phylogen- inflorescence, calyx and corolla structure, and nutlet characters (Bentham, 1848; Briquet, 1897; Pojarkova, 1954; Budantsev, 1993a; Budantsev and Lobova, 1997). Based on phylogenetic ⁎ Corresponding author. Tel.: +32 16 32 1535; fax: +32 16 32 1955. analysis of molecular data, Jamzad et al. (2003) concluded that E-mail address: [email protected] (H.-K. Moon). Nepeta is monophyletic but also that previous infrageneric 0034-6667/$ - see front matter © 2007 Elsevier B.V. All rights reserved. doi:10.1016/j.revpalbo.2007.12.001 H.-K. Moon et al. / Review of Palaeobotany and Palynology 149 (2008) 174–186 175 Table 1 Overview of palynological characters of species studied within Nepetinae based on acetolysed pollen grains Species T P (µm) E (µm) Shape AI CL (µm) Exine (µm) SE O So Os S Ps Sp P Agastache J.Clayton ex Gronov. A. palmeri A39–44.2–52 30–36.0–41 −− − −++++0.12–0.22 26–33.7–43 1.2–1.5 III-e A. scrophulariifolia A33–37.4–42 30–33.1–37 −− ± ± ++ + − 0.16–0.21 25–27.6–31 1.2–2.1 III A. urticifolia A32–37.6–41 26–27.6–29 −− − −− +++/27–29.1–34 1.1–1.2 III-e A. urticifolia A30–36.5–45 31–35.8–41 −− − −±+++0.12–0.15 24–30.4–37 1.0–1.4 III-e A. urticifolia C19–20.7–22 23–24.5–26 ++ ++−− − − 0.12–0.16 16–17.1–18 / III-e Cedronella Moench Cedronella canariensis A43–44.8–48 32–36.2–43 −− − −+ − ++ 0.14–0.23 30–32.8–35 0.9–1.8 III-d Dracocephalum L. D. fragile A49–52.2–57 35–38.6–44 −− − −− +++/35–40.3–45 1.1–1.3 I-a D. grandiflorum A47–54.6–60 40–44.0–52 −− − −±++ /38–41.7–45 1.0–1.3 I-a D. heterophyllum A33–37.5–42 28–32.9–41 −− + − ++ ++ ± 0.18–0.22 26–28.6–33 1.1–1.8 I-a D. imberbe A28–31.7–38 22–24.7–29 −− − − ±+++0.24–0.31 20–23.5–28 1.0–1.4 I-a D. komarovii A35–39.1–41 32–35.0–42 −− ± − ++ ++ − 0.12–0.14 27–29.8–32 0.9–1.2 I-a D. nutans A34–39.4–42 30–33.5–37 −− ± ± ++ ++ ± 0.10–0.14 27–31.3–35 1.0–1.4 III-d D. palmatum A42–49.1–61 38–45.0–53 −− + − ++ + − 0.10–0.17 34–37.5–45 1.0–1.3 III-d D. parviflorum A39–41.6–45 30–35.2–38 −− − −±++− /27–31.2–35 1.0–1.5 I-b D. pinnatum A39–42.0–45 33–38.2–45 −− ± ± ++ + − 0.08–0.15 28–33.1–38 1.1–1.6 I-a D. ruyschiana A40–42.4–45 30–33.4–37 −− − −±++±/30–32.9–35 1.1–1.5 I-b D. ruyschiana C29–30.4–31 24–28.6–34 −− + − ++ + − /24–26.4–29 / I-b D. scrobiculatum A41–49.2–55 31–35.4–43 −− − −− ++ ++ /29–36.7–42 1.0–1.5 I-b D. stamineum A31–35.1–39 29–31.7–34 −− ± − ++ ++ ± 0.08–0.16 25–28.3–32 0.9–1.0 III-a D. subcapitatum A32–37.8–46 29–37.1–45 −− ± − +++±0.16–0.25 28–32.7–38 0.9–1.1 I-b Drephanocaryum Pojark. Drepanocaryum sewerzowii A25–27.9–30 27–29.5–31 − ±++±+−−0.12–0.23 20–23.6–26 1.0–1.5 III-b Drepanocaryum sewerzowii C16–16.8–17 20–21.5–23 − +++−− − − /12–12.7–13 / III-b Glechoma L. Glechoma hederacea A35–37.5–44 30–35.2–46 − ±++− + −−0.12–0.20 30–34.1–39 0.9–1.2 I-a Glechoma hederacea C22–25.2–30 17–23.5–28 −− ± −− ++/17–21.0–25 / I-a Hymenocrater Fisch. & C.A.Mey. Hymenocrater bituminosus C27–29.6–32 26–28.4–30 −− ± − +++− 0.11–0.16 22–23.2–25 / I-c Hymenocrater longiflorus A49–53.1–59 40–44.2–50 −− ± − +++±0.11–0.18 40–44.1–49 1.5–1.7 I-c Lallemantia Fisch. & C.A.Mey. Lallemantia peltata A26–27.7–30 24–27.1–29 −− ++ − ++ ± − /21–22.9–25 1.1–1.4 I-b Lallemantia peltata C16–18.0–19 22–23.3–24 +++−−−−−/14–15.4–16 / I-b Lallemantia royleana A26–28.1–31 23–27.1–29 −− ++ + ++ + − 0.17–0.21 22–23.6–27 0.9–1.3 III-e Lallemantia royleana C16–17.3–19 20–20.5–21 ±+ ++−− − − 0.14–0.23 11–13.7–16 / III-e Lophanthus Adans. Lophanthus chinensis A35–41.4–51 30–35.4–40 −− − −+++±0.12–0.23 30–34.6–38 1.0–1.3 III-c Lophanthus laxiflorus A26–28.4–32 25–28.4–34 − ±+++±− 0.19–0.24 20–21.4–25 0.8–1.1 III-e Lophanthus tschimganicus A31–34.6–38 29–33.0–39 −− ++ ± + + − 0.14–0.17 24–28.1–31 1.5–2.1 III-b Lophanthus tschimganicus C21–24.3–26 17–19.2–21 −− − −++++/18–20.9–23 / III-b Marmoritis Benth. Marmoritis rotundifolia C28–30.6–36 30–31.3–33 −− ++ − ++− 0.13–0.18 23–26.6–31 / III-e Meehania Britton Meehania urticifolia A51–57.8–65 38–44.1–53 −− − −±+++/40–45.9–55 1.2–1.5 III-e Meehania urticifolia C35–42.9–51 21–32.7–37 −− − −− ++ + /25–34.4–42 III-e Nepeta L. N. assurgens A25–28.9–33 25–27.5–30 − ±±±+++− 0.14–0.22 18–20.5–25 0.9–1.1 III-e N. cataria A27–29.4–32 26–29.5–33 − ±++++±− 0.12–0.16 20–21.9–24 0.9–1.0 III-e N. cataria C21–23.2–25 15–20.9–27 − ++ −−−++/17–19.5–22 / III-e N. cephalote C18–21.0–25 22–23.0–25 − ++ + −− − − /16–18.2–20 / III-e (continued on next page) 176 H.-K.