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Article

Variation in Phenology and Morphological Traits of Seed-Propagated Laggera alata and Laggera crispata Forms

by
Noluthando Nonjabulo Nkosi
1,
Godfrey Elijah Zharare
2,
Clemence Zimudzi
3,
Brita Stedje
4 and
Nontuthuko Rosemary Ntuli
1,*
1
Department of Botany, Faculty of Science, Agriculture and Engineering, University of Zululand, KwaDlangezwa 3886, South Africa
2
Department of Agriculture, Faculty of Science, Agriculture and Engineering, University of Zululand, KwaDlangezwa 3886, South Africa
3
Department of Biological Sciences and Ecology, Faculty of Science, University of Zimbabwe, Harare P.O. Box MP167, Zimbabwe
4
Botanical Garden, Natural History Museum, University of Oslo, Blindern, P.O. Box 1172, 0318 Oslo, Norway
*
Author to whom correspondence should be addressed.
Diversity 2024, 16(8), 466; https://doi.org/10.3390/d16080466
Submission received: 21 June 2024 / Revised: 27 July 2024 / Accepted: 30 July 2024 / Published: 2 August 2024
Browse Figures
Versions Notes

Abstract

:
The phenological and morphological variation studies among forms of Laggera Sch. Bip. Ex. Benth. and Hook species are limited, despite the medicinal use of the genus. Therefore, this study aimed to document phenology and morphological variation in cultivated populations of Laggera alata and Laggera crispata forms from seedling to maturity. The forms were categorized as Laggera alata with a small capitulum (LA-SC), Laggera alata with a large capitulum (LA-BC), Laggera crispata from South Africa (LC-SA), and Laggera crispata from Zimbabwe (LC-ZIM). Seeds were germinated in Petri dishes, transplanted to plug trays, and later to field plots at 60 days. Phenological events were recorded when observed in at least one plant. Twelve qualitative and four morphometric traits were measured monthly on five plants per Laggera form. Analysis of variance and Tukey’s Honestly Significant Difference test (p < 0.05) were used for data analysis. Results indicated significant variation in phenology, qualitative traits, leaf traits, plant height, and stem diameter both within and between L. crispata and L. alata forms. Morphometric traits, such as leaf size and the number of leaves per plant, were identified as key descriptors for differentiating L. alata forms. These findings provide a foundation for the introduction of Laggera forms into farming systems for medicinal and commercial purposes.

1. Introduction

Plant phenology studies the time of recurring phenophases, such as new leaf foliage, leaf fall, flowering, and fruiting [1]. On the other hand, plant morphology is the science of plant form and development and can be differentiated into two types, namely, the morphology of systematics and the morphology of morphologists [2,3]. The latter is a character morphology and concludes in the character matrices of morphological cladistics [2]. Plant morphology is a backbone for all plant biology disciplines, such as plant diversity, phylogeny, evolution, physiology, molecular genetics, and systematics [4].
Biotic and abiotic factors greatly influence plant phenology [1,5]. The major abiotic factors affecting plant phenological patterns include photoperiod, temperature, and rainfall [6]. There is only limited general information about the phenological cycle of Laggera forms [7]. Documenting the phenology of Laggera forms will help determine the appropriate time for different phenological events, such as seed collection, before domestication or breeding efforts [8].
Several factors contribute to plant morphological variation, including season, elevation, species association, nutrients, and anthropogenic pressures [9]. Intraspecific morphometric variation in Laggera alata was recorded in Cameroon, and the researchers could not conclude whether these types were varieties or hybrids [10]. This proves that morphological comparative studies are crucial for species identification and differentiation [11]. Morphological identification of plants is accomplished by observing their organs’ size, color, shape, and other plant characteristics [3].
There are many similarities in the morphology of Laggera species and forms, resulting in taxonomic confusion, particularly when trying to identify the species and/or forms [12]. Most studies on the classification of Laggera species have used morphological characteristics of dried herbaria materials or live field plants [13,14,15]. Dry material is not always as easily observable as fresh material, making some morphological characters unrecognizable or deformed [16]. In addition, dried herbarium specimens exhibit significant differences in the plant organs’ dimensions compared to fresh material, but this is taxon-dependent [17]. Although dried herbaria material remains a valuable resource in plant systematics [18], there might be errors in the measurement data due to the shrinkage or expansion of the plant organs depending on the plant-drying method used [17]. Therefore, this study aimed to document phenology and quantify the morphological variation in Laggera alata and Laggera crispata forms in cultivated populations from seedling to maturity.

2. Materials and Methods

2.1. Plant Material and Growth Conditions

The seeds of four Laggera forms were germinated in Petri dishes: Laggera alata with large capitulum (LA-BC), Laggera alata with small capitulum (LA-SC), Laggera crispata from South Africa (LC-SA), and Laggera crispata from Zimbabwe (LC-ZIM). The emerging seedlings were then put in plug trays prefilled with a commercial potting mix under natural conditions in a net house at the Department of Botany (28.51° S, 31.50° E). The plug trays were placed in a water bath for irrigation purposes. Hydroponic fertilizer (5 g) was added to a water bath if the leaves of seedlings were turning yellow. At about 60 days after sowing, all seedlings were at the first to second genuine leaf stage and were transplanted onto the field. The 3–4–5 rule was used to lay out a rectangular field plot on uneven or irregular agricultural land to minimize the experimental error of the research findings [19]. The experimental plots were 2.25 m × 3 m and 1 m apart in size, and the seedlings were spaced at an intra-row spacing of 0.45 m and inter-row spacing of 0.75 m. The experiment adopted the randomized complete block design with three replications generated by R 4.2.1 software in the RStudio platform [20]. Each plot had 24 seedlings. Management practices, such as irrigation, weeding, and insecticide spraying, were carried out as needed. A sprinkler irrigation system was used to irrigate the field trial. Voucher specimens for each form were prepared and stored at the University of Zululand Herbarium. The field experiment was carried out for two seasons (from September 2021 to September 2022 and from September 2022 to September 2023) under the weather conditions detailed in Table 1.

2.2. Data Collection

Data collection began at 37 days after transplanting (DAT). Five plants from the inner rows of each plot were randomly selected, tagged with plastic tags, and used to collect morphological data every month. This represents a sample size of 15 plants per form. The collected data were presented as the mean of two seasons for the following parameters, outlined below.

2.2.1. Phenological Events

Radicle protrusion, vegetative growth, flower bud formation, first opened flower, flowering, seed set leaf senescence, seed dispersal, and resprouting were observed and recorded in each sampled plant. A particular phenological event’s initiation and completion dates were recorded [21]. For each sampled plant of a form, the beginning or end of a given phenological event was defined as the day on which at least one of the plants was seen to be experiencing it. Flowering was determined by the first opened flower seen on any plant and considered the first flowering day for that particular plant of the species, while the last flowering day was when only a single flower remained on any plant [5]. Then, for each form, phenophase calendars were created.

2.2.2. Morphological Traits

Morphological characterization was conducted using six quantitative morphometric traits (leaf area, stem diameter, number of leaves, plant height, and chlorophyll content) and six qualitative morphological traits (leaf color, leaf margin, stem wing shape, leaf shape, flower color, and leaf apex). A measuring tape measured plant height (mm) from aboveground level to the top of the spike. Direct counting was used to ascertain the number of leaves per plant. A meter ruler measured the leaf length (mm) and width (mm) of the third leaf from the apex. The leaf area (mm2) was calculated as the product of leaf length and leaf width and the correction factor. The correction factor of 0.74 was used in this study since Sclepias syriaca species’ leaves are as variable in shape as Laggera forms’ leaves (ovate, oblong, lanceolate, or elliptic) [22]. A SPAD 502 chlorophyll meter was used to determine the leaf chlorophyll content (mg cm−2). The stem diameter was measured at 5 cm aboveground using a Vernier caliper.

2.3. Statistical Analysis

The raw data on quantitative morphological traits were subjected to analysis of variance (ANOVA) using GenStat 15th edition software. Means were separated using Tukey’s Honestly Significant Difference (HSD) test at the 5% significance level.

3. Results

3.1. Phenology

It took an average of three to eight days for radicles to emerge in all Laggera forms (Figure 1). In all studied forms, the vegetative growth was the most prolonged phase, lasting about eight months, starting from mid-September until mid-April. There was phenological diversity in the leaf senescence phase within forms, where Laggera alata with a small capitulum (LA-SC) leaves turned yellow in early July, but this occurred in the middle of June for Laggera alata with a big capitulum (LA-BC). Similarly, great phenological diversity in the leaf senescence phase was recorded within Laggera crispata forms, with Laggera crispata from South Africa (LC-SA) leaves turning yellow in the middle of June, while those of Laggera crispata from Zimbabwe (LC-ZIM) started in early August. The leaves of Laggera forms did not fall, but dried leaves remained attached to the stems. The resprouting of shoots was only detected in September for LA-BC and LC-ZIM (Figure 1). These new plants were regenerated from the previous season’s rootstocks and stems (Figure 2).
Laggera alata forms were the earliest to form flower buds compared with the Laggera crispata forms (Figure 1). Among the studied Laggera forms, LA-BC opened their first flower early (3rd week of April), whereas LC-ZIM opened their first flower late (3rd week of June). The Laggera forms produced seeds in the same month (early June). However, comparisons between Laggera crispata forms indicated that LC-ZIM has late formation of seeds. The LA-BC started to disperse seeds in late July, while in LA-SC, this phenophase started in early August. In L. crispata forms, LC-SA was the earliest to disperse seeds (late July), followed by LC-ZIM a month later (early September).

3.2. Qualitative Traits

The qualitative traits of the studied forms were quite similar at the early and late stages of growth. In addition, the stems were woody at the bottom and bent for the decumbent growth form. However, the stem wings and leaf margins were the significant descriptors for differentiating Laggera alata forms from Laggera crispata forms in the present study (Table 2). Laggera alata forms had entire continuous stem wings, whereas those of Laggera crispata forms were interrupted and toothed. The leaf margins of Laggera alata forms were serrulate, whereas those of Laggera crispata forms were serrate. The young leaves of Laggera alata forms had a maroon pigment on the leaf edges, whereas this pigmentation was not present in Laggera crispata forms. The leaf trichrome stickiness of both Laggera alata forms was less sticky, whereas that of Laggera crispata forms differed, with LC-ZIM being more sticky than LC-SA.
Nevertheless, there were differences within the Laggera forms in terms of plant scent, leaf shape, and leaf color (Table 2). The plant scent of LA-SC was very strong, whereas that of LA-BC was moderately strong. The leaves of LA-SC were oblong and greyish-green, while those of LA-BC were ovate and dark green. Although the Laggera crispata forms had shiny light-green leaves, a lanceolate shape, and emitted a sweet-like plant scent, LC-ZIM leaves assumed a curly-like structure at maturity. The curliness of LC-ZIM leaves became a significant descriptor for differentiating these two L. crispata forms. In contrast, all studied forms shared several qualitative traits, such as the absence of the leaf petiole, acute leaf apex, pink flower color, and white seed color.

3.3. Leaf Size

There were significant differences in leaf length among Laggera forms, and the interaction between Laggera forms and days after transplanting (DAT) was also significant (Table S1). Significant differences in leaf length between forms were only evident at 91 and 105 days after transplant (Table 3). At 91 DAT, LA-BC seedlings had longer leaves (123.73 mm) than LA-SC (107.00 mm), but their leaf lengths were both similar to LC-SA and LC-ZIM. At 105 DAT, LA-BC and LC-ZIM had longer leaves than LA-SC. Furthermore, the leaf lengths of Laggera alata forms differed from each other from 91 to 105 DAT, while those of Laggera crispata forms were similar to each other from 37 to 105 DAT. All forms achieved their maximum leaf length at 91 DAT. The linear regression analysis for leaf length over days after transplanting showed a significant positive correlation for all Laggera forms studied (Figure S1).
Leaf width among Laggera forms differed significantly, and a significant interaction effect between DAT and Laggera forms on leaf width was recorded (Table S2). Variation in leaf width among Laggera forms was only recorded at 37, 91, and 105 DAT (Table 3). The LA-BC had wider leaves than LC-SA at 37 DAT. Leaves of LA-BC were also broader than those of LA-SC and LC-SA at both 91 and 105 DAT. The leaves of LA-SC did not expand as the days after transplanting increased. Leaf widening of LC-SA reached its maximum growth at 51 DAT, whereas the maximum widening of LA-BC and LC-ZIM leaves was at 91 DAT. The consistent increase in leaf width over days after transplanting was recorded in all Laggera forms (Figure S2).
The interaction between DAT and Laggera forms, as well as between plant and Laggera forms, had a significant effect on leaf area (Table S3). The leaves of LA-BC had a bigger leaf area in comparison to the other forms at 91 DAT (Table 3). Nevertheless, the leaf area was not significantly different from LC-ZIM. Once more, LA-BC exhibited the biggest leaf area at 105 DAT when compared with the other forms, yet the difference was not statistically significant when compared with the LC-ZIM. Leaf areas of LA-BC and LC-ZIM were the biggest, whereas those of LA-SC and LC-SA were the smallest. The leaves of LC-SA achieved their peak leaf area at 51 DAT. In contrast, the leaves of both LA-BC and LC-ZIM achieved their peak leaf area at 91 DAT. Unexpectedly, the leaf area of LA-SC remained significantly constant from 37 to 105 DAT. Leaf area demonstrated a positive linear relationship with days after transplanting across all studied forms (Figure S3).

3.4. Leaf Chlorophyll Content

The chlorophyll content significantly differed among species, and the interaction between individual plant and Laggera forms was significant (Table S4). The leaves of LA-SC had a higher chlorophyll content at 37 DAT compared with the other forms (Figure 3). The leaf chlorophyll content of Laggera alata forms differed significantly from 37 to 91 DAT. Among these forms, at 91 DAT, LA-SC had higher leaf chlorophyll content, while it was lower for LA-BC. The leaf chlorophyll content of Laggera crispata forms only differed significantly at 51 DAT, with LC-SA having the higher chlorophyll content and LC-ZIM the lowest. The maximum leaf chlorophyll content for LA-SC and LC-SA was recorded at 51 DAT. Chlorophyll content increased linearly with the number of days after transplanting for all forms (Figure S4).

3.5. Number of Leaves per Plant

The interaction between DAT and Laggera forms had a significant effect on the number of leaves per plant (Table S5). The variation in the number of leaves per plant emerged from 91 DAT, with LC-SA producing more leaves than LA-BC (Figure 4). Again, LC-SA produced the most leaves at 105 DAT, while LA-BC produced the fewest leaves. The maximum number of leaves was produced at 91 DAT for LA-BC, LA-SH, and LC-ZIM. A consistent increase in the number of leaves per plant was recorded for all Laggera forms (Figure S5).

3.6. Stem Diameter and Plant Height

Stem diameter significantly differed among Laggera forms, and individual plants had statistically significant effects on stem diameter (Table S6). Differences in stem diameter within forms emerged at 105 DAT, where LC-ZIM had wider stems than LA-SC (Table 4). At 91 DAT, Laggera alata forms had already reached their mature stem diameter, while Laggera crispata forms reached their mature stem diameter at 105 DAT. Stem diameter showed a positive linear relationship with days after transplanting across all studied Laggera forms (Figure S6).
Plant height significantly differed among Laggera forms (Table S7). Plant height drastically increased from 91 DAT for all forms, compared with 37 and 51 DAT (Figure 5). The plant heights of LA-SC and LA-BC forms were significantly similar at 105 DAT, as were those of LC-SA and LC-ZIM forms. Laggera crispata forms had significantly taller plants, reaching the heights of 874.6 and 833.3 mm, respectively, whereas Laggera alata forms had shorter plants, reaching 692.4 and 662.2 mm, respectively. The linear regression analysis for plant height over days after transplanting showed a significant positive correlation for all Laggera forms studied (Figure S7).

4. Discussion

4.1. Phenology

The prolonged vegetative phase documented in each of the studied Laggera forms (Figure 1) is consistent with the findings from Australia, where the southern New South Wales weed population of Chloris virgata Sw. had the longest vegetative period [23]. These results might indicate that these species are annuals and perennials and tend to produce leaves that have a longer lifespan [24]. Species with long vegetative growth phases may be more tolerant to environmental stress and often develop root systems that can extract water from deep soil profiles [25].
The high phenological diversity for the onset and completion of leaf senescence among forms (Figure 1 and Figure 2) can be ascribed to genetic differences, as leaf senescence is regulated by hundreds of senescence-associated genes [26]. The permanent attachment of leaves to stems after senescence in all studied Laggera forms was a similar phenomenon to that in species of the Lychnophorinae subtribe within the Asteraceae family [27]. The retention of dead leaves on the plant after senescence is called marcescence [28]. Species that retain the marcescent phytomass prefer nutrient-poor habitats, as marcescence postpones the release of nutrients retained in the dead leaves to the plant growth period [29,30]. It can, therefore, be assumed that the studied Laggera forms survive in nutrient-poor soil environments.
In the current study, resprouting was recorded for Laggera alata with a big capitulum (LA-BC) and Laggera crispata from Zimbabwe (LC-ZIM) in the rainy season (September). A comparable study in Kansas reported root resprouting after commercial harvest of a wild medicinal plant, Echinacea angustifolia DC [31]. Considering that resprouting is a key plant functional trait that enables the survival of individuals after a disturbance [32], Laggera forms might also exhibit the capacity for resprouting under similar circumstances. The capacity for spontaneous resprouting highlights the significance of sustainable approaches for harvesting medicinal plants [33].
The present study found that LA-BC and Laggera alata with a small capitulum (LA-SC) began flowering in mid-April and mid-May, respectively. In agreement with these findings, a previous botanic survey found that L. alata flowers year-round [34]. The flowering season for Laggera crispata from South Africa (LC-SA) and LC-ZIM began in mid-May and mid-June, respectively (Figure 1). These findings contradict those of an Indian survey that found that L. crispata flowers from January to April. Possibly, these differences are a result of the different environmental conditions in these two regions. A comparable study conducted in Uganda concluded that flowering in Asteraceae is partly dependent on annual rainfall [7]. Laggera alata forms in the present study were the earliest to form seeds, followed by LC-SA and LC-ZIM. The success of plant reproduction is highly dependent on effective seed dispersal [35]. Seed dispersal varied among Laggera forms in the current study and started from late July to early September. This variation in seed dispersal among and within species may be caused by seed size, plant height, and planting season [36,37].

4.2. Qualitative Traits

Laggera alata forms and/or Laggera crispata forms differed from each other by several morphological traits, but they also shared several morphological traits (Table 2). Intra- and inter-species or forms’ variations and similarities in morphology are brought about by genetic changes, environmental changes, and soil properties [38,39]. The current study found that Laggera forms had winged stems, whereby L. alata forms had entire, continuous stems, and L. crispata forms had interrupted and toothed stems. This is consistent with previous studies on the genus Laggera [14,40].
Considerable variability in plant scent, leaf shape, and leaf color was documented in the present study among L. alata forms. The stronger aroma in LA-SC than in LA-BC probably means that LA-SC have larger glandular trichomes or higher concentrations of aromatic compounds in the plant tissues that contribute to their stronger scent [41]. Laggera alata is known to be strongly aromatic, with a persistent thymol-like and sweet odor [40], but in this study, the sweet-like odor was present in both L. crispata forms. The current findings support a previous study in India, which found L. crispata species to be strongly aromatic [42].
The variation in leaf shape between oblong and ovate within L. alata forms in the present study may be caused by genetic, physiological, and ecological factors [43]. Apparently, intra-species changes in leaf shape may be a visual representation of the plant adaptation to environmental shifts [44]. On the other hand, this study found that the leaves of both L. crispata forms were lanceolate, with LC-ZIM leaves being particularly curly at maturity. Oblong- and ovate-shaped leaves exhibited higher photosynthetic efficiency and water use efficiency than lanceolate leaves [44].
The leaf color of LA-SC in the present study was greyish-green and comparable with that of Laggera tomentosa [40]. The dark green leaves in LA-BC and shiny light-green leaves in L. crispata forms probably showed variations in the pigment concentration of leaves, which resulted in variations in their color [45]. Glandular trichomes are hair-like structures found on the surface of vegetative and reproduction organs that produce secondary metabolites that are responsible for the stickiness of plant organs [46,47,48]. In the present study, leaf trichomes of LC-ZIM were the stickiest, followed by LC-SA, with LA-SC and LA-BC having the least. Similar studies have reported that L. crispata has sticky hair [42]. These sticky compounds on the leaf surfaces function as a direct defense and repel or kill antagonists [49]. The differences in leaf stickiness may also be attributed to secondary metabolites present in these plants [48].
Similarities between the Laggera forms, particularly in leaf petiole, leaf orientation, and the leaf apex (Table 2), indicate that these are some of the morphological traits that are used to taxonomically place these plants in the genus Laggera [50]. Pink flowers in all studied Laggera species and forms support that Laggera species have florets with pink or purple corollas [12]. In contrast, a study conducted in Nigeria reported that Laggera crispata has white flowers [51].

4.3. Leaf Traits

Leaf length and width varied significantly among all Laggera forms at 91 DAT in the present study. The leaves of LA-BC were the biggest when compared with other forms and ranged between 123.73 mm in length and 60.60 mm in width. In contrast, a much bigger maximum length of 160 mm and width of 70 mm was recorded for L. alata leaves in the flora Zambesiaca [13]. On the other hand, the leaf length and width of L. crispata fell within a range of 30–270 mm long and 8–110 mm wide [13]. These differences in leaf size are possibly caused by the trade-off between the leaf size and the number of leaves produced [52].
In the present study, LA-BC leaves exhibited the highest leaf area, significantly higher than the other three forms, while the leaf area of LA-SC was the lowest among the four. Variation in leaf area may be caused by temperature, aridity, and soil pH [53]. Larger leaves have a lower rate of heat exchange, and are thus more adapted to cooler, moister, and lower-irradiance environments [52,53,54]. Consequently, LA-BC can be well adapted to mesic environments. However, larger leaves might be more susceptible to herbivory [55]. Low leaf area is advantageous in low-resource or harsh situations; therefore, LA-SC may thrive in hot, dry, or nutrient-poor soil environments [56]. The number of leaves per plant is an essential trait for germplasm characterization [57]. This study found that LA-SC produced more leaves per plant than LA-BC at 91–105 DAT. This is possibly caused by the relatively small leaf size of LA-SC that results in a trade-off of high leafing intensity [58,59].
In the present study, LA-SC leaves had the highest chlorophyll content at 37, 51, and 91 DAT compared with the other forms, and it ranged from 41.86 to 51.97 mg cm−2. A comparable study in Iran reported a leaf chlorophyll content of 0.063 mg/L for an Asteraceae perennial herb [60]. Similarly, the average total chlorophyll content of 10 medicinal plants from the Asteraceae family recorded a chlorophyll content of 8.61 mg/g [61]. The present study used a non-destructive method, compared with the destructive extraction method used in the other two studies, hence the noted significant difference between the three [62]. The variation in leaf chlorophyll content in the current study might also have been caused by atmospheric and soil factors [44], because chlorophyll synthesis requires various elements, such as nitrogen and phosphorous, from the soil [63].

4.4. Stem Diameter and Plant Height

Significant variation in stem diameter between Laggera forms at 105 DAT (Table 4) might be caused by irreversible growth due to cell growth and enlargement [64]. The range of Laggera crispata forms’ plant height from 833 to 875 mm (Figure 5) were within the plant height range of 600–2400 mm reported for L. crispata in the flora Zambesiaca [13]. The current findings differ from what was reported in Côte d′Ivoire, where L. crispata had a maximum height of 1700 mm [65]. This study found that the maximum plant height for L. alata forms ranged between 630 and 686 mm, which is shorter than the plant height of 2000 mm for L. alata recorded in flora Zambesiaca [13]. This inconsistency was possibly caused by the cultivated L. alata population in the present study, compared to the naturally occurring population in flora Zambesiaca. Morphometric differences in wild-growing and cultivated individuals are due to the domestication process [66]. Also, in the present study, a decumbent growth form was recorded for all forms, which may negatively affect the plant height.

5. Conclusions

This study revealed significant diversity in the phenology, qualitative traits, leaf traits, plant height, and stem diameter within and between L. crispata forms and/or L. alata forms. This study was effective in determining the onset dates of vegetative and reproductive phenophases of seed-grown Laggera forms. These findings could be used to introduce Laggera forms into the farming systems for medicinal and/or commercial purposes. Laggera alata with a small capitulum (LA-SC) had a shorter vegetative stage, whereas Laggera crispata with a big capitulum had early flower bud formation compared to all other forms. Therefore, the farming process of these two L. alata forms can be more efficient and cost-effective. Although Laggera crispata from Zimbabwe had prolonged phenophases compared to other forms, it showed potential as an ideal plant for high essential oil yield extraction due to its very-sticky plant parts. However, future studies are needed to determine which Laggera form has a high essential oil content. Based on morphological traits, stem wing shape and leaf margins were the major traits used to classify L. alata and L. crispata. Morphometric traits, such as leaf size and number of leaves per plant, are the major descriptors that can be used to differentiate the L. alata forms. However, it is still unclear whether Laggera alata forms are varieties or hybrids. Molecular work studies should be conducted to clarify the taxonomic status of these types.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/d16080466/s1, Table S1: Summary of ANOVA results for leaf length; Table S2: Summary of ANOVA results for leaf width; Table S3: Summary of ANOVA results for leaf area; Table S4: Summary of ANOVA results for chlorophyll content; Table S5: Summary of ANOVA results for number of leaves; Table S6: Summary of ANOVA results for Stem diameter; Table S7: Summary of ANOVA results for plant height; Figure S1: Linear regression between leaf length and days after transplanting for (a) Laggera alata with a big capitulum, (b) Laggera alata with a small capitulum, (c) Laggera crispata from South Africa, and (d) Laggera crispata from Zimbabwe; Figure S2: Linear regression between leaf width and days after transplanting for (a) Laggera alata with a big capitulum, (b) Laggera alata with a small capitulum, (c) Laggera crispata from South Africa, and (d) Laggera crispata from Zimbabwe; Figure S3: Linear regression between leaf area and days after transplanting for (a) Laggera alata with a big capitulum, (b) Laggera alata with a small capitulum, (c) Laggera crispata from South Africa, and (d) Laggera crispata from Zimbabwe; Figure S4: Linear regression between chlorophyll content and days after transplanting for (a) Laggera alata with a big capitulum, (b) Laggera alata with a small capitulum, (c) Laggera crispata from South Africa, and (d) Laggera crispata from Zimbabwe; Figure S5: Linear regression between number of leaves per plant and days after transplanting for (a) Laggera alata with a big capitulum, (b) Laggera alata with a small capitulum, (c) Laggera crispata from South Africa, and (d) Laggera crispata from Zimbabwe; Figure S6: Linear regression between stem diameter and days after transplanting for (a) Laggera alata with a big capitulum, (b) Laggera alata with a small capitulum, (c) Laggera crispata from South Africa, and (d) Laggera crispata from Zimbabwe; Figure S7: Linear regression between plant height and days after transplanting for (a) Laggera alata with a big capitulum, (b) Laggera alata with a small capitulum, (c) Laggera crispata from South Africa, and (d) Laggera crispata from Zimbabwe.

Author Contributions

Conceptualization, N.N.N., N.R.N., G.E.Z., C.Z. and B.S.; methodology, G.E.Z. and N.N.N.; software, N.N.N. and N.R.N.; validation, N.N.N., N.R.N., G.E.Z., C.Z. and B.S.; formal analysis, N.N.N.; investigation, N.N.N.; resources, G.E.Z.; data curation, N.N.N.; writing—original draft preparation, N.N.N.; writing—review and editing, N.R.N.; visualization, N.N.N.; supervision, N.R.N., G.E.Z., C.Z. and B.S.; project administration, N.N.N.; funding acquisition, N.R.N. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Institutional Review Board Statement

Not applicable.

Data Availability Statement

The data is contained within the manuscript and Supplementary Materials.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. Phenophases of studied Laggera alata (A) and Laggera crispata (B) forms. The color-shaded rows indicate the whole period of appearance of different phenophases.
Figure 1. Phenophases of studied Laggera alata (A) and Laggera crispata (B) forms. The color-shaded rows indicate the whole period of appearance of different phenophases.
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Figure 2. Resprouting of shoots in Laggera alata with a big capitulum (A) and Laggera crispata from Zimbabwe (B).
Figure 2. Resprouting of shoots in Laggera alata with a big capitulum (A) and Laggera crispata from Zimbabwe (B).
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Figure 3. Leaf chlorophyll content of Laggera forms at different days after transplanting. LA-BC = Laggera alata with big capitulum; LA-SC = Laggera alata with small capitulum; LC-SA = Laggera crispata from South Africa; LC-ZIM = Laggera crispata from Zimbabwe. Means followed by different letter(s) within bars differ significantly (p < 0.05) according to Tukey’s Honestly Significance Difference (HSD) test.
Figure 3. Leaf chlorophyll content of Laggera forms at different days after transplanting. LA-BC = Laggera alata with big capitulum; LA-SC = Laggera alata with small capitulum; LC-SA = Laggera crispata from South Africa; LC-ZIM = Laggera crispata from Zimbabwe. Means followed by different letter(s) within bars differ significantly (p < 0.05) according to Tukey’s Honestly Significance Difference (HSD) test.
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Figure 4. Number of leaves per plant in Laggera forms on different days after transplanting. LA-BC = Laggera alata with big capitulum; LA-SC = Laggera alata with small capitulum; LC-SA = Laggera crispata from South Africa; LC-ZIM = Laggera crispata from Zimbabwe. Means followed by different letter(s) within bars differ significantly (p < 0.05) according to Tukey’s Honestly Significance Difference (HSD) test.
Figure 4. Number of leaves per plant in Laggera forms on different days after transplanting. LA-BC = Laggera alata with big capitulum; LA-SC = Laggera alata with small capitulum; LC-SA = Laggera crispata from South Africa; LC-ZIM = Laggera crispata from Zimbabwe. Means followed by different letter(s) within bars differ significantly (p < 0.05) according to Tukey’s Honestly Significance Difference (HSD) test.
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Figure 5. Plant height in Laggera forms at different days after transplanting. DAT = days after transplanting; LA-BC = Laggera alata with big capitulum; LA-SC = Laggera alata with small capitulum; LC-SA = Laggera crispata from South Africa; LC-ZIM = Laggera crispata from Zimbabwe. Means followed by different letter(s) within bars differ significantly (p < 0.05) according to Tukey’s Honestly Significance Difference (HSD) test.
Figure 5. Plant height in Laggera forms at different days after transplanting. DAT = days after transplanting; LA-BC = Laggera alata with big capitulum; LA-SC = Laggera alata with small capitulum; LC-SA = Laggera crispata from South Africa; LC-ZIM = Laggera crispata from Zimbabwe. Means followed by different letter(s) within bars differ significantly (p < 0.05) according to Tukey’s Honestly Significance Difference (HSD) test.
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Table 1. Weather data recorded during the first and second seasons of the experiment.
Table 1. Weather data recorded during the first and second seasons of the experiment.
Month 2021/2022 Season 2022/2023 Season
TMXTMNTtRHPRAINSRADTMXTMNTtRHPRAINSRAD
Sept.26.115.210.758134.616.32816.612.35947.916.6
Oct.25.915.610.867102.716.129.618.914.36292.816.9
Nov.27.618.112.9654319.52818.613.367358.518.7
Dec.29.520.114.868194.918.929.420.114.87019419
Jan.31.120.715.966102.423.432.620.616.65786.426.1
Feb.31.421.116.36594.822.929.821.415.678246.917.4
Mar.30.319.715.062102.719.63020.615.36982.718.1
Apr.2617.411.77435612.828.717.713.26234.415.8
May25.415.310.46466.112.925.71610.968128.610.6
Jun.2411.87.95471.411.725.513.29.4541011.8
Jul.25.513.49.55367.311.424.112.38.25681.711.3
Aug.25.213.89.55435.713.525.213.79.5574.614.1
Sept.2816.612.35947.916.628.215.812.05437.916.7
Mean27.416.812.162.2109.216.628.117.412.762.5108.216.4
TMX (maximum temperature (°C)), TMN (minimum temperature (°C)), Tt (thermal time), RHP (relative humidity PH (%)), RAIN (rainfall (mm)), and SRAD (solar radiation (MJ/m2/d)).
Table 2. Qualitative traits of Laggera alata forms and Laggera crispata forms based on field observations.
Table 2. Qualitative traits of Laggera alata forms and Laggera crispata forms based on field observations.
Qualitative TraitsLA-SCLA-BCLC-SALC-ZIM
StemProfusely branched, continuous, and entire wings Profusely branched, continuous, and entire wingsProfusely branched, interrupted, and toothed wings Profusely branched, interrupted, and toothed wings
Young leavesMaroon pigment on the leaf edges Maroon pigment on the leaf edgesNo pigmentationNo pigmentation
Leaf petioleAbsentAbsentAbsentAbsent
Leaf orientationAlternate AlternateAlternateAlternate
Plant scent Very strongModerately strongSweet-like odorSweet-like odor
Leaf shapeOblongOvateLanceolateLanceolate and curly
Leaf apexAcuteAcuteAcuteAcute
Leaf colorGreyish greenDark greenShiny light greenShiny light green
Leaf trichome stickinessLess stickyLess stickyModerately stickyVery sticky
Leaf marginsSerrulateSerrulateSerrateSerrate
Flower color PinkPinkPinkPink
Seed colorWhiteWhiteWhiteWhite
LA-BC = Laggera alata with big capitulum; LA-SC = Laggera alata with small capitulum; LC-SA = Laggera crispata from South Africa; LC-ZIM = Laggera crispata from Zimbabwe.
Table 3. Leaf size variability in Laggera forms.
Table 3. Leaf size variability in Laggera forms.
Leaf TraitsDATLA-BCLA-SCLC-SALC-ZIM
Leaf length (mm)3796.20 def91.80 ef83.80 f89.40 ef
5199.80 de99.20 de97.00 def101.47 cde
91123.73 a107.00 bcd118.73 ab120.73 ab
105126.80 a103.40 cde115.40 abc125.20 a
Leaf width (mm)3743.20 de40.00 def34.73 f37.33 ef
5145.93 cd44.73 de45.47 d44.60 de
9160.60 ab40.73 def44.33 de53.47 bc
10562.80 a37.20 ef45.27 d55.53 ab
Leaf area (mm2)372085 def1865 ef1457 f1678 ef
512309 de2207 de2204 de2246 de
913720 ab2161 de2603 cd3212 bc
1053964 a1915 def2583 cd3457 ab
DAT = days after transplanting; LA-BC = Laggera alata with big capitulum; LA-SC = Laggera alata with small capitulum; LC-SA = Laggera crispata from South Africa; LC-ZIM = Laggera crispata from Zimbabwe. Means followed by different letter(s) within columns and rows differ significantly (p < 0.05) according to Tukey’s Honestly Significance Difference (HSD) test.
Table 4. Stem diameter of Laggera forms on different days after transplanting.
Table 4. Stem diameter of Laggera forms on different days after transplanting.
DATLA-BCLA-SCLC-SALC-ZIM
370.47 e0.42 e0.36 e0.36 e
510.54 e0.52 e0.45 e0.47 e
911.17 bcd1.02 cd1.00 d1.17 bcd
1051.32 ab1.20 bc1.29 ab1.45 a
DAT = days after transplanting; LA-BC = Laggera alata with big capitulum; LA-SC = Laggera alata with small capitulum; LC-SA = Laggera crispata from South Africa; LC-ZIM = Laggera crispata from Zimbabwe. Means followed by different letter(s) within columns and rows differ significantly (p < 0.05) according to Tukey’s Honestly Significance Difference (HSD) test.
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Nkosi, N.N.; Zharare, G.E.; Zimudzi, C.; Stedje, B.; Ntuli, N.R. Variation in Phenology and Morphological Traits of Seed-Propagated Laggera alata and Laggera crispata Forms. Diversity 2024, 16, 466. https://doi.org/10.3390/d16080466

AMA Style

Nkosi NN, Zharare GE, Zimudzi C, Stedje B, Ntuli NR. Variation in Phenology and Morphological Traits of Seed-Propagated Laggera alata and Laggera crispata Forms. Diversity. 2024; 16(8):466. https://doi.org/10.3390/d16080466

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Nkosi, Noluthando Nonjabulo, Godfrey Elijah Zharare, Clemence Zimudzi, Brita Stedje, and Nontuthuko Rosemary Ntuli. 2024. "Variation in Phenology and Morphological Traits of Seed-Propagated Laggera alata and Laggera crispata Forms" Diversity 16, no. 8: 466. https://doi.org/10.3390/d16080466

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