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. 2025 Sep 2;262:105–128. doi: 10.3897/phytokeys.262.152205

Gap analysis of knowledge about the microstructure of Impatiens Riv. ex L. (Balsaminaceae) seeds using SEM techniques

Agnieszka Rewicz 1,✉, René Monzalvo 2, Kayla Grey 2, Katarzyna Sanek 1, Maja Mackiewicz 1,3, Monika Myśliwy 4
PMCID: PMC12421252  PMID: 40937399

Abstract

We conducted a literature review on the taxonomic importance of seed micromorphology in the genus Impatiens Riv. ex L. (Balsaminaceae), focusing on studies using scanning electron microscopy (SEM). Of 47 sources, SEM descriptions were available for more than 180 species, revealing significant gaps in micromorphological data within the genus. Our work provides a consolidated resource on Impatiens taxonomy and highlights the need for further SEM studies to improve species identification and phylogenetic analysis.

Key words: Micromorphological analysis, plant taxonomy, seed coat, systematic studies, taxonomic classification

Introduction

Owing to differences in the understanding of taxonomic features across different groups of plants, we can more accurately assess the usefulness and level of conservatism of specific taxonomic traits within plant groups. Undoubtedly, among the most important taxonomic features are those related to flowers (e.g. in Ranunculaceae, Asteraceae families) (Hoots 1991; Karanović et al. 2015; Zhai et al. 2019), and examples of their use can be found at all levels of the taxonomic hierarchy. Conversely, seed and fruit characters are less prevalent and indeed possess lesser taxonomic significance. However, there are botanical families where carpological research is the most useful taxonomic feature (e.g. in Caryophyllaceae or Brassicaceae families) (Abdel-Maksoud and Fawzi 2016; Ullah et al. 2019; Gabr 2018; Hashem AL-Blesh et al. 2021). An example is Caryophyllaceae, whose features such as tubercle shape, size, and distribution provide key diagnostic characters (Martín-Gómez et al. 2024a; 2024b). Unusual forms and arrangements include echinate, rugose, mammillate, papillose, and smooth seed tubercles, as well as epidermal cell wall patterns such as digitate, serrate, sinuous, and rectangular (Hubinský et al. 2024). But a crucial aspect lies in the extent of research into the micromorphology of seeds and fruits. Historically, descriptions of seeds and fruits were based on observations through light microscopes or binoculars, often failing to highlight interesting features of their structure, hence their role remained underappreciated by botanists (Hooker 1904–1906, 1909; Tardieu-Blot 1945–1946; Perrier de la Bathie 1948). Another challenge in carpological research was the limited availability of material. Botanists, typically emphasizing species descriptions, often focused on flowering specimens, and seeds were not always available on these specimens. It should also be remembered that only mature seeds or fruits can be the basis for correct taxonomic identification.

Thanks to progress in microscopy techniques, particularly the utilization of SEM (scanning electron microscope), research on seed and fruit cover has become more common, offering new perspectives and approaches to establishing boundaries between organisms (Brisson and Peterson 1976; Godfray and Knapp 2004). The first carpological research using this technology took place in the mid-1960s, but the real development of the use of SEM in botany, including carpology, occurred only in the 1990s (Barthlott 1981, 1984). This, in turn, resulted in the discovery of unusual forms, shapes and arrangements in known seed structures (e. g., net, reticulated and papillated-like patterns in the primary and secondary sculpture of cell walls) (Barthlott 1981) drawing attention to the fact that these features could be of significant importance in plant taxonomy (Heywood 1969; Stace 1992). Currently, an integrated approach to taxonomy can contribute to solving difficult or complicated cases (Ruchisansakun et al. 2015; Yu et al. 2015; Rahelivololona et al. 2018). Literature data indicate that seed and fruit characteristics serve as valuable tools at various taxonomic levels including families, e.g. Brassicaceae, Apiaceae, Rosaceae, and Caryophyllaceae (Crow 1979; Abdel-Maksoud and Fawzi 2016), genera, e.g. Silene (Fawzi et al. 2010; Atazadeh et al. 2017), Dianthus, Spergula, and Spergularia (Gvinianidze and Fedotova 1991), and even subspecies such as in Montia fontana, where they are solely distinguished based on seed morphology (Heywood 1974).

However, to ascertain the significance of seeds and fruits as a valuable tool in the taxonomy of a given group, thorough knowledge and description of these features are imperative. Therefore, the contemporary challenge in carpology lies not only in filling the knowledge gaps concerning various plant groups, but also in specifying how seed images are used in taxonomy. Both light microscopy and scanning electron microscopy (SEM) play an essential role in this context. Standard morphometric analyses typically include measurements of seed length, width, surface area, perimeter, and length-to-width ratio (aspect ratio), providing a fundamental quantitative basis for comparing taxa (Zhai et al. 2019; Martín-Gómez et al. 2024a). Moreover, seed shape analysis – increasingly performed using geometric morphometrics and elliptic Fourier analysis – opens new avenues for taxonomic and evolutionary studies, enabling objective comparisons across large datasets (Martín-Gómez et al. 2023). In addition to these parameters, SEM images allow for the detailed observation of seed coat cell arrangement and shape, as well as the morphology of periclinal and anticlinal cell walls, which serve as valuable diagnostic characters in species identification and classification (Rewicz et al. 2022c). In this work, our focus is on the genus Impatiens, which has been extensively studied in terms of systematics, ecology, and morphology. Impatiens is distinguished by its enormous species richness and extremely variable flowers, which makes it one of the most difficult groups to classify (Song et al. 2005; Cai et al. 2007; Janssens et al. 2010; Bhaskar 2012; Gogoi et al. 2018). New species are constantly being identified in various regions of the world (Fujimoto et al. 2024; Paing et al. 2024; Song et al. 2024), and research on them emphasizes the importance of seed micromorphology as an important feature in the process of species identification (Shimizu 1979; Lu and Chen 1991). Moreover, the ultrastructure of seeds can provide information about the phylogenetic relationships between them (Song et al. 2003, 2005; Yuan et al. 2004). It has been demonstrated that seed shape and seed coat micromorphology are effective tools for resolving taxonomic issues, including within the Balsaminaceae family, to which the genus Impatiens belongs (cf. Utami and Shimizu 2005; Chen et al. 2007; Cai et al. 2013; Zhang et al. 2016). Detailed morphometric analyses commonly involve measurements of seed length and width. In addition, scanning electron microscopy is employed to examine the ultrastructure of the testa, with particular attention to diagnostic characters such as the shape and arrangement of epidermal cells, the nature of anticlinal and periclinal walls, and specific surface ornamentation patterns. According to Zhang et al. (2016), features like reticulate or striate ornamentation, variations in cell wall elevation, and granule presence on the testa surface can be critical in differentiating species and understanding their phylogenetic relationships. Despite this, there is still much to be done, as the micromorphology of the seed coat has been studied using SEM in less than one fifth of Impatiens species (Maciejewska-Rutkowska and Janczak 2016). Literature data have shown that descriptions of Impatiens spp. seeds are typically brief, accompanied by microscopic observations, and sometimes supplemented with measurements, photographs or diagrams (Suppl. material 1). However, the absence of crucial details, such as measurements or SEM images, poses challenges for comprehensive analysis (Abid et al. 2011). In this study, we analyzed the gaps in available literature concerning the micromorphology of seeds in the genus Impatiens using SEM. The aim of our work was to develop the current state of knowledge on this issue, thus providing researchers with a comprehensive literature review on the ultrastructure of seeds.

Material and methods

A complete list of species of the genus Impatiens that have studies on seed carpology was prepared on the basis of available literature data, including information published in scientific articles and data from “gray literature”, such as local monographs and technical reports (Table 1). Our primary inclusion criteria comprised studies exclusively involving SEM analyses that incorporate SEM images. Subsequently, we gathered data on the missing species (species with no SEM images) considering the following variables: a) morphological description of the seeds, imagery including microscope and binocular photos, drawings and graphics and b) the geographical origin of the analyzed material (country) (Supp. material 1).

Table 1.

List of species of the genus Impatiens with SEM images of seeds.

Number Species name Source Remarks
1 Impatiens acaulis Arn. Utami and Shimizu 2005 accepted by WFO (placed as the accepted name of a taxon)
2 Impatiens aconitoides Y.M.Shui & W.H.Chen Shui et al. 2011 accepted by WFO (placed as the accepted name of a taxon)
3 Impatiens alboflava Miq. Utami and Shimizu 2005 accepted by WFO (placed as the accepted name of a taxon)
4 Impatiens alpicola Y.L.Chen & Y.Q.Lu Lu and Chen 1991 accepted by WFO (placed as the accepted name of a taxon)
5 Impatiens anjawensis Borah, Kandwal, Chhetri & Gogoi Rewicz et al. 2020a accepted by WFO (placed as the accepted name of a taxon)
6 Impatiens apalophylla Hook.f. Yu et al. 2015 accepted by WFO (placed as the accepted name of a taxon)
7 Impatiens apsotis Hook.f. Song et al. 2005 accepted by WFO (placed as the accepted name of a taxon)
8 Impatiens aquatilis Hook.f. Song et al. 2005 accepted by WFO (placed as the accepted name of a taxon)
9 Impatiens aquatilis Hook.f. Zhang et al. 2016 accepted by WFO (placed as the accepted name of a taxon)
10 Impatiens aquatilis Hook.f. Xia et al. 2019 accepted by WFO (placed as the accepted name of a taxon)
11 Impatiens arguta Hook.f. & Thomson Cai et al. 2013 accepted by WFO (placed as the accepted name of a taxon)
12 Impatiens arguta Hook.f. & Thomson Song et al. 2005 accepted by WFO (placed as the accepted name of a taxon)
13 Impatiens arriensii (alliensii) (Zoll.) T. Shimizu Shimizu 1987 accepted by WFO (placed as the accepted name of a taxon)
14 Impatiens arriensii (Zoll.) T.Shimizu Utami and Shimizu 1995 accepted by WFO (placed as the accepted name of a taxon)
15 Impatiens aureliana Hook.f. Cai et al. 2013 synonym of Impatiens violiflora Hook.f.
16 Impatiens aureliana Hook.f. Song et al. 2005 synonym of Impatiens violiflora Hook.f.
17 Impatiens badrinathii Pusalkar & D.K.Singh Pusalkar and Singh 2010 synonym of Impatiens chungtienensis Y.L.Chen
18 Impatiens bahanensis Hand.- Mazz. Zhang et al. 2016 accepted by WFO (placed as the accepted name of a taxon)
19 Impatiens bahanensis Hand.-Mazz. Song et al. 2005 accepted by WFO (placed as the accepted name of a taxon)
20 ? Impatiens baishaensis B.Ding & H. P. Deng Ding et al. 2017 not found on WFO
21 Impatiens balfourii Hook. f. Najberek et al. 2020 accepted by WFO (placed as the accepted name of a taxon)
22 Impatiens balfourii Hook. f. Song et al. 2005 accepted by WFO (placed as the accepted name of a taxon)
23 Impatiens balsamina L. Song et al. 2005 accepted by WFO (placed as the accepted name of a taxon)
24 Impatiens balsamina L. Utami and Shimizu 2005 accepted by WFO (placed as the accepted name of a taxon)
25 Impatiens balsamina L. Chen et al. 2007 accepted by WFO (placed as the accepted name of a taxon)
26 Impatiens barbata Comber Yu et al. 2015 accepted by WFO (placed as the accepted name of a taxon)
27 Impatiens begoniifolia S.Akiyama & H.Ohba Song et al. 2005 accepted by WFO (placed as the accepted name of a taxon)
28 Impatiens benthamii Steenis Utami and Shimizu 2005 accepted by WFO (placed as the accepted name of a taxon)
29 Impatiens bicolor subsp. Bicolor Abid et al. 2011 synonym of Impatiens bicolor Royle
30 Impatiens bicolor Royle subsp. pseudo-bicolor (Grey-Wilson) Y.J.Nasir Abid et al. 2011 synonym of Impatiens bicolor Royle
31 Impatiens bicornis L.Joseph & Bhaskar Bhaskar 2012 accepted by WFO (placed as the accepted name of a taxon)
32 Impatiens blepharosephala E.Pritz. Cai et al. 2013 accepted by WFO (placed as the accepted name of a taxon)
33 Impatiens blinii H.Lév. Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
34 Impatiens bodinieri Hook.f. Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
35 Impatiens brachycentra Kar. & Kir. Zhang et al. 2016 accepted by WFO (placed as the accepted name of a taxon)
36 ? Impatiens brachycentra Kar. & Kir. var. jacquemontii (Hook.f.) Abid et al. 2011 not found on WFO
37 Impatiens burtonii Hook.f. Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
38 Impatiens capensis Meerb. Rewicz et al. 2020b accepted by WFO (placed as the accepted name of a taxon)
39 Impatiens chekiangensis Y.L.Chen Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
40 Impatiens chimiliensis Comber Cai et al. 2013 accepted by WFO (placed as the accepted name of a taxon)
41 Impatiens chinensis L. Chen et al. 2007 accepted by WFO (placed as the accepted name of a taxon)
42 Impatiens chinensis L. Song et al. 2005 accepted by WFO (placed as the accepted name of a taxon)
43 Impatiens chinensis L. Utami and Shimizu 2005 accepted by WFO (placed as the accepted name of a taxon)
44 Impatiens chloroxantha Y.L.Chen Wang et al. 2020 accepted by WFO (placed as the accepted name of a taxon)
45 Impatiens chungtienensis Y.L.Chen Song et al. 2005 accepted by WFO (placed as the accepted name of a taxon)
46 Impatiens clavigera Hook.f. Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
47 Impatiens clavigera var. auriculata S.H.Huang Lei et al. 2015 synonym of Impatiens apalophylla Hook.f
48 Impatiens commelinoides Hand.-Mazz. Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
49 Impatiens conchibracteata Y.L.Chen & Y.Q.Lu Lu and Chen 1991 accepted by WFO (placed as the accepted name of a taxon)
50 Impatiens cordata Wight Utami and Shimizu 2005 accepted by WFO (placed as the accepted name of a taxon)
51 Impatiens cornutisepala S.X.Yu, Y.L.Chen & H.N.Qin Yu et al. 2009 accepted by WFO (placed as the accepted name of a taxon)
52 Impatiens cyanantha Hook.f. Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
53 Impatiens cyathiflora Hook.f. Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
54 Impatiens cymbifera Hook.f. Zhang et al. 2016 accepted by WFO (placed as the accepted name of a taxon)
55 Impatiens damingensis S.X.Yu, Chang Y. Xia & H. P.Deng Xia et al. 2019 accepted by WFO (placed as the accepted name of a taxon)
56 Impatiens davidii Franch. Chen et al. 2007 accepted by WFO (placed as the accepted name of a taxon)
57 Impatiens davidii Franch. Song et al. 2005 accepted by WFO (placed as the accepted name of a taxon)
58 Impatiens decipiens Hook.f. Rewicz et al. 2020a accepted by WFO (placed as the accepted name of a taxon)
59 Impatiens delavayi Franch. Yu et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
60 Impatiens delavayi Franch. Song et al. 2005 accepted by WFO (placed as the accepted name of a taxon)
61 Impatiens devendrae Pusalkar Pusalkar and Singh 2010 accepted by WFO (placed as the accepted name of a taxon)
62 Impatiens dicentra Franch. ex Hook.f. Song et al. 2021c accepted by WFO (placed as the accepted name of a taxon)
63 Impatiens dicentra Franch. ex Hook.f. Chen et al. 2007 accepted by WFO (placed as the accepted name of a taxon)
64 Impatiens dolichoceras E.Pritz. ex Diels Song et al. 2005 accepted by WFO (placed as the accepted name of a taxon)
65 Impatiens drepanophora Hook.f. Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
66 Impatiens drepanophora Hook.f. Rewicz et al. 2020a accepted by WFO (placed as the accepted name of a taxon)
67 Impatiens edgeworthii Hook.f. Abid et al. 2011 accepted by WFO (placed as the accepted name of a taxon)
68 Impatiens elwiraurzu-lae Eb. Fisch., Abrah., Holstein & S.B. Janssens Fischer et al. 2021 accepted by WFO (placed as the accepted name of a taxon)
69 Impatiens faberi Hook.f. Lu and Chen 1991 accepted by WFO (placed as the accepted name of a taxon)
70 Impatiens falcifera Hook.f. Zhang et al. 2016 accepted by WFO (placed as the accepted name of a taxon)
71 Impatiens fanjingshanica Y.L.Chen Cai et al. 2013 accepted by WFO (placed as the accepted name of a taxon)
72 Impatiens fenghwaiana Y.L.Chen Song et al. 2005 accepted by WFO (placed as the accepted name of a taxon)
73 Impatiens flemingii Hook.f. Abid et al. 2011 accepted by WFO (placed as the accepted name of a taxon)
74 Impatiens forrestii Hook.f. & W.W. Sm. Song et al. 2005 accepted by WFO (placed as the accepted name of a taxon)
75 Impatiens fragicolor C.Marquand & Airy Shaw Cai et al. 2013 accepted by WFO (placed as the accepted name of a taxon)
76 Impatiens fragicolor C.Marquand & Airy Shaw Zhang et al. 2016 accepted by WFO (placed as the accepted name of a taxon)
77 Impatiens fugongensis K.M.Liu & Y.Y.Cong Cai et al. 2013 accepted by WFO (placed as the accepted name of a taxon)
78 Impatiens fugongensis K.M.Liu & Y.Y.Cong Cong et al. 2008 accepted by WFO (placed as the accepted name of a taxon)
79 Impatiens galactica Eb.Fisch., Raheliv. & Abrah. Fischer et al. 2017 accepted by WFO (placed as the accepted name of a taxon)
80 Impatiens gardneriana Wight. Utami and Shimizu 2005 accepted by WFO (placed as the accepted name of a taxon)
81 Impatiens garrettii Craib Utami and Shimizu 2005 synonym of Impatiens porrecta Wall. ex Hook.f. & Thomson
82 Impatiens gesneroidea Gilg Fischer et al. 2021 accepted by WFO (placed as the accepted name of a taxon)
83 Impatiens glandulifera Royle Najberek et al. 2020 accepted by WFO (placed as the accepted name of a taxon)
84 Impatiens glandulifera Royle Abid et al. 2011 accepted by WFO (placed as the accepted name of a taxon)
85 Impatiens glandulifera Royle Maciejewska-Rutkowska and Janczak 2016 accepted by WFO (placed as the accepted name of a taxon)
86 Impatiens gongshanensis Y.L.Chen Song et al. 2005 accepted by WFO (placed as the accepted name of a taxon)
87 Impatiens goughii Wight Utami and Shimizu 2005 accepted by WFO (placed as the accepted name of a taxon)
88 Impatiens guizhouensis Y.L.Chen Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
89 Impatiens hainanensis Y.L.Chen Bi et al. 2010 accepted by WFO (placed as the accepted name of a taxon)
90 Impatiens harae H.Ohba & S. Akiyama Zhang et al. 2016 accepted by WFO (placed as the accepted name of a taxon)
91 Impatiens henslowiana Arn. Utami and Shimizu 2005 accepted by WFO (placed as the accepted name of a taxon)
92 Impatiens hirta L.Joseph & Bhaskar Bhaskar 2012 accepted by WFO
93 Impatiens holocentra Hand.-Mazz. Cai et al. 2013 accepted by WFO (placed as the accepted name of a taxon)
94 Impatiens huangyanensis X.F.Jin & B.Y.Ding subsp. attenuata X.F.Jin & Z.H.Chen Song et al. 2022 accepted by WFO (placed as the accepted name of an infraspecific taxon of the species Impatiens huangyanensis X.F.Jin & B.Y.Ding)
95 Impatiens huangyanensis X.F.Jin & B.Y.Ding subsp. attenuata X.F.Jin & Z.H.Chen Jin et al. 2008 accepted by WFO (placed as the accepted name of an infraspecific taxon of the species Impatiens huangyanensis X.F.Jin & B.Y.Ding)
96 Impatiens huangyanensis X.F.Jin & B.Y.Ding subsp. huangyanensis Jin et al. 2008 accepted by WFO (placed as the accepted name of an infraspecific taxon of the species Impatiens huangyanensis X.F.Jin & B.Y.Ding)
97 Impatiens hunanensis Y.L.Chen Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
98 Impatiens imbecilla Hook.f. Lu and Chen 1991 accepted by WFO (placed as the accepted name of a taxon)
99 Impatiens infirma Hook.f. Song et al. 2005 accepted by WFO (placed as the accepted name of a taxon)
100 Impatiens jenjittikuliae Ruchis. & Suksathan Ruchisansakun and Suksathan 2019 accepted by WFO (placed as the accepted name of a taxon)
101 Impatiens jinggangensis Y.L.Chen Cai et al. 2013 accepted by WFO (placed as the accepted name of a taxon)
102 Impatiens jinpingensis Y.M.Shui & G.F.Li Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
103 Impatiens jurpia Buch.-Ham. ex Hook.f. & Thomson Rewicz et al. 2020a accepted by WFO (placed as the accepted name of a taxon)
104 Impatiens kivuensis Eb.Fisch., Abrah., Holstein & S.B.Janssens Fischer et al. 2021 accepted by WFO (placed as the accepted name of a taxon)
105 Impatiens larsenii T.Shimizu Utami and Shimizu 2005 accepted by WFO (placed as the accepted name of a taxon)
106 Impatiens lateristachys Y.L.Chen & Y.Q.Lu Lu and Chen 1991 accepted by WFO (placed as the accepted name of a taxon)
107 Impatiens latipetala S.H.Huang Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
108 Impatiens laxiflora Edgew. Zhang et al. 2016 accepted by WFO (placed as the accepted name of a taxon)
109 Impatiens leggei Pusalkar & D.K.Singh Pusalkar and Singh 2010 synonym of Impatiens laxiflora Edgew.
110 Impatiens lemannii Hook.f. & Thomson subsp. kurramensis Grey-Wilson Abid et al. 2011 accepted by WFO (placed as the accepted name of an infraspecific taxon of the species Impatiens lemannii Hook.f. & Thomson)
111 Impatiens lemannii Hook.f. & Thomson subsp. lemannii Abid et al. 2011 accepted by WFO (placed as the accepted name of an infraspecific taxon of the species Impatiens lemannii Hook.f. & Thomson)
112 Impatiens lemeei H.Lév. Chen et al. 2007 accepted by WFO (placed as the accepted name of a taxon)
113 Impatiens lepida Hook.f. Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
114 Impatiens leptocaulon Hook.f Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
115 Impatiens liboensis K.M.Liu & R.P.Kuang Kuang et al. 2014 accepted by WFO (placed as the accepted name of a taxon)
116 Impatiens linearisepala S.Akiyama, H.Ohba & S.K.Wu Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
117 Impatiens lobulifera S.X.Yu, Y.L.Chen & H.N.Qin Yu et al. 2009 accepted by WFO (placed as the accepted name of a taxon)
118 Impatiens longialata E.Pritz. Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
119 Impatiens longicornuta Y.L.Chen Cai et al. 2013 accepted by WFO (placed as the accepted name of a taxon)
120 Impatiens longipes Hook.f. & Thomson Zhang et al. 2016 accepted by WFO (placed as the accepted name of a taxon)
121 Impatiens longirostris S.H.Huang Cai et al. 2013 accepted by WFO (placed as the accepted name of a taxon)
122 ^Impatiens longshanensis Y.Y.Cong & Y.X.Song Song et al. 2021c unplaced by WFO (this name is currently unchecked and awaiting taxonomic scrutiny), found on IPNI
123 Impatiens ludewigii Eb.Fisch., Abrah., Holstein & S.B.Janssens Fischer et al. 2021 accepted by WFO (placed as the accepted name of a taxon)
124 Impatiens lutzmanniae Eb.Fisch., Abrah., Holstein & S.B.Janssens Fischer et al. 2021 accepted by WFO (placed as the accepted name of a taxon)
125 Impatiens macrovexilla var. macrovexilla Yu 2007 accepted by WFO (placed as the accepted name of an infraspecific taxon of the species Impatiens macrovexilla Y.L.Chen)
126 Impatiens macrovexilla Y.L.Chen var. yaoshanensis S.X.Yu, Y.L.Chen & H.N.Qin Yu 2007 accepted by WFO (placed as the accepted name of an infraspecific taxon of the species Impatiens macrovexilla Y.L.Chen)
127 Impatiens maculifera S.X.Yu & Chang Y.Xia Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
128 Impatiens maculifera S.X.Yu & Chang Y.Xia Xia et al. 2019 accepted by WFO (placed as the accepted name of a taxon)
129 Impatiens margaritifera Hook.f. Zhang et al. 2016 accepted by WFO (placed as the accepted name of a taxon)
130 Impatiens menghuochengensis Q.Luo Luo et al. 2014 accepted by WFO (placed as the accepted name of a taxon)
131 Impatiens mengtszeana Hook.f. Song et al. 2005 synonym of Impatiens pulchra Hook.f. & Thomson.; accepted name in the genus Impatiens
132 Impatiens meruensis Gilg Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
133 Impatiens meyana Hook.f. Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
134 Impatiens monticola Hook.f. Cai et al. 2013 synonym of Impatiens pulchra Hook.f. & Thomson.; accepted name in the genus Impatiens
135 Impatiens monticola Hook.f. Song et al. 2022 synonym of Impatiens pulchra Hook.f. & Thomson.; accepted name in the genus Impatiens
136 Impatiens monticola Hook.f. Lu and Chen 1991 synonym of Impatiens pulchra Hook.f. & Thomson.; accepted name in the genus Impatiens
137 Impatiens monticola Hook.f. Cong et al. 2008 synonym of Impatiens pulchra Hook.f. & Thomson.; accepted name in the genus Impatiens
138 Impatiens morsei Hook.f. Yu et al. 2015 accepted by WFO (placed as the accepted name of a taxon)
139 Impatiens morsei Hook.f. Yu et al. 2009 accepted by WFO (placed as the accepted name of a taxon)
140 ^Impatiens muscicola Craib Utami and Shimizu 2005 unplaced by WFO (this name is currently unchecked and awaiting taxonomic scrutiny), found on IPNI
141 Impatiens nagorum Gogoi, Moaakum & S.Dey Moaakum et al. 2017 accepted by WFO (placed as the accepted name of a taxon)
142 Impatiens namkatensis T.Shimizu Utami and Shimizu 2005 accepted by WFO (placed as the accepted name of a taxon)
143 Impatiens napoensis Y.L.Chen Cai et al. 2013 accepted by WFO (placed as the accepted name of a taxon)
144 Impatiens nasuta H.Perrier Song et al. 2022 synonym of Impatiens kanchigandhiana Rasingam, Karthig. & Gogoi.; accepted name in the genus Impatiens
145 Impatiens noli-tangere L. Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
146 Impatiens noli-tangere L. Song et al. 2005 accepted by WFO (placed as the accepted name of a taxon)
147 Impatiens noli-tangere L. Utami and Shimizu 2005 accepted by WFO (placed as the accepted name of a taxon)
148 Impatiens noli-tangere L. Chen et al. 2007 accepted by WFO (placed as the accepted name of a taxon)
149 Impatiens noli-tangere L. Jin et al. 2008 accepted by WFO (placed as the accepted name of a taxon)
150 Impatiens nyimana C.Marquand & Airy Shaw Rewicz et al. 2020a accepted by WFO (placed as the accepted name of a taxon)
151 Impatiens nyimana C.Marquand & Airy Shaw Zhang et al. 2016 accepted by WFO (placed as the accepted name of a taxon)
152 Impatiens obesa Hook.f. Shimizu 1987 accepted by WFO (placed as the accepted name of a taxon)
153 Impatiens obesa Hook.f. Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
154 Impatiens obesa Hook.f. Bi et al. 2010 accepted by WFO (placed as the accepted name of a taxon)
155 Impatiens oblongipetala K.M.Liu & Y.Y.Cong Cai et al. 2013 accepted by WFO (placed as the accepted name of a taxon)
156 Impatiens oblongipetala K.M.Liu & Y.Y.Cong Cong et al. 2010 accepted by WFO (placed as the accepted name of a taxon)
157 Impatiens omeiana Hook.f. Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
158 Impatiens omissa Hook.f. Utami and Shimizu 2005 accepted by WFO (placed as the accepted name of a taxon)
159 Impatiens oxyanthera Hook.f. Lu and Chen 1991 accepted by WFO (placed as the accepted name of a taxon)
160 Impatiens parviflora DC. Song et al. 2005 accepted by WFO (placed as the accepted name of a taxon)
161 ^Impatiens patula Craib Utami and Shimizu 2005 unplaced by WFO (this name is currently unchecked and awaiting taxonomic scrutiny), found on IPNI
162 Impatiens pianmaensis S.H.Huang Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
163 Impatiens pinetorum Hook.f. ex W.W.Sm. Cai et al. 2013 accepted by WFO (placed as the accepted name of a taxon)
164 Impatiens pingxiangensis H.Y.Bi & S.X.Yu Bi et al. 2010 accepted by WFO (placed as the accepted name of a taxon)
165 Impatiens piufanensis Hook.f. Chen et al. 2007 accepted by WFO (placed as the accepted name of a taxon)
166 Impatiens platychlaena Hook.f. Lu and Chen 1991 accepted by WFO (placed as the accepted name of a taxon)
167 Impatiens platysepala Y.L.Chen Yu et al. 2015 accepted by WFO (placed as the accepted name of a taxon)
168 Impatiens platysepala Y.L.Chen Wang et al. 2020 accepted by WFO (placed as the accepted name of a taxon)
169 Impatiens poculifer Hook.f. Song et al. 2005 accepted by WFO (placed as the accepted name of a taxon)
170 Impatiens polyneura K.M.Liu Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
171 Impatiens polyneura K.M.Liu Chen et al. 2007 accepted by WFO (placed as the accepted name of a taxon)
172 Impatiens principis Hook.f. Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
173 Impatiens principis Hook.f. Zhang et al. 2016 accepted by WFO (placed as the accepted name of a taxon)
174 Impatiens pritzelii Hook.f. Chen et al. 2007 accepted by WFO (placed as the accepted name of a taxon)
175 Impatiens pseudoviola Gilg Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
176 Impatiens psittacina Hook.f. Utami and Shimizu 2005 accepted by WFO (placed as the accepted name of a taxon)
177 Impatiens pterosepala Hook.f. Cai et al. 2013 accepted by WFO (placed as the accepted name of a taxon)
178 Impatiens pterosepala Hook.f. Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
179 Impatiens purpurea Hand.-Mazz. Song et al. 2005 accepted by WFO (placed as the accepted name of a taxon)
180 Impatiens purpureifolia S.H.Huang & Y.M.Shui Shui et al. 2011 accepted by WFO (placed as the accepted name of a taxon)
181 Impatiens purpureoviolacea Gilg Fischer et al. 2021 accepted by WFO (placed as the accepted name of a taxon)
182 Impatiens quadriloba K.M.Liu & Y.L.Xiang Cai et al. 2013 accepted by WFO (placed as the accepted name of a taxon)
183 Impatiens quadriloba K.M.Liu & Y.L.Xiang Cong et al. 2010 accepted by WFO (placed as the accepted name of a taxon)
184 Impatiens racemosa DC. Zhang et al. 2016 accepted by WFO (placed as the accepted name of a taxon)
185 Impatiens radiata Hook.f. Zhang et al. 2016 accepted by WFO (placed as the accepted name of a taxon)
186 Impatiens rapiformis Y.Y.Cong & Y.X.Song Song et al. 2021b accepted by WFO (placed as the accepted name of a taxon)
187 Impatiens rapiformis Y.Y.Cong & Y.X.Song Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
188 Impatiens rectangula Hand.-Mazz. Zhang et al. 2016 accepted by WFO (placed as the accepted name of a taxon)
189 Impatiens reptans Hook.f. Chen et al. 2007 synonym of Impatiens procumbens Franch.; accepted name in the genus Impatiens
190 Impatiens rhombifolia Y.Q.Lu & Y.L.Chen Lu and Chen 1991 synonym of Impatiens procumbens Franch.; accepted name in the genus Impatiens
191 Impatiens rostellata Franch. Lu and Chen 1991 accepted by WFO (placed as the accepted name of a taxon)
192 Impatiens rostellata Franch. Song et al. 2005 accepted by WFO (placed as the accepted name of a taxon)
193 Impatiens rubro-striata Hook.f. Cai et al. 2013 accepted by WFO (placed as the accepted name of a taxon)
194 Impatiens rubro-striata Hook.f. Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
195 Impatiens rugata S.H.Huang & Y.M.Shui Shui et al. 2011 accepted by WFO (placed as the accepted name of a taxon)
196 Impatiens ruiliensis S.Akiyama & H.Ohba Song et al. 2005 accepted by WFO (placed as the accepted name of a taxon)
197 Impatiens ruiliensis S.Akiyama & H.Ohba Zhang et al. 2016 accepted by WFO (placed as the accepted name of a taxon)
198 Impatiens rupestris K.M.Liu & X.Z.Cai Cai et al. 2013 accepted by WFO (placed as the accepted name of a taxon)
199 Impatiens rupestris K.M.Liu & X.Z.Cai Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
200 Impatiens scabrida DC. Song et al. 2005 accepted by WFO (placed as the accepted name of a taxon)
201 Impatiens scabrida DC. Abid et al. 2011 accepted by WFO (placed as the accepted name of a taxon)
202 Impatiens scabrida DC. Zhang et al. 2016 accepted by WFO (placed as the accepted name of a taxon)
203 Impatiens scullyi Hook.f. Zhang et al. 2016 accepted by WFO (placed as the accepted name of a taxon)
204 Impatiens shennongensis Qiang Wang & H.P.Deng Wang et al. 2016 accepted by WFO (placed as the accepted name of a taxon)
205 Impatiens siamensis T.Shimizu Utami and Shimizu 2005 accepted by WFO (placed as the accepted name of a taxon)
206 ? Impatiens siculifer Hook.f. Song et al. 2005 Found on WFO as Impatiens siculifera, but originally named as Impatiens siculifer by Hooker (1908)
207 ? Impatiens siculifer Hook.f. Zhang et al. 2016 Found on WFO as Impatiens siculifera, but originally named as Impatiens siculifer by Hooker (1908)
208 Impatiens siculifera Hook.f. Chen et al. 2007 accepted by WFO (placed as the accepted name of a taxon)
209 ? Impatiens siculifera var. porphyrea Hook.f. Chen et al. 2007 accepted by WFO (placed as the accepted name of an infraspecific taxon of the species Impatiens siculifera Hook.f.)
210 ? Impatiens siculifera var. mitis Lingelsh. & Borza Song et al. 2022 accepted by WFO (placed as the accepted name of an infraspecific taxon of the species Impatiens siculifera Hook.f.)
211 Impatiens sikaiensis Q. Luo & Ying Yuan Yuan et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
212 Impatiens soulieana Hook.f. Song et al. 2005 accepted by WFO (placed as the accepted name of a taxon)
213 Impatiens spirifera Hook.f. & Thomson Rewicz et al. 2020a accepted by WFO (placed as the accepted name of a taxon)
214 Impatiens stenantha Hook.f. Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
215 Impatiens stenosepala E.Pritz. Chen et al. 2007 accepted by WFO (placed as the accepted name of a taxon)
216 Impatiens sterilis Y.Y. Cong & Y.X.Song Song et al. 2021a accepted by WFO (placed as the accepted name of a taxon)
217 Impatiens stocksii Hook.f. & Thomson Utami and Shimizu 2005 accepted by WFO (placed as the accepted name of a taxon)
218 Impatiens stuhlmannii Warb. Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
219 Impatiens sulcata Wall. Cai et al. 2013 accepted by WFO (placed as the accepted name of a taxon)
220 Impatiens sulcata Wall. Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
221 Impatiens sulcata Wall. Abid et al. 2011 accepted by WFO (placed as the accepted name of a taxon)
222 Impatiens sulcata Wall. Zhang et al. 2016 accepted by WFO (placed as the accepted name of a taxon)
223 Impatiens sulcata Wall. Rewicz et al. 2020a accepted by WFO (placed as the accepted name of a taxon)
224 Impatiens sunkoshiensis S.Akiyama, H.Ohba & Wakab. Zhang et al. 2016 synonym of Impatiens laxiflora Edgew.; accepted name in the genus Impatiens
225 Impatiens sutchuenensis Franch. ex Hook.f. Cai et al. 2013 accepted by WFO (placed as the accepted name of a taxon)
226 Impatiens teitensis sub. teitensis Song et al. 2022 accepted by WFO (placed as the accepted name of an infraspecific taxon of the species Impatiens teitensis Grey-Wilson)
227 Impatiens textori Miq. Utami and Shimizu 2005 accepted by WFO (placed as the accepted name of a taxon)
228 Impatiens thomsonii Hook.f. Abid et al. 2011 accepted by WFO (placed as the accepted name of a taxon)
229 Impatiens tomentella Hook.f. Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
230 Impatiens tongbiguanensis S.Akiyama & H.Ohba Cai et al. 2013 synonym of Impatiens leptoceras DC.; accepted name in the genus Impatiens
231 Impatiens tortisepala Hook.f. Luo et al. 2014 accepted by WFO (placed as the accepted name of a taxon)
232 Impatiens tripetala Roxb. & DC. Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
233 Impatiens troupinii Eb.Fisch., Abrah., Holstein & S.B.Janssens Fischer et al. 2021 accepted by WFO (placed as the accepted name of a taxon)
234 Impatiens tsangshanensis Y.L.Chen Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
235 Impatiens tuberculata Hook.f. & Thomson Zhang et al. 2016 accepted by WFO (placed as the accepted name of a taxon)
236 Impatiens uliginosa Franch. Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
237 Impatiens undulata Y.L.Chen & Y.Q.Lu Lu and Chen 1991 accepted by WFO (placed as the accepted name of a taxon)
238 Impatiens unguiculata K.M.Liu & Y.Y.Cong Cong et al. 2013 accepted by WFO (placed as the accepted name of a taxon)
239 Impatiens urticifolia Wall. Zhang et al. 2016 accepted by WFO (placed as the accepted name of a taxon)
240 Impatiens urundiensis Gilg Fischer et al. 2021 synonym of Impatiens purpureoviolacea Gilg.; accepted name in the genus Impatiens
241 Impatiens versicolor Eb. Fisch., Abrah., Holstein & S.B.Janssens Fischer et al. 2021 accepted by WFO (placed as the accepted name of a taxon)
242 Impatiens walleriana Hook.f. Song et al. 2005 accepted by WFO (placed as the accepted name of a taxon)
243 Impatiens walleriana Hook.f. Utami and Shimizu 2005 accepted by WFO (placed as the accepted name of a taxon)
244 Impatiens wawuensis Bo Ding & S.X.Yu Ding et al. 2016 accepted by WFO (placed as the accepted name of a taxon)
245 Impatiens wenshanensis S.H.Huang Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
246 Impatiens wilsonii Hook.f. Lu and Chen 1991 accepted by WFO (placed as the accepted name of a taxon)
247 Impatiens wuerstenii S.B. Janssens & Dessein Janssens et al. 2018 accepted by WFO (placed as the accepted name of a taxon)
248 Impatiens wuyiensis J.S.Wang, Y.F.Lu & X.F.Jin Wang et al. 2020 accepted by WFO (placed as the accepted name of a taxon)
249 Impatiens wuyuanensis Y.L.Chen Cai et al. 2013 accepted by WFO (placed as the accepted name of a taxon)
250 Impatiens xanthina H.F.Comber Song et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
251 Impatiens xanthina H.F.Comber Song et al. 2005 accepted by WFO (placed as the accepted name of a taxon)
252 Impatiens xanthina H.F.Comber Chen et al. 2007 accepted by WFO (placed as the accepted name of a taxon)
253 Impatiens xanthina H.F.Comber Cong et al. 2008 accepted by WFO (placed as the accepted name of a taxon)
254 Impatiens xanthinoides G.W.Hu Cai et al. 2015 accepted by WFO (placed as the accepted name of a taxon)
255 ^Impatiens xishuangbannaensis S.H.Huang Song et al. 2022 unplaced by WFO (a taxonomist hasn’t yet placed the name in the taxonomy), found on IPNI
256 Impatiens yilingiana X.F.Jin, Shu Z.Yang & L.Qian Jin et al. 2008 accepted by WFO (placed as the accepted name of a taxon)
257 Impatiens yingjiangensis S.Akiyama & H.Ohba Cai et al. 2013 accepted by WFO (placed as the accepted name of a taxon)
258 Impatiens yunlingensis S.X.Yo, Chang Y.Xia & J.H.Yu Yu et al. 2022 accepted by WFO (placed as the accepted name of a taxon)
259 ^Impatiens zhaojueensis Q.Luo Luo et al. 2023 unplaced by WFO (a taxonomist hasn’t yet placed the name in the taxonomy), found on IPNI
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^ - species name unchecked and awaiting taxonomic scrutiny by WFO ? - species name not found in WFO

We utilized the PRISMA Flowchart as a visual tool to outline the systematic review process, from the initial literature search to the final inclusion of studies (Page et al. 2021https://estech.shinyapps.io/prisma_flowdiagram/) (Fig. 1).

Figure 1.

Figure 1.

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PRISMA flow chart – visualization of the systematic review process, from the initial literature search to the final inclusion of data in studies.

The species nomenclature validation was adopted from The World Flora Online (WFO 2024), Plants of the World Online (POWO 2024) and International Plant Names Index (IPNI 2024), especially in the most recent publications.

In Table 1, species names are divided into four categories: 1) names confirmed in The World Flora Online (WFO) database – Accepted nomenclature; 2) names that have not yet been confirmed in The World Flora Online (WFO) database – Unchecked and awaiting taxonomic scrutiny; 3) names that appeared under synonyms in the articles - the current names are placed in the column Remarks; 4) species name not found in WFO.

Additionally, three graphs were generated based on three aspects: 1.-number of publications presenting seeds processed with SEM for taxa of the Impatiens genus, 2.-number of Impatiens taxa with seeds processed with SEM over time, and 3.-number of publications on seeds without SEM photos but with seeds information for taxa of the Impatiens genus (Figs 2, 3, 5 respectively)

Figure 2.

Figure 2.

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Number of publications featuring SEM-processed seeds for taxa of the genus Impatiens (based on data from Table 1).

Figure 3.

Figure 3.

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Number of Impatiens taxa with SEM-processed seeds over time (based on data from Table 1).

Figure 5.

Figure 5.

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Number of publications and records (online databases) on seeds without SEM photos but with seed information for taxa of the genus Impatiens (based on Suppl. material 1).

Finally, to provide a clear visual representation of the countries and species with the highest number of records under the SEM criterion, a heat map was generated in Microsoft Excel 2018 (Fig. 5). Additionally, a word cloud was generated using the “wordcloud” package (Fellows et al. 2018) in R (R Development Core Team 2012) (Fig. 6).

Figure 6.

Figure 6.

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Wordcloud of Impatiens species with frequencies of SEM studies. The symbol ‘-’ was added to some species, subspecies and varieties to recognize them as a single word.

Results

Literature data with SEM photos

A total of 47 references in the literature pertain to the carpology of seeds within the genus Impatiens, including attached SEM images, and encompass information on 183 unique spp. and 4 subspecies/variations (with accepted nomenclature by WFO), (Table 1, Fig. 1).

Single carpological descriptions have been recorded for most species. Only a few species had information available from more than one literature source: for instance, I. arguta, I. balfourii, and others I. drepanophora and others were documented in two sources; I. aquatilis, I. balsamina, I. chinensis in three; and I. xanthina in four. In the case of three species (I. noli-tangere, I. pulchra, and I. sulcata), information was found in five publications (Table 1, Fig. 6).

Information regarding the seed carpology of the genus Impatiens (with SEM photos) was found in 47 articles, with the earliest ones dating back to the early 1990s. From 1991 to 2004, individual articles on the carpology of the Impatiens genus were published sporadically. Subsequently, there was a fluctuating level of publishing activity on this topic, with significant increases in 2016, 2017, 2020, and 2023 (Fig. 2).

Despite the limited number of studies on the carpology of Impatiens genus seeds since the early 1990s, notable contributions were made by Song et al. (2005) with 50 taxa, followed by Chen et al. (2007) with 16, and Abid et al. (2011) with 14 taxa. Subsequent significant works providing carpological descriptions for large groups of Impatiens taxa included those of Cai et al. (2013) with 25 taxa, Zhang et al. (2016) with 26, and Song et al. (2022) with 51 taxa (Fig. 3).

Most of the material originated from Asia, particularly from China (187 taxa), India (21), Pakistan (11) and Thailand (9). A few individual samples were sourced from Europe, including Italy (2), Switzerland (2) and Poland (2), but also Japan (2) and Kenya (4) (Fig. 4).

Figure 4.

Figure 4.

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Country of origin of the material of Impatiens genus (based on data from Table 1).

Literature data without SEM photos

Through our literature review, we identified around 336 accepted species, six subspecies and one hybrid of seeds belonging to various taxa of the genus Impatiens. These descriptions encompassed a range of information, including morphological characteristics, binocular photographs, and illustrations (Suppl. material 1). The publications containing these descriptions do not include SEM photographs.

An analysis of publications on the carpology of the genus Impatiens that do not contain SEM images reveals various trends and variability in the number of publications over the past 200 years. During this period, there were 128 publications on this topic. The earliest publications date back to the XIX century (year 1824). The greatest increase in interest occurred after 2000, when articles on specimens of the Impatiens genus, most frequently including descriptions of the seeds, began to appear more often (Fig. 5), especially due to the presence of online databases (Suppl. material 1).

Discussion and conclusion

Investigating the microstructure of seeds (e. g., primary, and secondary seed coat ornamentation, periclinal cell wall patterns) is essential not only for taxonomic purposes but also as a tool for gaining insights into plant developmental processes, including the adaptation of plants to changing environmental conditions (Utami and Shimizu 2005; Yu et al. 2015; Zhang et al. 2016; Rewicz et al. 2022b). Understanding these adaptations is crucial for comprehending how plants thrive in diverse environments and how they can be protected, particularly in the face of challenges posed by climate change (Bailly and Gomez Roldan 2023). A key element is understanding the micromorphology of seeds and fruits (Van Waveren 2019; Rewicz et al. 2020a; Rewicz et al. 2020b). To facilitate robust conclusions and data comparisons, carpological research, especially regarding seed coat morphology, should be based on high-quality photographs, preferably obtained through scanning electron microscopy. This approach helps mitigate potential discrepancies across different descriptive systems and minimizes researcher subjectivity (Zhao et al. 2023). The introduction of scanning electron microscopy (SEM) was an important moment in scientific advancement, and research using this technology flourished in the late 1990s.

Our analysis of the literature on the microstructure of seeds from the genus Impatiens confirms this trend, as the first attempts to use SEM in research on this genus Impatiens date back to 1990 (Fig. 2). Unfortunately, despite a significant increase in the number of carpological publications using SEM images, our analysis of literature sources showed that only 21% of taxa from the entire genus, which includes more than 1,000 species, have data with SEM images (Grey-Wilson 1980; Zhao-Cen et al. 2020; Tanaka et al. 2022). The analysis of literature sources also shows that the majority of species are described in only one publication, and the material was collected from a single locality (Table 1). There are, however, exceptions, where several species have garnered more research attention, such as I. aquatilis, I. bahanensis, I. balsamina, I. chinensis, I. fugongensis, I. monticola, I. noli-tangere, I. scabrida, I. sulcata, and I. xanthina. However, it’s worth noting that for Impatiens species found in Europe, almost all (with the exception of I. parviflora) have SEM images of seeds available. This is particularly significant because species like I. glandulifera or I. capensis are considered invasive. Therefore, any insight into their biology in non-native ranges becomes crucial for effective management and control strategies (Table 1). Our literature analysis revealed that carpological data for over 300 species of Impatiens are available, but without descriptions using SEM (Suppl. material 1). At the same time, it should be noted that the seeds of this genus have long been a subject of interest for researchers (Hooker 1910; Fischer 1934).

Our research also revealed a significant oversight in the carpological studies of Impatiens, namely the neglect of seed phenotypic variability. Analyzing the variability of quantitative and qualitative characteristics across individuals from diverse locations, especially those covering a wide geographic range, provides valuable insights into the extents of phenotypic variability and the underlying determinants, such as habitat and climate. Such data also serve as a vital resource for understanding the biology and ecology of the studied species (Rewicz et al. 2016; Kostrakiewicz-Gierałt et al. 2022). Furthermore, analyzing phenotypic plasticity enables us to ascertain whether a given structure exhibits stability or substantial intraspecific variability in response to habitat or climatic influences (Stace 1992). Our literature review on Impatiens seeds identified only one study, specifically by Rewicz et al. (2020b), that described the phenotypic plasticity of seeds, focusing on an invasive species in Europe - I. capensis (Rewicz et al. 2022a).

In our opinion, material availability plays a key role in carpological research. It is noteworthy that research on material from herbarium collections is not always possible due to two main reasons. Firstly, the quantity of material on an herbarium sheet is sometimes limited, rendering it impossible for collection or comprehensive analyses. Secondly, there is a risk of mechanical damage to the material, such as when pressing it into an herbarium sheet, which impedes its utility for research purposes.

Furthermore, it is important to recognize that collecting material from areas of high biodiversity is often restricted for legal reasons. These constraints may pose a significant obstacle in obtaining representative material for carpological research.

The analysis of the literature pertaining to the genus Impatiens indicates that majority of the material (seeds) originates from China, accounting for 87% of the cases examined. Following China, the next region with considerably higher seed representation is Madagascar and East-Central Africa. We posit that this distribution pattern is attributed to the abundance of material available from the genus Impatiens, as these regions are among the five hot spots with the highest species richness (Grey-Wilson 1980; Yuan et al. 2004; Song 2006; Tabak and von Wettberg 2008; Zika 2009; Ruchisansakun et al. 2014).

Further carpological research on the genus Impatiens is imperative for advancing our understanding of these plants. Identifying new species, elucidating their geographical distribution, life cycles, and ecology are among the numerous benefits of such research. Additionally, carpological research can also shed new light on the evolutionary history of these plants and the interrelationships among individual species. Moreover, research on Impatiens carpology can be extremely helpful in monitoring the population status of these plants. Given the threats facing many species and their natural habitats, understanding population dynamics is crucial for effective conservation efforts.

In conclusion, carpological research on the genus Impatiens is not only of scientific importance, but also of practical significance for the protection of representatives of this genus.

Acknowledgments

We would like to express our sincere gratitude to PhD Wojciech Adamowski for his valuable feedback and insightful suggestions during the development of this article.

Citation

Rewicz A, Monzalvo R, Grey K, Sanek K, Mackiewicz M, Myśliwy M (2025) Gap analysis of knowledge about the microstructure of Impatiens Riv. ex L. (Balsaminaceae) seeds using SEM techniques. PhytoKeys 262: 105–128. https://doi.org/10.3897/phytokeys.262.152205

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Use of AI

No use of AI was reported.

Funding

Co-financed by the Minister of Science under the “Regional Excellence Initiative” Program for 2024–2027 (RID/SP/0045/2024/01). This research was partially funded by International researcher competitions of the University of Lodz financed as part of the subsidy increased by 2% for the universities that joined the Initiative of Excellence – Research University (IDUB) competition (B2211001000147.07) granted to AR, and partially by the statutory funds of the University of Lodz. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.

Author contributions

Investigation: AR, RM, MM, MMA, Methodology: AR, RM, KS, KG, Visualization: AR, RM, MMA, MM, KS, Writing – original draft: AR, RM, KG, MM, Writing – review and editing: AR, RM, KG, KS, MMA, MM.

Author ORCIDs

Agnieszka Rewicz https://orcid.org/0000-0002-8228-6406

René Monzalvo https://orcid.org/0000-0003-1049-5468

Monika Myśliwy https://orcid.org/0000-0001-6573-6455

Data availability

All of the data that support the results of this study are available in the manuscript and in the Supplementary Information (see Suppl. material 1).

Supplementary materials

Supplementary material 1

Impatiens species with seed information but no SEM photographs

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.

Agnieszka Rewicz, René Monzalvo, Kayla Grey, Katarzyna Sanek, Maja Mackiewicz, Monika Myśliwy

Data type

docx

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Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Supplementary Materials

Supplementary material 1

Impatiens species with seed information but no SEM photographs

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.

Agnieszka Rewicz, René Monzalvo, Kayla Grey, Katarzyna Sanek, Maja Mackiewicz, Monika Myśliwy

Data type

docx

phytokeys-262-105_article-152205__-s001.docx (727.9KB, docx)

Data Availability Statement

All of the data that support the results of this study are available in the manuscript and in the Supplementary Information (see Suppl. material 1).

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