Next Article in Journal
Evaluation of Two Different Methods of Fulvic Acid Application (Seed Priming and Foliar Spray) on Growth, Yield, and Nutritional Quality of Pea (Pisum sativum L.)
Next Article in Special Issue
Phylogenetic Relationships in the Miracle Berry Genus, Synsepalum, Sensu Lato, and Relatives (Sapotaceae)
Previous Article in Journal
Pseudoplagiostoma Causing Leaf Spot Disease in Key Tropical Fruit Crops in Thailand
 
 
Search for Articles:
Title / Keyword
Author / Affiliation / Email
Journal
Article Type
 
 
Advanced Search
 
Section
Special Issue
Volume
Issue
Number
Page
 
Journals
Plants
Volume 13
Issue 23
10.3390/plants13233378
plants-logo
Submit to this Journal Review for this Journal Propose a Special Issue
► Article Menu

Article Menu

share Share announcement Help format_quote Cite question_answer Discuss in SciProfiles
first_page
settings
Order Article Reprints
Font Type:
Arial Georgia Verdana
Font Size:
Aa Aa Aa
Line Spacing:
Column Width:
Background:
Article

Study on the Genus Drymaria (Caryophyllaceae)—A New Species from North-East India

by
Sindhu Arya
1,
Harsh Singh
2,
Kalarikkal Walsan Vishnu
3 and
Duilio Iamonico
4,*
1
PG & Research Department of Botany, PSG College of Arts & Science, Tamil Nadu 641014, India
2
Department of Botany, Centre for Advanced Studies in Botany, North-Eastern Hill University, Shillong 793022, India
3
Medicinal Plant Research Section, Regional Ayurveda Research Centre, Dimapur 797112, India
4
Department of Environmental Biology, University of Rome Sapienza, Piazzale Aldo Moro 5, 00185 Rome, Italy
*
Author to whom correspondence should be addressed.
Plants 2024, 13(23), 3378; https://doi.org/10.3390/plants13233378
Submission received: 16 October 2024 / Revised: 22 November 2024 / Accepted: 25 November 2024 / Published: 30 November 2024
(This article belongs to the Special Issue Plant Taxonomy, Phylogeny, and Evolution)
Browse Figures
Versions Notes

Abstract

:
A new species of Drymaria from the Kohima District (Nagaland of North-East India) is described and illustrated based on both molecular data (the ITS region of nuclear DNA) and morphometric analyses (clustering, PCA, DA, and MANOVA). The new species resembles D. cordata but differs with respect to the shape of the sepal (oblong, incurved at the margin vs. lanceolate, not incurved at the margin), petals (oblong or linear vs. ovate–lanceolate), and bract (foliaceous, glabrous, non-prominent vs. non-foliaceous, pubescent, and prominent). For nomenclatural purposes, the typification of Drymaria villosa by Duke (in 1961) was corrected according to Art. 9.10 of the Shenzhen Code.

1. Introduction

Caryophyllaceae Juss. is one of the most diverse families in the order Caryophyllales, comprising approximately 100 genera and 3000 species, and is mainly distributed in north-temperate, montane, and alpine areas with a center of diversity in the Eastern Mediterranean and Irano-Turanian regions, while its presence in the tropics and the Southern Hemisphere is limited and mostly at higher elevations [1,2,3]. The family is monophyletic, as circumscribed by Bittrich [4], but the traditional classification into three subfamilies [4,5], which is based on stipule, petal, sepal, and fruit features, does not provide monophyletic groups and should be replaced with the tribe-based scheme [1,2]. According to The Caryophyllales Network [6], 11 tribes can be recognized at the moment: Alsineae Lam. & DC. (with 15 genera), Arenarieae Kitt. (3 genera), Caryophylleae Lam. & DC. (14 genera), Corrigioleae Dumort. (2 genera), Eremogoneae Rabeler & W.L.Wagner (3 genera), Paronychieae Dumort. (5 genera), Polycarpaeae DC. (25 genera), Sagineae J.Presl (14 genera), Sclerantheae DC. (10 genera, including the recently described genus Maguirellaria Iamonico [7]), Sileneae DC. (9 genera), and Sperguleae Dumort. (3 genera); a further 4 genera (Calycotropis Turcz., Dadjoua Parsa, Dolophragma Fenzl, and Pentastemonodiscus Rech.f.) are unplaced.
According to phylogenetic analyses of the whole family [1,2,7], the genus Drymaria Willd. ex Schultes belongs to the tribe Polycarpeae, a well-supported clade which is basal to a large group including many other tribes of Caryophyllaceae, i.e., Sperguleae, Sclerantheae/Sagineae, and Ermogoneae/Caryophylleae/Sileneae. This genus is represented by 57 species native mainly to tropical and subtropical regions of the world [8].
Drymaria is a poorly known genus. In fact, the only comprehensive taxonomic revision was published about 70 years ago by Duke [9], who recognized 48 species and proposed a classification of the genus by recognizing 17 series. Hartman [10], Villarreal and Estrada [11], and Montesinos-Tubeé et al. [12] stated that Duke’s series were not validly published, being nomina nuda. However, by checking Duke’s paper, it transpires that each series is in fact valid from a nomenclatural point of view (see Art. 38.1 of the Shenzhen Code [13]), having an own description or, for monospecific series, a description of the single species [9] (pagg. 186, 189, 193, 200, 201, 204, 210, 212, 216, 220, 222, 223, 230, 240, 245). No molecular study on all the species belonging to Drymaria has been published until now.
The flora of India includes three Drymaria species, i.e., D. cordata (L.) Willd. ex Schult. [alien in Haryana State (N-India [14]), Assam (NE India [15,16]), Upper Siang District in Arunachal Pradesh (NE India [17])], D. diandra Blume (hillock regions of India, from the Western to the Eastern Himalaya [18]), and D. villosa Schltdl. & Cham. (Eastern, Western, and Central Himalaya and Southern India [18,19,20]), all occurring in the Kerala region.
As a part of ongoing studies of the family Caryophyllaceae in India (see, e.g., [21,22]), we found an interesting population of Drymaria from Kohima District (Nagaland, NE India). Based on both morphometric and molecular analyses, as well as a critical examination of the literature, we reached the conclusion that the population from Nagaland can be described as a species new to science.

2. Material and Methods

2.1. Morphometric Data

The work was carried out via field surveys in Nagaland during the monsoon season (August to November) of the years 2022–2023, examination of herbarium specimens housed at BM, CALI, CAL, P, UCBD, and TBGT (acronyms according to [23]), and analysis of the relevant literature (cited below throughout the text).
The data matrix [characters (columns) × individuals (=specimens; rows)] was processed by means of the software NCSS 2007 ver. 01.1.21 (Kaysville, UT, USA). Seven populations, 14–19 individuals per population (we were not able to collect more individuals per population since they were too reduced) were investigated (Table 1) and 21 characters (1 discrete, 20 continuous; Table 2) were measured in 90 individuals (30 for each native species, i.e., Drymaria diandra, D. villosa, and the new population, hereafter named “Drymaria sp.”). After a preliminary recognition of the morphological characters of Drymaria throughout its distribution area (herbarium specimens), we decided to collect populations in Nagaland State, where more similarities between them were expected. An explorative cluster analysis was performed by the UPGMA method. The data matrix was also submitted to ordination (Principal Component Analysis, PCA; a correlation matrix of only continuous characters) and Discriminant Analysis (DA). The DA was performed using the first three components derived from the PCA, which explain about 85% of the total variability. As a supervised technique, we performed the DA on groups classified as species; the matrix of actual/predicted groups was analyzed by comparing the values among these groups, especially regarding the diagonal, whose values reveal the matching of actual and predicted observations for each group. The value of correct classification reported in the results is the classification accuracy achieved by the actual discriminant functions over what is expected. A multivariate analysis of variance (MANOVA) was also performed to test the significance of differences between response (=dependent) variables (morphological characters) and factor variables (=taxonomic groups).
Photographs of specimens were taken using a Canon 1500 camera and, for macromorphology, they were taken using a Carl Zeiss camera. Seed and pollen grains were examined using SEM (Carl Zeiss, Oberkochen, Germany).

2.2. Molecular Analysis

Fresh leaf samples weighing 50 mg were crushed in liquid nitrogen using a sterile mortar and pestle, yielding a fine powder. DNA extraction was carried out using a Plant/Fungi DNA isolation kit or the CTAB method [24]. PCR reactions were carried out using a Thermo Fisher thermal cycler (Applied BioSystems, Waltham, MA, USA) under the following conditions: initial denaturation at 94 °C for 5 min, followed by 40 cycles of denaturation at 94 °C for 15 s, annealing at 60 °C for 20 s and extension at 72 °C for 1:00 min, with a final extension at 72 °C for 5 min. The PCR reactions were performed in a 20 μL volume beaker containing Emarald PCR Master mix at 1× final concentration [25], 500 ng of genomic DNA, and 0.2 micromoles of primers. The ITS region was used for phylogenetic analysis of the new species.
The obtained sequences were compared with other available sequences in the NCBI database using the Basic Local Alignment Search Tool (BLAST) [26]. DNA sequences of the rbcL region were submitted to NCBI/GENBANK (Table 3). Molecular phylogenetic analysis for the rbcL gene was carried out by using the MEGA X software var. 11.0.10. The phylogenetic tree was constructed using the neighbor joining (NJ) method with a Jukes–Cantor substitution model [27], selecting bootstrap values with the resampling number set to 1000 [28] to view and download the constructed phylogenetic tree. All positions containing gaps and missing data were excluded during phylogenetic analysis.
We considered the molecular studies by Harbaugh et al., Greenberg & Donoghue, and Dillenberger & Kadereit [1,2,7] to select genera related to Drymaria, i.e., the following members of the tribe Polycarpeae: Achyronychia cooperi A.Gray, Cardionema ramosissimum (Weinm.) A.Nelson & J.F.Macbr., Dicheranthus plocamoides Webb, Illecebrum verticillatum L., Loeflingia hispanica L., Sphaerocoma aucheri Boiss., Spergula arvensis L., Spergularia media (L.) C.Presl, Spergularia rubra (L.) J.Presl & C.Presl, Polycarpon tetraphyllum L. subsp. tetraphyllum, Pteranthus dichotomus Forssk., Pycnophyllum bryoides (Phil.) Rohrb., Pycnophyllum spathulatum Mattf., and Scopulophila rixfordii (Brandegee) Munz & I.M.Johnst. Further, other groups were selected based on Figure 1B–D by Greenberg and Donoghue [2], i.e., the generitypes of Dianthus (Dianthus caryophyllus L. from the tribe Caryophylleae), Stellaria L. (Stellaria holostea L. for Alsineae Lam. & DC.), Arenaria L. (Arenaria serpyllifolia L. for Arenarieae Kitt.), Sleranthus L. (Sleranthus annuus L. for Slerantheae), Sagina L. (Sagina procumbens L. for Sagineae), and Silene L. (Silene gallica L. for Sileneae).

3. Results and Discussion

3.1. Morphometric Data

The analyses of the characters allow the recognition of three distinct groups.
Concerning the hierarchical clustering (Figure 1), the cutoff dissimilarity point (1.35) defines three clusters, with each one comprising specimens referring to single species, i.e., Drymaria diandra, D. villosa, and Drymaria sp.
The PCA analysis shows that the cumulative percentage of eigenvalues for the first two axes is 78.76%, with the contribution given by the first component being 58.06% and that by the second component being 20.70%. The component1 vs. component2 graphs show three well-separated groups along the first component, whereas, along component2, two groups are partially overlapped (Figure 2). These three groups correspond to the populations of Drymaria diandra, D. villosa, and Drymaria sp., as displayed by the clustering. The highest contribution to the first component is given by the following characters: length and width of fruits, length and width of sepals, and length of stamens. The second component is mainly related to the length and width of gynoecium, length of petiole, seed diameter, width of petals, and length of bracts.
The DA, carried out using the names of the species (three groups: Drymaria diandra, D. villosa, and Drymaria sp.), predicted three clearly separated groups (Figure 3) based on the first two discriminant functions, which explain 100.0% of the total variation [eigenvalues: 79.7% (first function) and 20.3% (second function)]. These groups correspond to D. diandra, D. villosa, and Drymaria sp.
The results of the MANOVA show significant differences at both species and population levels. The probability level is less than 0.000001 for all the statistical tests considered (Wilks’ lambda, Hotelling–Lawley trace, Pillai’s trace, and Roy’s largest root). F-ratios are high, ranging from F = 27.03 to 2020.7 (populations as groups; Table 4) and F = 1639.56 to 4011.09 (species as groups; Table 5).

3.2. Molecular Analysis

The molecular sequencing ITS region of the nuclear DNA supported Drymaria sp. as distinct from the other allied species of Drymaria (Figure 4). The new species is separated in the second clade, which comprises D. cordata, D. mollugineae, D. grandulosa, and D. laxiflora.
A total of 658 aligned bp of the ITS sequence were considered for comparing Drymaria sp. with D. cordata. From this, 15 sites distinguished Drymaria sp. from D. cordata. The out groups are also well separated in the dendrogram. Thus, the results of phylogenetic analysis also support the status of Stellaria mcclintockiae as a distinct species.

3.3. Taxonomic Treatment

Both the morphological and molecular results obtained in the present research demonstrated that the Drymaria populations growing in Kohima District (Nagaland region, NE India) are different to the other Indian Drymaria species. Consequently, here, we propose describing the population of Drymaria as a new species for science. A complete taxonomic treatment of Drymaria taxa occurring in India follows.
1. 
Drymaria anilii S. Arya, Harsh & Iamonico, sp. nov. (Figure 5).
Holotype: INDIA. Nagaland: Kohima District, Kisama Village, 1800 m a.s.l., 25°37′35″ N 94°60′48.9″ E, 20 October 2021, Arya 730 (DMP!; isotypes: DMP!, RO!).
Diagnosis. Drymaria anilii resembles D. diandra, from which it differs in terms of inflorescence (sessile in a single terminal flower vs. cymose with 10–15 flowers), bracts (foliaceous, glabrous, 2 cm long vs. non-foliaceous, pubescent, less than 5 mm long) pedicel (glabrous vs. glandular), sepal (obtuse or rounded at the apex, incurved at the margin vs. acute at the apex, not incurved at the margin), petal (obtuse or rounded at the apex and linear vs. acute at the apex and ovate), fruit (ellipsoidal, 6 mm long vs. ovate, 3 mm), pollen (spherical with depressed non-prominent pores vs. ellipsoid with prominent pores), and seed surface architecture (small mamillated spots which form a star-like radiating pattern vs. large spinulate spots that cross each other).
Description (macromorphology). An annual herb (therophyte) which is slender, branching from the base, and 10–12 cm long. The root is thin, producing many short horizontal spreading branches, sometimes originating from the lower stem nodes. The stem is prostrate or ascending and erect pubescent with glandular multiseriate trichomes. The leaves are opposite, orbicular to cordate (1.3–2.1 cm × 1.2–1.5 cm), glabrous in the adaxial side, and sparsely pubescent on the abaxial surface; the apex is rounded or acute, and the base is cordate; the petioles are 1–5 mm long, hairy; and the stipules are minute, at 0.1 mm long. There are inflorescences in the reduced cyme (up to three-flowered); the peduncles are 1.5–2.5 mm long and hairy; the bracts are linear to lanceolate and 2 mm long; the pedicels of the flowers are 1–2 mm long and sparsely pubescent; the five sepals are, 1.2–1.4 × 0.6–0.7 mm long, lanceolate, and the apex is acute or obtuse and hairy; the five petals are 1.2–1.3 mm long and bifid (rounded lobes); the stamens are five fewer in number than the sepals with a 0.5 mm long filament and oblong, yellow-brown anthers; the staminode is absent; the ovary is ovoid or globose and 0.7–0.8 mm, and the style is 0.4–0.6 mm long and parted into four at the apex with glands. The capsules are ovoid, 1.7–1.9 mm long, equaling or slightly exceeding the sepals, and opening by four valves to the base. The seeds are brown and reniform.
Description (micromorphology; Figure 6). The pollen grains are spheroidal, poly-pantoporate, and 8.16–10.64 µm in diameter, with 5–6 visible pores. The pore margin or aperture is well demarcated, at about 2 µm. The pores are 3.2–3.3 µm in size. The surface is micro-echinate; the echinates are blunt or appear like dots which are fused. Ektexinous bodies are present inside the pore, minute, and number 4–8 fused, but the apex is sharp and free.
The seeds are reniform with a circular base (0.548–0.765 × 0.614–0.761 mm). The margin is wavy, and the tubercles are short and prominent; small mamillated spots which form a star-like radiating pattern are visible on the surface. The dots form a “V” shape that do not cross each other.
Etymology. The species is named in honor of Dr. V.S. Anil Kumar, Principal of Government College Kasargode, Kerala, India, in recognition of his outstanding contributions to the field of plant taxonomy of the Western Ghats. Dr. Anil Kumar is acknowledged as a great teacher and motivator in the field.
Habitat distribution (Figure 7) and phenology. Drymaria anilii is known from only the locus classicus (Kohima, Nagaland, NE India at an elevation of 1800 m a.s.l.), where it grows along the muddy slope. Each population spreads over an area of less than 5 km. Flowering and fruiting times are June to November.
Conservation status. Even though the plant propagates through vegetative propagation (rooting at the nodes), the number of individuals is very small and they are vulnerable to the severe grazing. There are possibilities of the species occurring in other parts. More extensive studies on the species are required to reach a conclusion on its conservation status. Therefore, by following the IUCN Red List criteria [29], Drymaria anilii is assessed as Data Deficient (DD).
Taxonomic notes. The three Drymaria species occurring in India (D. cordata, D. diandra, and D. villosa) usually have a restricted distribution area and can be found across the Himalayan belts and the hilltops of the Western and Eastern Ghats. Based on the morphology of D. anilii, especially its stipulate leaves, terminal solitary cyme, bifid petals, 2–5 stamens, usual three styles, and one or many seeds, it would belong to the series Cordatae [9] (p. 245). The series is represented by five species, four subspecies, and two varieties in Duke’s monograph (Drymaria gracilis Schltdl. & Cham., D. gracilis subsp. carinata (Brandegee) A.J.Duke, D. glandulosa C.Presl var. glandulosa, D. glanduosa var. galeottiana (Briq.) A.J.Duke, D. xerophylla A.Gray, D. ladewih Rusby, D. cordata (L.) Willd. ex Roem. & Schult. and D. diandra Blume). All these mentioned species, except for D. cordata and D. diandra, have a distribution in the Americas (the USA, Argentina, and Mexico [8]). Note, however, that a comprehensive molecular study to demonstrate that Duke’s classification is natural is lacking. Consequently, the inclusion of D. anilli in the Ser. Cordatae is preliminary and deserves further investigations.
2. 
Drymaria cordata (L.) Willd. ex Schult., in J.J.Roemer & J.A.Schultes, Syst. Veg., ed. 15[bis]. 5: 406 (1819) ≡ Holosteum cordatum L., Sp. Pl. 1: 88 (1753) ≡ Cerastium cordatum (L.) Crantz in Inst. Rei Herb. 2: 400 (1766).
Lectotype [designated by Burger in Cafferty and Jarvis [30] (p. 1052): [Icon] Alsine americana nummulariae folio [31] (t. 11)]; image available at https://bibdigital.rjb.csic.es/viewer/13586/?offset=#page=35&viewer=picture&o=bookmark&n=0&q= (accessed on 26 November 2024).
3. 
Drymaria diandra Blume, Bijdr. Fl. Ned. Ind.: 62 (1825) ≡ Drymaria cordata subsp. diandra (Blume) J.A.Duke in Ann. Missouri Bot. Gard. 48: 253 (1961).
Lectotype [designated by Mizushima [32] (p. 81)]: INDONESIA. Java, in paludosis montanis, October, Blume 1549 (L99-143-199, non vidi fide Mizushima 1957: 81); isotypes: BM000946422! (image available at https://data.nhm.ac.uk/object/0db637d6-3591-4232-b01a-f014f2715810/1704153600000, accessed on 26 November 2024), MO216778! (image available at https://plants.jstor.org/stable/viewer/10.5555/al.ap.specimen.mo-216778, accessed on 26 November 2024).
Note. The isotypes listed above (BM000946422 and MO216778) were not reported by neither Mizushima [32] (p. 81) nor Duke [5] (p. 253).
4. 
Drymaria villosa Cham. & Schltdl., Linnaea 5(2): 232–233 (1830) subsp. villosa ≡ Drymaria cordata var. villosa (Cham. & Schltdl.) Rohrb. in C.F.P.von Martius & auct. suc. (eds.), Fl. Bras. 14(2): 260 (1872).
Lectotype [designated by Duke [9] (p. 226)] as “isotype”, corrected here according to Art. 9.10 of the Shenzhen Code [13]): MEXICO. In aquosis prope Jalapam, Chamisso 505 (LECB0000548!, image available at https://plants.jstor.org/stable/viewer/10.5555/al.ap.specimen.lecb0000548, accessed on 24 November 2024).
Note. Duke [9] (p. 226) stated “Drymaria villosa Cham. & Schlecht. in Linnaea 5:232. 1830. (HOLOTYPE: Schiede & Deppe 505; in aquosis prope Jalapam, B, probably destroyed; isotype at LE!”. Collections at B were mostly destroyed during World War II, as highlighted by many authors (e.g., [33,34,35,36,37]). According to Art. 9.10 of the Shenzhen Code [13], the term “isotype” used by Duke [9] (p. 226) is to be corrected to lectotype.

3.4. A Diagnostic Key of the Members of Drymaria Series Cordatae (Sensu Duke 1961)

1a. Petals equal the sepals; seeds numerous, less than 1 mm, broad………………………………………..2
1b. Petals shorter than sepals; seeds one to many, 0.5–2.0 mm broad………………………………………4
2a. Leaves ovate–reinform; stipules longer than the petiole; petals divided by about half
the length; lobes 2–4, nerved………………………………………………………D. glandulosa var. galeottiva
2b. Leaves deltoid; long petiole; stipules shorter than the petiole; petals divided by more
than half its length; 1 lobe, nerved………………………………………………………………………………3
3a. Stipules lacerate; seeds coarsely tuberculate; tubercles prominent………….D. gracilis subsp. gracilis
3b. Stipules entire; seed merely verucate with remote protuberance……………………….subsp. carinata
4a. Sepals slightly longer than petals; seeds 0.5–0.8 mm; stipules lacerate…………………………………5
4b. Sepals twice the length of petals; seeds 0.7–2.0 mm; stipules entire…………………………………….6
5a. Flowers in terminal cymes; leaves petiolate; petals bifid;
stamens 3–5…………………………………………………………………………D. glandulosa var glandulosa
5b. Flowers clustered in the axil of the leaves; petals bifid or ligulate;
stamens 2–3……………………………………………………………………………………………D. xerophylla
6a. Leaves deltoid–ovate; stipules entire; seeds 3–5, styles united for half the length…………..D. ladewii
6b. Leaves reniform, stipules lacerate, seeds 1–12, style free…………………………………………………7
7a. Flowers campanulate; pedicels glandular; seeds 1–12, broad……………………………………………8
7b. Flowers pyriform; pedicles eglandular; seeds 1–2, broad……………………………………..D. diandra
8a. Seeds have small mamillated spots with a star-like radiating pattern; pollen spherical,
depressed with non-prominent pores……………………………………………………………………D.anilii
8b. Seeds have large spinulate spots that cross each other; pollen oblong with prominent
pores………………………………………………………………………………………………………D. cordata

3.5. Conclusions

Drymaria, a genus belonging to the large tribe Polycarpaeae of the Caryophyllacecae family, is poorly known from a taxonomical point of view. In fact, a recent comprehensive study is lacking, and the last one was published about 70 years ago by Duke [9]. Also, molecular data are few, and those available are included in a wider paper on the whole family [2]. Among the important gaps in morphological and molecular studies of Drymaria, there are, e.g., the Andean species [12], but also Asian species (three—D. cordata, D. diandra, and D.villosa—according to the Plants of the World Online database [8]) which were not investigated in detail. So, our study, which deals with these three species, represents the first one for the continent. Furthermore, the combined use of both morphometric and molecular techniques appears to have never been considered before us for Drymaria and proved to be very useful for analyzing taxonomically critical plant taxa, as highlighted by other authors regarding Caryophyllales groups, e.g., Allmania R.Br. ex Wight [38], Dianthus L. [39], Limonium Mill. [40], or Salicornia L. [41].

Author Contributions

Conceptualization, D.I. and S.A.; methodology, D.I. and S.A.; investigation, D.I. and S.A.; resources, D.I. and S.A.; data curation, D.I. and S.A.; writing—original draft, D.I.; writing—reviewing of drafts and editing, S.A., H.S. and K.W.V. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Data Availability Statement

Data are contained within the article.

Acknowledgments

Thanks are due to the Directors and Curators of the Herbaria quoted in this paper for their support during visits and the loaning of specimens/photographs. Special thanks to A. Giuliani (Istituto Superiore di Sanità, Rome, Italy) for helping with the statistical analyses.

Conflicts of Interest

The authors declare no conflicts of interest.

References

  1. Harbaugh, D.T.; Nepokroeff, M.; Rabeler, K.; McNeill, J.; Zimmer, E.A.; Wagner, W.L. A new lineage–based tribal classification of the family Caryophyllaceae. Int. J. Plant Sci. 2010, 171, 185–198. [Google Scholar] [CrossRef]
  2. Greenberg, A.K.; Donoghue, M.J. Molecular systematics and character evolution in Caryophyllaceae. Taxon 2011, 60, 1637–1652. [Google Scholar] [CrossRef]
  3. Hernández-Ledesma, P.; Berendsohn, W.G.; Borsch, T.; von Mering, S.; Akhani, H.; Arias, S.; Castaneda-Noa, I.; Eggli, U.; Eriksson, R.; Flores-Olvera, H.; et al. A taxonomic backbone for the global synthesis of species diversity in the angiosperm order Caryophyllales. Willdenowia 2015, 45, 281–383. [Google Scholar] [CrossRef]
  4. Bittrich, V. Caryophyllaceae Juss. In The Families and Genera of Vascular Plants, Magnoliid, Hamamelid and Caryophyllid Families; Kubitzki, K., Rohwer, J., Bittrich, V., Eds.; Springer: Berlin, Germany, 1993; Volume 2, pp. 206–236. [Google Scholar]
  5. Pax, F.; Hoffmann, K. Caryophyllaceae Juss. In Die Natürlichen Pflanzenfamilien, 2nd ed.; Engler, A., Harms, H., Eds.; Wilhelm Engelmann: Leipzig, Germany, 1934; Volume 16, pp. 275–364. [Google Scholar]
  6. The Caryophyllales Network 2015+: A Global Synthesis of Species Diversity in the Angiosperm Order Caryophyllales. Available online: https://caryophyllales.org/ (accessed on 19 October 2024).
  7. Dillenberger, M.S.; Kadereit, J.W. Maximum polyphyly: Multiple origins and delimitation with plesiomorphic characters require a new circumscription of Minuartia (Caryophyllaceae). Taxon 2014, 63, 64–88. [Google Scholar] [CrossRef]
  8. POWO [Continuously Updated]. Amaranthus L. Plant of the World Online. Available online: https://powo.science.kew.org/ (accessed on 19 October 2024).
  9. Duke, J.A. Preliminary revision of the genus Drymaria. Ann. Mo. Bot. Gard. 1961, 48, 173–268. [Google Scholar] [CrossRef]
  10. Hartman, R.L. Drymaria. In Flora of North America: North of Mexico; Flora of North America Editorial Committee, Ed.; Oxford University Press: Oxford, UK, 2005; Volume 12, pp. 9–14. [Google Scholar]
  11. Villarreal, J.A.; Estrada, A.E. A new species of Drymaria (Caryophyllaceae) from northeastern Mexico. Brittonia 2008, 60, 329–331. [Google Scholar] [CrossRef]
  12. Montesinos-Tubée, D.B.; Tovar, C.; Iberico-Vela, G.; Montoya-Quino, J.; Sanchez-Vega, I. Drymaria veliziae (Caryophyllaceae), a new species from the Andes of Cajamarca (North Peru). PhytoKeys 2020, 140, 47–56. [Google Scholar] [CrossRef]
  13. Turland, N.J.; Wiersema, J.H.; Barrie, F.R.; Greuter, W.; Hawksworth, D.L.; Herendeen, P.S.; Knapp, S.; Kusber, W.-H.; Li, D.-Z.; Marhold, K.; et al. International Code of Nomenclature for Algae, Fungi, and Plants (Shenzhen Code). In Regnum Vegetabile, Proceedings of the Nineteenth International Botanical Congress, Shenzhen, China, 19 July 2017; Koeltz Botanical Books: Glashütten, Germany, 2018; Volume 159, pp. 1–254. [Google Scholar] [CrossRef]
  14. Balkrishna, A.; Joshi, B.; Srivastava, A.; Shukla, B.K. New plant records to the flora of Haryana. Indian J. For. 2018, 41, 117–127. [Google Scholar] [CrossRef]
  15. Sarmah, K.K.; Borthakur, S.K. A checklist of angiospermic plants of Manas national park in Assam, India. Pleione 2009, 3, 190–200. [Google Scholar]
  16. Barooah, C.; Ahmed, I. Plant Diversity of Assam: A Checklist of Angiosperms and Gymnosperms; Assam Science Technology and Environment Council: Assam, India, 2014. [Google Scholar]
  17. Choudhary, R.K.; Srivastava, R.C.; Das, A.K.; Lee, J. Floristic diversity assessment and vegetation analysis of Upper Siang district of eastern Himalaya in North East India. Korean J. Plant Taxon. 2012, 42, 222–246. [Google Scholar] [CrossRef]
  18. Majumdar, N.C. Caryophyllaceae Juss. In Flora of India; Sharma, B.D., Balakrishnan, N.P., Eds.; BSI: Calcutta, India, 1993; pp. 502–595. [Google Scholar]
  19. Nayar, T.S.; Beegam, A.R.; Sibi, M. Flowering Plants of Western Ghats; Jawaharlal Nehru Tropical Botanical Garden and Research Institute: Kerala, India, 2014; Volume 1. [Google Scholar]
  20. Press, J.R.; Shrestha, K.K.; Sutton, D.A. Annotated Checklist of the Flowering Plants of Nepal Natural History Museum Publications. Available online: http://www.efloras.org/florataxon.aspx?flora_id=110 (accessed on 19 September 2024).
  21. Arya, S.; Kumar, V.N.S.A.; Nathanpillai, A.V.; Philip, A.A.; Sojan, J.; Suresh, V. Polycarpaea palakkadensis (Caryophyllaceae), a new species from Kerala, south-west India. Phytotaxa 2021, 57, 11–30. [Google Scholar] [CrossRef]
  22. Arya, S.; Kumar, V.N.S.A.; Pillai, A.V.; Alen, A.P.; Sojan, J.; Suresh, V. Three new species of Polycarpaea (Caryophyllaceae) from Kerala, South India. PhytoKeys 2022, 213, 95. [Google Scholar] [CrossRef]
  23. Thiers, B. [Continuously Updated] Index Herbariorum: A Global Directory of Public Herbaria and Associated Staff; New York Botanical Garden’s Virtual Herbarium: New York, NY, USA, 2016; Available online: http://sweetgum.nybg.org/science/ih/ (accessed on 19 September 2024).
  24. Doyle, J.J.; Doyle, J.L. Isolation of plant DNA from fresh tissue. Focus 1990, 12, 13–15. [Google Scholar]
  25. Taberlet, P.; Gielly, G.L.; Pautou, G.; Bouvet, J. Universal primers for amplification of three noncoding regions of chloroplast DNA. Plant Mol. Biol. 1991, 17, 1105–1109. [Google Scholar] [CrossRef] [PubMed]
  26. McGinnis, S.; Madden, T.L. BLAST: At the core of a powerful and diverse set of sequence analysis tools. Nucleic Acids Res. 2004, 32 (Suppl. S2), W20–W25. [Google Scholar] [CrossRef] [PubMed]
  27. Saitou, N.; Nei, M. The neighbor-joining method: A new method for reconstructing phylogenetic trees. Mol. Biol. Evol. 1987, 4, 406–425. [Google Scholar]
  28. Robinson, O.; Dylus, D.; Dessimoz, C. Phylo.io: Interactive viewing and comparison of large phylogenetic trees on the web. Mol. Biol. Evol. 2016, 33, 2163–2166. [Google Scholar] [CrossRef]
  29. IUCN. Guidelines for Using the IUCN Red List Categories and Criteria, Version 14. Prepared by the Standards and Petitions Subcommittee of the IUCN Species Survival Commission. 2024. Available online: https://cmsdocs.s3.amazonaws.com/RedListGuidelines.pdf (accessed on 19 September 2024).
  30. Cafferty, S.; Jarvis, C.E. Typification of Linnaean Plant Names in Caryophyllaceae. Taxon 2004, 53, 1049–1054. [Google Scholar] [CrossRef]
  31. Hermann, P. Paradisus Batavus; apud Abrahamum Elzevier: Lugduni Batavorum, 1698. [Google Scholar]
  32. Mizushima, M. A revision of Drymaria cordata Willd. J. Jpn. Bot. 1957, 32, 69–81. [Google Scholar]
  33. Eggli, U.; Leuenberger, B.E. Type specimens of Cactaceae names in the Berlin Herbarium (B). Willdenowia 2008, 38, 213–280. [Google Scholar] [CrossRef]
  34. Hassemer, G.; Iamonico, D.; Rønsted, N.; Di Pietro, R. Typification of the Linnaean names Plantago serraria and P. subulata (Plantago subgenus Coronopus, Plantaginaceae). Taxon 2017, 66, 738–741. [Google Scholar] [CrossRef]
  35. Iamonico, D.; Hassemer, G.; Rønsted, N.; Di Pietro, R. The intricate nomenclatural questions around Plantago holosteum (Plantaginaceae). Phytotaxa 2017, 306, 75–84. [Google Scholar] [CrossRef]
  36. Nobis, M.; Klichowska, E.; Wolanin, M.; Nobis, A.; Nowak, A. Typification of five plant names described based on specimens collected by Józef Warszewicz in Central and South America. PhytoKeys 2022, 192, 45–61. [Google Scholar] [CrossRef] [PubMed]
  37. Montesinos-Tubée, D.B.; Iamonico, D. The genus Paronychia (Caryophyllaceae) in South America: Nomenclatural review and taxonomic notes with the description of a new species from North Peru. Plants 2023, 12, 1064. [Google Scholar] [CrossRef] [PubMed]
  38. Sindhu, A.; Iamonico, D.; Suresh, V.; Venugopalan Nair Saradamma, A.K. First molecular and morphometric data for the genus Allmania (Amaranthaceae), with the description of a new species from India. Phytotaxa 2022, 559, 221–237. [Google Scholar] [CrossRef]
  39. Franzoni, J.; Astuti, G.; Bartolucci, F.; Bernardo, L.; Fior, S.; Casazza, G.; Conti, F.; Iamonico, D.; Iberite, M.; Minuto, L.; et al. Different species or altitudinal morphotypes? Testing the taxonomic value of Dianthus brachycalyx (Caryophyllaceae). Taxon, 2024; early view. [Google Scholar] [CrossRef]
  40. Iamonico, D.; de Castro, O.; di Iorio, E.; Nicolella, G.; Iberite, M. Taxonomy complexity of some Tyrrhenian endemic Limonium species belonging to L. multiforme group (Plumbaginaceae): New insights from molecular and morphometric analyses. Plants 2022, 11, 3163. [Google Scholar] [CrossRef]
  41. Sciuto, K.; Marion, A.F.; Sfriso, A.; Brancaleoni, L.; Iberite, M.; Iamonico, D. Molecular and morphometric update on Italian Salicornia (Chenopodiaceae), with a focus on the species S. procumbens s.lat. Plants 2023, 12, 375. [Google Scholar] [CrossRef]
Plants 13 03378 g001
Figure 1. Dendrogram (UPGMA method) showing the relationships between the specimens examined in the present study (the vertical axis is dissimilarity). The red line refers to the cutoff dissimilarity point (1.35). Clades: Drymaria sp. (green), D. villosa (blue), and D. diandra (yellow).
Figure 1. Dendrogram (UPGMA method) showing the relationships between the specimens examined in the present study (the vertical axis is dissimilarity). The red line refers to the cutoff dissimilarity point (1.35). Clades: Drymaria sp. (green), D. villosa (blue), and D. diandra (yellow).
Plants 13 03378 g001
Plants 13 03378 g002
Figure 2. Ordination (PCA) based on quantitative morphological characters measured in the present study (component1 vs. component2). Groups: Drymaria sp. (yellow), D. diandra (blue), and D. villosa (green).
Figure 2. Ordination (PCA) based on quantitative morphological characters measured in the present study (component1 vs. component2). Groups: Drymaria sp. (yellow), D. diandra (blue), and D. villosa (green).
Plants 13 03378 g002
Plants 13 03378 g003
Figure 3. DA. The first two discriminant functions explain 100.0% of the total variation [eigenvalues: 79.7% (1st function) and 20.3% (2nd function)]. Dots: Drymaria diandra (red), D. villosa (blue), and Drymaria sp. (green).
Figure 3. DA. The first two discriminant functions explain 100.0% of the total variation [eigenvalues: 79.7% (1st function) and 20.3% (2nd function)]. Dots: Drymaria diandra (red), D. villosa (blue), and Drymaria sp. (green).
Plants 13 03378 g003
Plants 13 03378 g004
Figure 4. Neighbor joining dendrogram. Drymaria sp. is “Dryamria anilii” (proposed binomial for the new species; see paragraph “3.3. Taxonomic treatment”) in the dendrogram.
Figure 4. Neighbor joining dendrogram. Drymaria sp. is “Dryamria anilii” (proposed binomial for the new species; see paragraph “3.3. Taxonomic treatment”) in the dendrogram.
Plants 13 03378 g004
Plants 13 03378 g005
Figure 5. Drymaria anilii (a) habit; (b) detail of inflorescence; (c) hairiness at nodes; (d) stem; (e) closed flower; (f) bracteole; (g) tepal; (h) petal; (i) gynoecium; (j) fruit; (k) seed (photos by Arya Sindhu).
Figure 5. Drymaria anilii (a) habit; (b) detail of inflorescence; (c) hairiness at nodes; (d) stem; (e) closed flower; (f) bracteole; (g) tepal; (h) petal; (i) gynoecium; (j) fruit; (k) seed (photos by Arya Sindhu).
Plants 13 03378 g005
Plants 13 03378 g006
Figure 6. SEM images of Drymaria anilii: (a) seed; (b) magnification of seed surface; (c) pollen grain; (d) magnification of pollen grain.
Figure 6. SEM images of Drymaria anilii: (a) seed; (b) magnification of seed surface; (c) pollen grain; (d) magnification of pollen grain.
Plants 13 03378 g006
Plants 13 03378 g007
Figure 7. Location of Drymaria anilii in NE India.
Figure 7. Location of Drymaria anilii in NE India.
Plants 13 03378 g007
Table 1. Drymaria populations studied in the present research.
Table 1. Drymaria populations studied in the present research.
LocalityTaxon
India, Nelliyampathy, Palakkad, Kerala, on muddy slopes D. diandra
India, Pothundi, Palakkad, Kerala, on plain landD. diandra
India, Cheruthoni, Idukki, Kerala, on slopesD. diandra
India, Kohima, Nagaland, on muddy slopesDrymaria sp.
India, Kohima, Nagaland, on slopesDrymaria sp.
India, Adimali, Idukki, Kerala, on muddy slopesD. villosa
India, Minnampara, Palakkad, Kerala, on muddy slopesD. villosa
Table 2. Characters measured for the morphometric analysis. The characters labeled with an asterisk are qualitative, and the other ones are quantitative.
Table 2. Characters measured for the morphometric analysis. The characters labeled with an asterisk are qualitative, and the other ones are quantitative.
1. Height of plant (cm)
2. Length of leaf blade (mm)
3. Width of leaf blade (mm)
4. Length of petiole (mm)
5. Diameter of petiole (mm)
6. Number of flowers per inflorescence *
7. Length of bracteoles (mm)
8. Width of bracteoles (mm)
9. Length of bracts (mm)
10. Width of bracts (mm)
11. Length of sepals (mm)
12. Width of sepals (mm)
13. Length of petals (mm)
14. Width of petals (mm)
15. Length of stamens (mm)
16. Length of gynoecium (mm)
17. Width of gynoecium (mm)
18. Length of fruit (mm)
19. Width of fruit (mm)
20. Minor diameter of seed (mm)
21. Major diameter of seed (mm)
Table 3. GenBank accession numbers and taxa selected for molecular analysis.
Table 3. GenBank accession numbers and taxa selected for molecular analysis.
Sl No.SpeciesGenBank Accession Number
1Spergularia mediaEU812820
2Spergularia rubraOQ150721
3Spergularia marinaMG234808
4Drymaria diandraPP0712821
5Loeflingia hispanicaKX282251
6Scopulophia rixfordiiMF963915
7Drymaria aniliiPP068051
8Drymaria cordataKF737457
9Drymaria mollugineaJN589128
10Drymaria laxifloraAY286528
11Achyronychia cooperiMF964174
12Pteranthus dichotomusAY936250
13Sphaerocoma aucheriAJ310979
14Sphaerocoma hookeriHE586023
15Pycnophyllum bryoidesJN589108
16Dicheranthus plocamoidesKF737513
17Polycarpon tetraphyllumOQ412391
18Cardionema ramosissimumMG235912
19Illecebrum verticillatumKX167481
20Scleranthus annuusOQ150654
21Sagina procumbensOR593647
22Arenaria serpyllifoliaMG101851
23Stellaria holosteaKX183996
24Dianthus caryophyllusMT923226
Table 4. MANOVA applied to populations as groups.
Table 4. MANOVA applied to populations as groups.
Statistic TestTest ValueF-Ratiop (0.05)
Wilks’ Lambda0.000176258.10.00000001
Hotelling–Lawley Trace182.958906809.760.00000001
Pillai’s Trace1.98437727.030.00000001
Roy’s Largest Root146.0745662020.70.00000001
Table 5. MANOVA applied to species as groups.
Table 5. MANOVA applied to species as groups.
Statistic TestTest ValueF-Ratiop (0.05)
Wilks’ Lambda0.0001942008.380.00000001
Hotelling–Lawley Trace175.5896222458.250.00000001
Pillai’s Trace1.9656321639.560.00000001
Roy’s Largest Root139.92164011.090.00000001
Disclaimer/Publisher’s Note: The statements, opinions and data contained in all publications are solely those of the individual author(s) and contributor(s) and not of MDPI and/or the editor(s). MDPI and/or the editor(s) disclaim responsibility for any injury to people or property resulting from any ideas, methods, instructions or products referred to in the content.

Share and Cite

MDPI and ACS Style

Arya, S.; Singh, H.; Vishnu, K.W.; Iamonico, D. Study on the Genus Drymaria (Caryophyllaceae)—A New Species from North-East India. Plants 2024, 13, 3378. https://doi.org/10.3390/plants13233378

AMA Style

Arya S, Singh H, Vishnu KW, Iamonico D. Study on the Genus Drymaria (Caryophyllaceae)—A New Species from North-East India. Plants. 2024; 13(23):3378. https://doi.org/10.3390/plants13233378

Chicago/Turabian Style

Arya, Sindhu, Harsh Singh, Kalarikkal Walsan Vishnu, and Duilio Iamonico. 2024. "Study on the Genus Drymaria (Caryophyllaceae)—A New Species from North-East India" Plants 13, no. 23: 3378. https://doi.org/10.3390/plants13233378

APA Style

Arya, S., Singh, H., Vishnu, K. W., & Iamonico, D. (2024). Study on the Genus Drymaria (Caryophyllaceae)—A New Species from North-East India. Plants, 13(23), 3378. https://doi.org/10.3390/plants13233378

Note that from the first issue of 2016, this journal uses article numbers instead of page numbers. See further details here.

Article Metrics

Back to TopTop