1
RESEARCH ARTICLE
Seed micromorphology and its taxonomic evidence in
subfamily Alsinoideae (Caryophyllaceae)
Fazal Ullah
1,2,3
| Alessio Papini
4
| Syed Nasar Shah
1
| Wajid Zaman
3,5
|
Aamir Sohail
6
| Majid Iqbal
1
1
Department of Plant sciences, Quaid-i-Azam
University, Islamabad, Pakistan
2
CAS Key Laboratory of Mountain Ecological
Restoration and Bioresource Utilization &
Ecological Restoration and Biodiversity
Conservation Key Laboratory of Sichuan
Province, Chengdu Institute of Biology,
Chengdu, Sichuan, China
3
University of Chinese Academy of Sciences,
Beijing, China
4
Department of Biology, University of
Florence, Florence, Italy
5
State Key Laboratory of Systematic and
Evolutionary Botany, Institute of Botany,
Beijing, China
6
Department of Botany, University of
Peshawar, Peshawar, Pakistan
Correspondence
Fazal Ullah, Department of Plant sciences,
Quaid-i-Azam University, Islamabad, 45320,
Pakistan.
Email: fazalbotanist@gmail.com
OR
Alessio Papini, Department of Biology
University of Florence, Italy.
Email: alessio.papini@unifi.it
Funding information
University of Peshawar
Review Editor: George Perry
Abstract
Seed micromorphology of 13 species, belonging to four genera of subfamily Alsinoideae
(Caryophyllaceae) were investigated with scanning electron microscopy (SEM), in order to assess
their diagnostic significance at generic level and provide additional evidence on species delimita-
tion, as well as correct identification and phylogenetic position. Genera and species of subfamily
Alsinoideae exhibit great variation in ultrastructure and a high diversity of novel micromorphologi-
cal characters were observed. Variation in seed shape, color, hilum, anticlinal wall, epidermal cell,
cell surface, margins, and quantitative characters as length and width were studied in detail, com-
pared, illustrated, and their taxonomic significant were discussed. Seed shapes of the species were
classified as reniform, round, angular, subcircular, subreniform, and elliptical pyriform, with sub-
central, central, basal, and nearly basal hilum. Wavy, irregular, tetragonal, and elongated epidermal
cells structure has been observed as an exomorphological character. The present findings show
that the micromorphology of subfamily Alsinoideae provides taxonomic information and is helpful
to distinguish different species. The results also explained that SEM morphology of seeds provide
important data about affinity among taxa and give potential characters in delimitation of members
of subfamily Alsinoideae at generic and species level. A principal component analysis allowed to
highlight the most outsiders among seed micromorphology with a possible explanation. Taxo-
nomic keys were developed based on micromorphological characters to delimit the species and
useful for their quick identification within subfamily Alsinoideae.
KEYWORDS
Alsinoideae, hilum, seed morphology, seed surface, SEM
1 | INTRODUCTION
Caryophyllaceae is one of the largest dicot family consisting of three
subfamilies, that is, Alsinoideae, Caryophylloideae, and Paronychioideae
and 11 tribes (Bittrich, 1993), with 88 genera and 2,200 species
(Harbaugh et al., 2010). In Flora of Pakistan the family is represented by
110 species and 26 genera distributed throughout the country
(Perveen & Qaiser, 2006; Ullah, Zafar, Ahmad, Dilbar, et al., 2018). The
family is primarily distributed in the Holarctic but with rich diversity in
Irano-turanian and Mediterranean regions (Hernández-Ledesma et al.,
2015), most of the species grow in relatively moist habitats and temper-
ate forest of the world (Vandelook, Van de Moer, & Van Assche, 2008).
Members of the family as Stellaria media (L.) Vill. also grow as important
weeds among several crops (Ullah, Ullah, & Sohail, 2016). The dominant
features of the subfamily Alsinoideae are the presence of nectar glands
on the base of abaxial side of the episepalous stamens and they can be
distinguished from Caryophylloideae by tubular calyx and clawed petals
(Bittrich, 1993; Harbaugh et al., 2010; Ullah, Zafar, Ahmad, Dilbar, et al.,
2018; Ullah, Zafar, Amhad, Shah, et al., 2018).
According to the taxonomic classification by McNeill (1980) in
subfamily Alsinoideae, Cerastium L., and the Stellaria-Cerastium com-
plex are included, together with Myosoton Moench, Moenchia Ehrh.,
Holosteum L., and Pseudostellaria Pax., as members of tribe Alsineae
Lam & Dc. Genus Cerastium is traditionally divided into two
Received: 19 September 2018 Accepted: 9 October 2018
DOI: 10.1002/jemt.23167
Microsc Res Tech. 2018;1–10. wileyonlinelibrary.com/journal/jemt © 2018 Wiley Periodicals, Inc. 1