Bull 65 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Shamso et al. Bull. Iraq nat. Hist. Mus. (2024) 18 (1): 65-90. https://doi.org/10.26842/binhm.7.2024.18.1.0065 ORIGINAL ARTICLE PALYNOLOGICAL AND FOLIAR EPIDERMAL STUDIES OF GENUS OXALIS LINNAEUS, 1753 (OXALIDALES, OXALIDACEAE) FROM EGYPT Eman M. Shamso*, Hasnaa A. Hosni *, Amal A. Draz**, Salwa A. Kawashty** and Sameh R. Hussein** *The Herbarium, Botany & Microbiology Department, Faculty of Science, Cairo University, Giza 12613 Egypt. **Phytochemistry & Plant Systematics Department, National Research Centre. 33 El Bohouth St., Dokki, Giza, P.O. 12622, Egypt. Corresponding author: eshamso@sci.cu.edu.eg Received: 28 September 2023, Revised:26 Nov. Accepted: 3 December 2023, Published:20 June 2024 This work is licensed under a Creative Commons Attribution 4.0 International License ABSTRACT By using light microscopy (LM) and scanning electron microscopy (SEM), the pollen grains and foliar epidermal morphology of four species and two varieties of Egyptian Oxalis Linnaeus, 1753 belonging to three sections of the subgenus Oxalis Linnaeus, 1753 were studied. The objective was to determine the importance of pollen and foliar epidermal characters as taxonomic evidence. The current study revealed that the pollen grains of all taxa examined are monads, radially symmetric, isopolar, and of medium size; the shape varied from sub-spheroidal to subprolate. The pollen apertures were tricolpate with ornamented colpal membranes (rugulate, warty, congregated granules). Exine is semi-tectate, with micro- reticulate to reticulate ornamentation, perforate in O. debilis Kunth, 1822. The study also revealed the presence of four types of stomata: actinocytic, anisocytic, anomocytic, and an unusual 4-celled anisocytic. Both abaxial and adaxial epidermal cells were irregular or polygonal in shape, with undulate or straight anticlinal walls and semi-swollen to swollen periclinal walls with dispersed epicuticular wax. Two forms of non-glandular, unicellular trichomes were observed: short clavate and long with a tapering apex and papillate surface. The study further discussed numerical results based on combined morphological, palynological, and foliar epidermal characters. This study is the first report dealing with anatomical and palynological features of the genus Oxalis in Egypt. Keywords: Egypt, Foliar epidermis, Numerical analysis, Oxalis, Pollen morphology. INTRODUCTION Oxalis Linnaeus, 1753 is a cosmopolitan genus and presents about 566 accepted species (POWO, 2023), distributed within four subgenera: Monoxalis (Small) Lourt., Trifidus Lourt., Thamnoxys (Endl.) Reiche and Oxalis. It is believed that South America and Southern Africa BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Iraq Natural History Research Center & Museum, University of Baghdad https://jnhm.uobaghdad.edu.iq/index.php/BINHM/Home Copyright © Bulletin of the Iraq Natural History Museum Online ISSN: 2311-9799, Print ISSN: 1017-8678 https://doi.org/10.26842/binhm.7.2024.18.1.0065 https://orcid.org/0000-0002-0068-0224 https://orcid.org/0000-0001-7609-9383 https://orcid.org/0000-0002-1979-511X https://orcid.org/0000-0003-3914-3423 https://orcid.org/0000-0003-3657-7410 mailto:eshamso@sci.cu.edu.eg https://creativecommons.org/licenses/by/4.0/ https://jnhm.uobaghdad.edu.iq/index.php/BINHM/Home 66 Bull. Iraq nat. Hist. Mus. 18 (1): 65-90. Palynological and foliar epidermal studies are the centers of distribution, and South America is probably the place of origin of this genus (Knuth, 1930; Denton, 1973; Lourteig, 1994, 2000; De Azkue, 2000). The genus presents many difficulties in identification, mainly because of the use of morphological characteristics that are not very clear (Abreu et al., 2012). Pollen morphology has a taxonomic importance and can support the morphological and the phylogenetic studies (Chen and Xia, 2011; Tripathi et al., 2017). The taxonomic significance of pollen morphology for the genus Oxalis was a topic for many studies. These palynological studies were performed on Oxalis from various aspects such as morphology (Perveen and Qaiser, 2003; Rosenfeldt and Galati, 2007), type of pollen grains (Ghosh and Verma, 1985; Dreyer, 1996), formation of aberrant pollen (Dreyer and Van Wyk, 1998), and orbicule morphology (Rosenfeldt and Galati, 2005, 2008; Lòpez and Rosenfeldt, 2015, 2016). A comprehensive study on the pollen morphology of American and African members of the genus Oxalis has been performed by Huynh (1969 a, b), who identified four pollen types and 13 pollen subtypes based on the presence, or absence of a distinct supra-reticulum and the simple or complex constitution of apertures (colpate, porate, or colporate); in addition to the differences in grain size that were observed in anthers from the different stamen whorls of Oxalis flowers. Dreyer (1996) examined the pollen micro-morphology of all the Southern African members of Oxalis (270 taxa) using LM, SEM, and TEM techniques. Four main pollen types and 19 subtypes were described based on tectum structure and the number and arrangements of apertures. The study also recorded the presence of variable percentages of aberrant pollen grains in some taxa of Oxalis. The occurrence of aberrant pollen in this genus has been reported previously by Dreyer and Van Wyk (1998). They studied the pollen from both stylar whorls to detect the relation between palynological variation and tristyly. They showed that the aberrations always affect the number and arrangement of the apertures, but never alter the exine structure of the grain and also revealed that reticulate pollen displays a low percentage of aberrations. The macro- and micro-morphology of the epidermis and its appendages have been used by various authors to classify certain taxa of the angiosperms (Adedeji et al., 2007; Chukwuma et al., 2017; Bahadur et al., 2022; Chukwuma et al., 2022). The most important epidermal traits are trichome types, shape of epidermal cells, stomata types, stomata position, and subsidiary cells. These traits are useful for identification purposes and for the determination of phylogenetic relationships between taxa (Metcalf and Chalk, 1950, 1979; Jones, 1986; Judd et al., 2008; Ogundipe et al., 2009; Araujo et al., 2010; Zhang et al., 2018; Bahadur et al., 2023). Oxalis is tremendously variable in the number and shape of leaflets, leaf and leaflet size, degree of leaflet conduplication, petiole length and shape, nature of the epidermis, and indumentum attributes (Salter, 1944). However, only a few studies have focused on the leaf anatomy of this genus. Metcalfe and Chalk (1950); dos Reis and Alvim (2013); Jooste (2015); and Richetti et al. (2022) are among the authors who have investigated and described the leaf 67 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Shamso et al. anatomical features of the genus Oxalis. Singh (2010) showed that anatomical features have played an important role in the clarification of phylogenetic relationships among South American Oxalis species. Jooste et al. (2016) assessed the variation of fifty-nine leaflet anatomical traits of 109 Southern African species in a phylogenetic context and stated that a combination of six leaflet anatomical traits (stomatal position, adaxial epidermal cell types, abaxial epidermal cell types, mesophyll type, presence or absence of a sheath around vascular tissue, and degree of leaflet conduplication) is useful and supports various Southern African taxa clades previously defined by DNA-based phylogenetic work (Oberlander et al., 2011). In Egypt, this genus is represented by four species and two varieties belonging to three sections in the subgenus Oxalis (Draz et al., 2021; Shamso et al., 2021). The current study presents a description of the macro- and micromorphology of pollen grains and foliar epidermal cells using both LM and SEM for the evaluation of their taxonomic significance and uses numerical taxonomy to better understand the phenetic relationships among the studied taxa within the genus. MATERIALS AND METHODS Plant materials: The present study is based on field studies and herbarium specimens held at the following herbaria: Cairo University (CAI), National Research Centre (CAIRC); Tanta University (TANE), and the Agriculture Museum, Flora and Phytotaxonomy Researches (CAIM) [Herbaria acronyms according to Thiers (2023); continuously updated] (Tab. 1). The nomenclature of all taxa has been updated according to several websites: POWO (2023) and IPNI (2023). The morphological data were obtained from both fresh and herbarium specimens. For each taxon, at least 3-4 specimens were chosen (if available) to encompass the range of variations. Light investigation: The following steps were carried to examine the pollen and lamina samples: 1. Pollen examination: Pollen samples were taken from mature anthers of flowering buds. For light microscopy (LM), the pollen grains were prepared using the technique of Erdtman (1960) and then mounted in glycerin jelly. Light microscopic observations were made using a Serico research microscope under (E 40, 0.65) with a 16-x eye piece. The measurements of pollen grains were based on at least 15-20 pollen grains for each specimen. Various pollen characters: shape, size, aperture type, and exine sculpture were determined (Tab. 2). Pollen terminology is followed Punt et al. (2007); Hesse et al. (2009) and Halbritter et al. (2018). 2. Epidermal examination: Three pieces of lamina were embedded in KOH 5% for 24-48 hrs. Transverse sections of all studied taxa were taken in lamina through hand cutting, and stained with a Safranin-fast green standard double stain to show the shape of epidermal cells (Sass, 1961). In an additional method, the epidermal peels were obtained from fresh leaves after being washed in water, painted with colorless nail polish on abaxial and adaxial surfaces; each epidermal peel was fixed and examined under a light microscope. The micromorphological characters of both surfaces were studied at different 68 Bull. Iraq nat. Hist. Mus. 18 (1): 65-90. Palynological and foliar epidermal studies magnifications (10X, 40X objectives) and photographed using an Olympus - cx41 light microscope equipped with an Olympus digital camera. The qualitative characteristics include the shape of epidermal cells, stomata, and trichomes, their presence and absence, and their types (Tab. 3). Terminology was followed by Metcalfe and Chalk (1950) and Barthlott (1981). Voucher specimens and slides were kept in the Cairo University Herbarium (CAI). SEM Microscopy investigation: For each taxon, non-acetolyzed pollen grains and small pieces of the median portion of dry lamina were mounted onto stubs with double sided adhesive tape, and then these stubs were sputter-coated with gold. After coating, they were examined using a field emission scanning electron microscope (FE-SEM) QUANTA FEG250Y at an accelerating voltage of 20 KV, at the Electron Microscopy Unit, National Research Centre, Dokki. Data Analysis: Forty-eight morphological, palynological, and foliar epidermal traits, divided into 113 character states, were analyzed and scored. The data matrix was constructed of six OTUs (App. 1). In this study, the similarity will be measured at the species level (represented by specimens). An equal number of specimens of each species (three specimens, if available) were used. The resemblance between the fundamental taxonomic units was determined in two steps: first, measure the similarity values (or distance values) between all possible pairs of specimens under study of the studied characters and character states; and second, form the similarity matrix. This matrix was analyzed using the numerical taxonomy technique supplied in the Minitab program, version 20 (Minitab, 2020). Table (1): The collection data of the taxa included in the present study: Taxon Locality and date of collection Collector and Number Herbarium and specimen code Section: Cernuae Knuth, 1914 O. anthelmintica A. Rich., 1847 O. pes-caprae L., 1753 -Mountain tributary, Wadi Akwametra, Gebel Elba, 27-2-1967. -Wadi Darawein, Gebel Elba, 3-2-1979. -Bramly’s grotto, Burg El-Arab, 28-2-2019 -Behig, 4-5-1976 -Maruit, 23-3-2006 Osborn & Helmy s.n. Boulos 12924 A. Draz s.n. Ahmed & Mokhtar 71 D.A.Ahmed 2937 (CAI), cai.46.240.722.18 (CAIRC), A306 (CAIRC), A304 (CAI), cai.46.240.721.33 (TANE), Not coded 69 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Shamso et al. Section: Corniculatae DC., 1824 O. corniculata var. corniculata O. corniculata var. repens (Thunb.) Zucc., 1831 - El Bagor, Monufia,12- 4-2019 - Bir Romani, Budkhulu, Dakhla, 12-2-1952; - Wadi Feiran, 19-10- 2012 - National research center garden, Giza, 13-5-2019 - ibid, 18-4-2021 A. Draz s.n. Täckholm & Kassas 185 Keshta s.n. A. Draz s.n A. Draz s.n (CAIRC), A301 (CAI), cai.46.240.723.1.10 (TANE), Not coded (CAI), cai.240.723.2.1 (CAIRC), A302 Section: Ionoxalis Small, 1903 O. debilis Kunth, 1822 (= O. corymbosa DC) O. latifolia Kunth,1822 -Giza, 12-4-2020 -Dokki, 20-2-2021 -Arab El-Ghadir, Qalubia, French bean, 20-5-2003 -Giza, Cairo University Garden, 15-4-2021 A. Draz s.n. H. Hosni s.n. El Khanagry s.n. E. Shamso s.n. (CAIRC), A303 (CAI), cai.46.240.723.5.11 (CAIM), Not coded (CAI), cai.46.240.723.8.4 RESULTS AND DISCUSSION The current study of pollen morphology and the epidermal features of leaflet lamina using LM & SEM have revealed some valid characters for identification and classification of the studied Oxalis taxa (Tab. 2, 3). I-Pollen grains morphology: Palynological characters of six taxa belonging to the genus Oxalis were studied under LM and SEM for detecting various characteristics such as polarity, size, shape, aperture type, aperture arrangements, aperture membrane sculpture, and exine sculpture of pollen grains. Detailed pollen morphological features of the investigated taxa are summarized in Table (2), and representative pollen grains are illustrated in Plates (1-3). The morphological characters showed that the pollen grains of all studied taxa were monads, isopolar, radially symmetrical, and circular in polar view. Pollen size medium (medium to large in O. pes-caprae) with mean polar axis ranges from 26.9µm ± 1.1 in O. corniculata var. corniculata to 38.6 µm ±0.94 in O. anthelmintica; mean equatorial diameter ranges from 22.6 µm ±1.2 in O. debilis to 33.8 µm ±1.2 in O. corniculata var. repens. The shape of pollen grains varied from sub-spheroidal to sub-prolate in an equatorial view. The sub-spheroidal grains ranged from prolate-spheroidal in O. corniculata var. corniculata (P/E= 1.08) to oblate-spheroidal in O. corniculata var. repens (P/E= 0.97). The results of this study agree with Dreyer (1996) that the shape of pollen is of minor importance among the southern 70 Bull. Iraq nat. Hist. Mus. 18 (1): 65-90. Palynological and foliar epidermal studies African members of Oxalis, and most grains ranged from spherical to oblate (seldomly prolate) in an equatorial view. The present study showed that the pollen grains are normally tricolpate, and tetracolpate grains are also observed within the same species, as in O. debilis, O. corniculata var. repens, and O. latifolia (Pls. 2, 3). These findings are in agreement with Ghosh and Verma (1985); Dreyer and Van Wyk (1998). Moore (1973) and Goldblatt (1987) suggested that the presence of two types of grains (tri- and tetracolpate) within the same species is the result of polyploidy. Colpi apocolpate, vary from elliptic to fusiform or oblong; the colpal membrane was densely granulated to rugulated in O. corniculata (Pl. 1 C), scabrous to warty in O. corniculata var. repens, with congregated granules distributed irregularly in O. pes-caprae (Pl. 1 G), and warty in O. anthelmintica, O. latifolia, and O. debilis (Pl. 1 J, M). The exine of the studied taxa was semi-tectate, brochate, uniform, and generally micro- reticulate to reticulate ornamentation, and rarely perforate in O. debilis (Pl. 1 O). Brochi are circular to polygonal, diminishing in size towards the colpi. Lumina with free single columella in O. pes-caprae (Pl. 1 I). Huynh (1969 a, b) and Dreyer (1996) indicated that the reticulate pollen type is the most common type among the South African members of Oxalis. Our results coincide with those of previous studies, and the exine features have little taxonomic value for identification and species delimitation. Key to the examined taxa of Oxalis based on pollen morphology: 1. Pollen prolate-spheroidal or oblate-spheroidal (Pl. 1 A, D)… …..……….…………….… 2 - Pollen sub-prolate (Pl. 1 G, J, M)……………………………………….…... ………..…...3 2. Pollen prolate-spheroidal, with P/E= 1.08 µm, tri-colpate, colpus 15-20 µm long, granulate to rugulate membrane (Pl. 1 A-C and Pl. 2 A) …………... O. corniculata var. corniculata - Pollen oblate-spheroidal, with P/E=0.97µm, tri- or tetra-colpate, colpus 20-25 µm long, scabrous to warty membrane (Pl. 1 D-F and Pl. 3 B) ……....… O. corniculata var. repens 3. Pollen with tricolpate (Pl. 2 B, C).………………………..…………………………...….. 4 - Pollen with tricolpate and tetracolpate (Pl. 2 A) ……...........................………………..…. 5 4. Exine sculpture micro-reticulate, colpal membrane warty (Pl. 1 K, L) .…O. anthelminthica - Exine sculpture reticulate with free single columella in lumina, colpal membrane with congregated granules (Pl. 1 G, I) ……..……………………...………...….….O. pes-caprae 5. Pollen grains mean polar axis 29.1 µm, mean equatorial diameter 22.57 µm. Exine sculpture perforate (Pl. 1 O) ...………………………………….………………. O. debilis - Pollen grains mean polar axis 34.75 µm, mean equatorial diameter 28.75 µm. Exine sculpture reticulate (Pl. 3 B) ………………………………………….……..… O. latifolia 71 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Shamso et al. E x in e sc u lp tu re M ic ro - re ti cu la te R et ic u la te lu m in a w it h fr ee s in g le co lu m el la M ic ro - R et ic u la te to r et ic u la te R et ic u la te P er fo ra te to m ic ro re ti - cu la te R et ic u la te C o lp u s m em b ra n e w ar ty W it h co n g re g at ed g ra n u le s d is tr ib u te d ir re g u la rl y D en se ly g ra n u le s t o r u g u la te S ca b ro u s t o w ar ty W ar ty W ar ty S iz e ca te g o ry : S m al l: 1 0 -2 5 μ m , M ed iu m s iz ed : 2 6 -5 0 μ m , la rg e 5 1 -1 0 0 μ m , an d s h ap e cl as se s w er e ca te g o ri ze d b as ed o n t h e ra ti o o f th e m ea n p o la r ax is ( P ) an d m ea n e q u at o ri al d ia m et er ( E ) (E rd tm an , 1 9 5 2 ). C o lp u s le n g th 2 0 - 3 0 2 0 - 4 0 1 5 - 2 0 2 0 - 2 5 2 0 - 2 5 2 0 - 3 0 . C o lp u s fu si o n A p o co - lp at e A p o co - lp at e A p o co - lp at e A p o co - lp at e A p o co - lp at e A p o co - lp at e T a b le ( 2 ): P o ll en m o rp h o lo g ic al c h ar ac te rs o f in v es ti g at ed t ax a. C o lp u s sh ap e E ll ip ti c F u si fo rm E ll ip ti c O b lo n g F u si fo rm F u si fo rm A p er tu re - ty p e T ri co lp at e T ri co lp at e T ri co lp at e T ri - an d te tr ac o - lp at e T ri - an d T et ra co - lp at e T ri - an d T et ra co - lp at e P o ll en s iz e M ed iu m si ze d M ed iu m t o la rg e si ze d M ed iu m si ze d M ed iu m si ze d M ed iu m si ze d M ed iu m si ze d P o ll en sh ap e S u b -p ro - la te S u b -p ro - la te P ro la te - sp h er o id al O b la te - sp h er o id al S u b -p ro - la te S u b - p ro la te P /E R at io (μ m ) 1 .2 1 .2 1 .1 1 1 .3 1 .2 E q u at o ri al d ia m et er ( E ) M ea n 3 1 .4 ± ( 1 .3 ) 3 0 .8 ± ( 2 .9 ) 2 4 .7 ± ( 1 .4 ) 3 3 .8 ± ( 1 .2 ) 2 2 .6 ± ( 1 .2 ) 2 8 .8 ± ( 1 .1 ) R an g e (μ m ) 2 5 - 4 5 1 5 - 6 0 2 0 - 3 5 2 5 - 4 0 1 5 - 3 5 2 6 - 4 5 P o la r ax is ( P ) M ea n 3 8 .6 ± ( 0 .9 4 ) 3 7 ± ( 1 .5 ) 2 6 .9 ± ( 1 .1 ) 3 3 .1 ± ( 0 .7 ) 2 9 .1 ± ( 1 .5 ) 3 4 .8 ± ( 1 .2 ) R an g e (μ m ) 3 5 - 4 5 2 5 - 6 0 2 5 - 3 5 3 0 - 3 5 2 0 - 3 5 3 0 - 3 9 T a x o n O . a n th el m in ti ca O . p es -c a p ra e O . co rn ic u la ta v a r. c o rn ic u la ta O . co rn ic u la ta v a r. r ep en s O . d eb il is O . la ti fo li a 72 Bull. Iraq nat. Hist. Mus. 18 (1): 65-90. Palynological and foliar epidermal studies T r ic h o m e ty p e C la v at e u n ic el lu la r L o n g u n ic el lu la r L o n g u n ic el lu la r L o n g u n ic el lu la r C la v at e an d l o n g u n ic el lu la r so m et im es m u lt ic el lu - la r u n is er ia te C la v at e an d lo n g u n i- ce ll u la r S to m a ta l sh a p e O b lo n g E ll ip so id O b lo n g S il t- li k e E ll ip so id E ll ip so id S to m a ta l p a tt e r n A n is o cy ti c an d u n u su al 4 -c el le d an is o cy ti c A n is o cy ti c an d u n u su al 4 -c el le d an is o cy ti c A n is o cy ti c, u n u su al 4 -c el le d an is o cy ti c an d so m et im es an o m o cy ti c A n is o cy ti c an d so m et im es ac ti n o cy ti c A n is o cy ti c, u n u su al 4 -c el le d an is o cy ti c an d so m et im es an o m o cy ti c A n is o cy ti c, u n u su al 4 -c el le d an is o cy ti c an d so m et im es an o m o cy ti c T a b le ( 3 ): F o li ar e p id er m al c h ar ac te ri st ic s o f in v es ti g at ed t ax a. ta x a. S to m a ta l p o si ti o n H y p o st o - m at ic H y p o st o - m at ic A m p h is to - m at ic A m p h is to - m at ic A m p h is to - m at ic H y p o st o - m at ic E p ic u ti c - u la r w a x ab se n t P re se n t P re se n t P re se n t P re se n t P re se n t A d a x ia l E p id er m a l c el ls P e ri c li n a l w a ll S em i- sw o ll en S w o ll en S li g h tl y S em i- sw o ll en S w o ll en S em i- sw o ll en S w o ll en A n ti c li n a l w a ll p a tt e r n st ra ig h t S tr ai g h t t o s li g h tl y cu rv ed U n d u la te S tr ai g h t U n d u la te S tr ai g h t S h a p e P o ly g o n - n al ± P o ly g o - n al Ir re g u la r ± P o ly g o - n al Ir re g u la r P o ly g o n al A b a x ia l E p id er m a l ce ll s P e r ic li n a l w a ll s u r fa c e g le b u la te g le b u la te S m o o th S m o o th ± S m o o th W it h ce n tr al p ap il la e P e r ic li n a l w a ll S em i- sw o ll en S w o ll en S em i- sw o ll en S w o ll en S em i- sw o ll en S w o ll en A n ti cl in a l w a ll p a tt e r n st ra ig h t S tr ai - g h t to sl ig h tl y cu rv ed U n d u l- at e S tr ai - g h t U n d u l- at e S tr ai - g h t S h a p e P o ly g o - n al ± P o ly g o - n al Ir re g u - la r ± P o ly - g o n al Ir re g - u la r P o ly g - o n al T a x o n O . a n th el m in ti ca O . p es -c a p ra e O . co rn ic u la ta v a r. co rn ic el a ta O . co rn ic u la ta v a r. r ep en s O . d eb il is O . la ti fo li a 73 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Shamso et al. II- Foliar epidermal characteristics: The current study primarily focuses on micro- morphological features of trichomes, epidermal cells, and stomata. The data on the foliar epidermal characters of the taxa examined are presented in Table (3). Trichomes on plants are extremely variable in their presence, density, and form; therefore, their morphology and structure could be of taxonomical importance in certain plant groups (Che Amri et al., 2018; Bahadur et al., 2023). According to previous studies by Metcalf and Chalk (1950); dos Reis and Alvim (2013); and Jooste (2015), Oxalis is recognized for having two types of foliar trichomes: glandular and non-glandular trichomes. In the present study, the majority of the studied taxa were sparsely pubescent, restricted to the midrib, abaxial surface, and margin of the leaf blade. Unicellular trichomes were the main type, with two forms: short, clavate unicellular (Pl. 4 C) and long unicellular with a tapering apex and papillate surface (Pl. 4 A, B); while multicellular uniseriate trichomes were observed only in O. debilis (Pl. 4D). The presence of glandular trichomes is not reported in Egyptian taxa, and trichome characters were rather uniform and of little taxonomic importance. Epidermal anticlinal walls have low interspecific variation in Oxalis and can be regarded as a taxonomically stable character (Jooste, 2015; Jooste et al., 2016). In the present study, both abaxial and adaxial epidermal cells were polygonal or irregular in shape with a straight to undulate anticlinal wall pattern (Pl. 6). The variability in the anticlinal cell wall pattern may be related to the environmental conditions (Stace, 1965; Metcalf and Chalk, 1979). All studied taxa are listed as having semi-swollen to swollen periclinal walls (Pl. 5), and their surfaces were smooth in O. corniculata and O. debilis (Pl. 6C); glebulate in O. anthelminthica and O. pes-caprae (Pl. 6B), and with central papillae in O. latifolia (Pl. 6A). Jooste (2015) reported the occurrence of swollen and papillose epidermal cells in some African Oxalis species, which act as lenses to focus light through the epidermis into the chloroplast-rich palisade cells (Poulson and Vogelmann, 1990; Myers et al., 1994; Vogelmann et al., 1996). Moreover, Gkikas et al. (2015) reported the production of epicuticular wax by those papillose cells that functioned as a water repellent material, a character that was assigned in the present study for O. corniculata, O. pes-caprae, O. debilis, and O. latifolia. The types and distribution of stomata provide taxonomically useful information in some taxa (Metcalf and Chalk, 1950). According to Jooste et al. (2016), stomata display great variations in their types and distribution, and recorded the presence of epistomatic, hypostomatic, and amphistomatic leaflets in southern African Oxalis. In this study, the majority of the studied taxa have hypostomatic leaves, in the case of O. anthelminthica, O. pes-caprae, and O. latifolia. However, in the case of O. corniculata and O. debilis leaves, were amphistomatic, similar to previous studies by dos Reis and Alvim (2013). Anomocytic and paracytic stomatal complex types have been reported in the Oxalidaceae by Metcalf and Chalk (1950); dos Reis and Alvim (2013); while Jooste et al. (2016) reported four types of stomata: anomocytic, anisocytic, actinocytic, and an unusual 4-celled anisocytic stomatal type. These four types of stomata were observed in the studied taxa, where the anisocytic and an unusual 4- celled anisocytic types of stomata were present in all studied taxa 74 Bull. Iraq nat. Hist. Mus. 18 (1): 65-90. Palynological and foliar epidermal studies (Pl. 7). However, the actinocytic type was encountered in O. corniculata var. repens and O. latifolia, while the anomocytic type was restricted to O. corniculata var. corniculata and O. debilis. Baranova (1992) suggested that the multiple stomatal types within a leaf are a common occurrence, which is taxonomically significant. Despite the general consensus that stomatal complex types can be taxonomically informative, we could not detect any clear pattern based on these traits among the studied taxa. The subsequent key was based on the foliar epidermal cell characters, which might be used to distinguish between the studied taxa: 1- Both abaxial and adaxial epidermal cells irregular with undulated anticlinal walls (Pl. 6 B, C) ………………………………………………………………………………..…..… 2 - Both abaxial and adaxial epidermal cells polygonal with straight anticlinal walls (Pl. 6 A) …...…...………………………………………….………………………………..….... 3 2- Stomata oblong; trichomes long unicellular (Pl. 4 A, B and Pl. 8 B) ……………………………………………………….…. O. corniculata var. corniculata - Stomata ellipsoid; trichomes clavate and long unicellular sometimes multicellular uniseriate (Pl. 4 and Pl. 8 A) ….………………..............................………. O. debilis 3- Epidermal cells with smooth periclinal wall, amphistomatic stomata (Pl. 6 C)…………………………………………………………….…O. corniculata var. repens - Epidermal cells with glebulate or with central papillae periclinal wall, hypostomatic stomata (Pl. 6 A, B) …………………………………………………………..…..……. 4 4- Epidermal cells with central papillae periclinal wall (Pl. 6 A) ……………….O. latifolia - Epidermal cells with glebulate periclinal wall (Pl. 6 B) ….…...…………………...… 5 5- Epidermal cells with semi-swollen periclinal wall and epicuticular wax absent. Trichomes clavate (Pl. 4 C, Pl. 5 A) ……………..…………………….…………O. anthelminthica - Epidermal cells with swollen periclinal wall and epicuticular wax present. Trichomes long unicellular (Pl. 4 A and Pl. 5 C) …………………………….….…..…….O. pes-caprae III-Numerical analysis The 113 morphological, palynological, and foliar epidermal character states and their binary codes (0, 1) are given in Appendix (1). The numerical analysis obtained is represented as a dendrogram (Diag. 1). The dendrogram shows that the studied taxa were classified into two main series (I, II), which included three clusters (I, II, and III) at a similarity level of 34.18%. Series (I) included four taxa, namely O. anthelmintica, O. pes-caprae, O. debilis, and O. latifolia. The series is characterized by geophytes, stemless herbs with bulbous-rooted, basal rosette leaves, inflorescence on long, erect scape-like peduncle, and pollen subprolate with fusiform colpate. At a 50.36% similarity level, this series is subdivided into two distinct clusters based on the presence or absence of vertical rhizomes, bulb morphology, and the number of pollen apertures. The first cluster (I) represented O. anthelmintica and O. pes- caprae (Section: Cernuae), which are very close with a 72.91% similarity level. Morphologically, these two species are very similar, except for their leaf morphology, flower color, exine sculpture, and epidermal anticlinal wall. The second cluster (II) included O. 75 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Shamso et al. debilis and O. latifolia (section: Ionoxalis) with a 53.38% similarity level, and are differentiated by the leaf shape, bulb, and foliar eidermal characteristics (Pl. 6 A, C). Series (II) included a single cluster (III), represented by the two taxa; O. corniculata var. corniculata and O. corniculata var. repens, with a 74.02% similarity level. Morphologically, these two taxa are very close, except for their leaf color, fruit and seed characters, pollen apertures, and foliar epidermal walls (Pl.1A-C, 2A-C). In general, the numerical analysis based on the combined pollen morphology and foliar epidermal characters, along with morphological traits, is consistent with Lourteig’s (2000) taxonomic treatment concerning the sectional affinities of the studied taxa. 76 Bull. Iraq nat. Hist. Mus. 18 (1): 65-90. Palynological and foliar epidermal studies Plate (1): SEM micrographs of the Oxalis taxa pollen grains; (A-C) O. corniculata var. corniculata: A- Pollen in equatorial view, B- Pollen in polar view, C- Exine sculpture. (D-F) O. corniculata var. repens: D- Pollen in equatorial view, E- Pollen in polar view, F- Exine sculpture. (G-I) O. pes-caprae: G- Pollen in equatorial view, H- Pollen in polar view, I- Exine sculpture. (J-L) O. anthelmintica: J- Pollen in equatorial view, K- Pollen in polar view, L- Exine sculpture. (M-O) O. debilis: M- Pollen in equatorial view, N- Pollen in polar view, O- Exine sculpture. 77 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Shamso et al. Plate (2): Pollen with tricolpate apertures; (A) Oxalis corniculata var. corniculate, (B) O. pes-caprae and (C) O. anthelmintica. (A. polar view; B & C. polar and equatorial view). Plate (3): Pollen with tri- and tetra-colpate apertures; (A) Oxalis corniculata var. repens, (B) O. latifolia. Plate (4): Types of foliar trichomes; (A) Long unicellular with tapering apex (LM, X40), (B) Long unicellular with papillate surface (SEM, X 1600), (C) Short, clavate unicellular (SEM, X 1600), (D) Multicellular uniseriate (LM, X40). 78 Bull. Iraq nat. Hist. Mus. 18 (1): 65-90. Palynological and foliar epidermal studies Plate (5): T.S. in lamina to show the periclinal wall of the epidermal cells; (A) Semi swollen on abaxial surface in O. debilis, (B & C) Swollen on both surfaces in O. latifolia and O. corniculata var. repens respectively (LM, X40). Plate (6): Lamina surface to show the anticlinal wall pattern and periclinal wall surface; (A) straight anticlinal wall with papillate surface in O. latifolia, (B) slightly curved anticlinal wall with glebulate surface in O. pes-caprae, (C) undulate anticlinal wall with smooth surface in O. corniculata var. corniculata. (LM, X 40). Plate (7): Lamina abaxial surface to show stomatal types;(A) Anisocytic and unusual 4- celled anisocytic stomata in O. anthelmintica, (B) anomocytic stomata in O. debilis, (C & D) actinocytic stomata in O. corniculata var. repens and O. latifolia respectively (LM, X40). A C B A B C D 79 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Shamso et al. Plate (8): Lamina adaxial surface showing; (A) Ellipsoid and sunken stomata in O. debilis, (B) Oblong and slightly sunken stomata, unicellular clavate trichomes in O. corniculata var. corniculata (SEM, X1600). Diagram (1): Cluster dendrogram of the six taxonomic taxa based on a similarity matrix using a single linkage analysis software. CONCLUSIONS Despite that pollen morphology and foliar epidermal cell characters of the six taxa of Egyptian Oxalis described here show considerable agreement with previous characteristics found in other Oxalis taxa, their potential taxonomic significance may be limited. Nevertheless, when these data and morphological traits are combined, they become valuable for identifying and classifying the taxa at specific and sectional levels. Thus, this study can indeed serve as a reference for future studies of Oxalis taxa. 80 Bull. Iraq nat. Hist. Mus. 18 (1): 65-90. Palynological and foliar epidermal studies ACKNOWLEDGMENTS The authors are grateful to Dr. Amal Hosny (Botany and Microbiology Department, Faculty of Science, Cairo University) for valuable comments and critical revision of the manuscript. CONFLICT OF INTEREST STATEMENT "The authors declare no conflict of interest". LITERATURE CITED Abreu, M. C., Silva, M. J. and Sales, M. F. 2012. Análise cladística de Oxalis sec. Thamnoxys (Oxalidaceae) baseado em dados morfológicos. Rodriguésia, 63(4): 755-761. [CrossRef] Adedeji, O., Ajuwon, O. Y. and Babawale, O. O. 2007. Foliar epidermal studies, organographic distribution and taxonomic importance of trichomes in the family Solanaceae. 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[(1) Oxalis anthelmintica; (2) O.pes- caprae; (3) O.corniculata var. corniculata; (4) O.corniculata var. repens; (5) O. debilis; (6) O. latifolia]; (0 = absent; 1 = present) Character Sub- character Character state 1 2 3 4 5 6 1- Life form Epiphytes 0 0 1 1 0 0 Geophytes 1 1 0 0 1 1 2- Duration Perennial 1 1 0 0 1 1 Annual or perennial 0 0 1 1 0 0 3- Bulb Present 1 1 0 0 1 1 Absent 0 0 1 1 0 0 4- Number of bulbs Solitary 1 1 0 0 0 1 Numerous 0 0 0 0 1 0 5- Bulb shape Ovoid to pyriform 1 1 0 0 0 1 Globose 0 0 0 0 1 1 6- Rhizome Present 1 1 0 0 0 0 Absent 0 0 1 1 1 1 7- Rhizome colour Brownish 1 0 0 0 0 0 Whitish 0 1 0 0 0 0 8- Root type Fibrous 0 0 1 1 0 0 Contractile 1 1 0 0 0 1 Translucent tuberous 0 0 0 0 1 0 9- Stem presence Present 0 0 1 1 0 0 Absent 1 1 0 0 1 1 10- Stem surface Hairy 0 0 1 1 0 0 Glabrous 1 1 0 0 1 1 11- Leaves arrangement Alternate 0 0 1 1 0 0 Basal rosette 1 1 0 0 1 1 12- Petiole surface Hairy 0 0 1 1 1 1 Glabrous 1 1 0 0 0 0 https://doi.org/10.1016/j.jop.2018.01.003 86 Bull. 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Palynological and foliar epidermal studies 13- Petiole flexuous near the base Present 0 0 0 0 1 0 Absent 1 1 1 1 0 1 14- Stipule shape Broadly oblong 0 0 1 1 0 0 Ovate – oblong 1 1 1 0 0 0 Elliptic 1 0 0 0 0 0 Oblong-lanceolate 0 0 0 0 1 0 Ovate or rectangular 0 0 0 0 0 1 15- Auriculate apex of stipules Present 0 0 1 0 1 1 Absent 1 1 0 1 0 0 16- Stipules with nerves Present 0 0 0 0 1 1 Absent 1 1 1 1 0 0 17- Leaflet shape Obdeltoid 0 0 0 0 0 1 Obcordate 0 1 1 1 1 0 Subcircular to obovate 1 0 0 0 0 0 18- Leaflt's lobes symmetry Symmetrical 1 0 1 1 1 1 Asymmetrical 0 1 0 0 0 0 19- Leaflet dotting Present 1 1 0 0 1 0 Absent 0 0 1 1 0 1 20- Sinus width Wide 0 1 0 1 0 1 Narrow 1 0 1 0 1 0 21- Sinus depth Deep 0 0 0 1 0 0 Shallow 1 1 1 0 1 1 22- Peduncle surface Hairy 1 0 1 1 1 1 Glabrous 0 1 0 0 0 0 23- Bracts relative lengths Equal bracts 1 1 0 0 1 1 Unequal bracts 0 0 1 1 0 0 24- Bract apex with brownish calli Present 0 1 0 0 1 0 Absent 1 0 1 1 0 1 25- Bracteole presence Present 0 0 1 0 1 0 87 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Shamso et al. Absent 1 1 0 1 0 1 26- Flower color Yellow 0 1 1 1 0 0 Pink to purple 1 0 0 0 1 1 27- Sepal shape Oblong- lanceolate 0 1 1 1 0 0 Lanceolate to ovate-lanceolate 1 0 0 0 1 1 28- Sepal's calli Present 1 1 0 0 1 1 Absent 0 0 1 1 0 0 29- Petal shape Spathulate 1 1 1 1 0 0 Obovate 0 0 0 0 1 1 30- Long stamens with appendage Present 0 1 0 0 0 0 Absent 1 0 1 1 1 1 31-Long stamens surface Hairy 1 0 0 0 1 1 Glabrous 0 1 1 1 0 0 32-Shape of stigma Capitate 1 1 1 1 0 1 Laciniate 0 0 0 0 1 0 33- Pollen shape in equatorial view Prolate- spheroidal 0 0 1 0 0 0 Subprolate 1 1 0 0 1 1 Oblate- spheroidal 0 0 0 1 0 0 34- Pollen aperture type Tri-zonocolpate 1 1 1 0 0 0 Tri- or tetra- zonocolpate 0 0 0 1 1 1 35- Colpus shape Elliptic 0 0 1 0 0 0 Fusiform 0 1 0 0 1 1 Oblong 0 0 0 1 0 0 Fusiform or oblong 1 0 0 0 0 0 36- Colpus membrane sculpture Densely granulated to rugulate 0 0 1 0 0 0 With congregated granules distributed irregularly 0 1 0 0 0 0 Scabrous to warty 0 0 0 1 0 0 Warty 1 0 0 0 1 1 88 Bull. 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Palynological and foliar epidermal studies 37- Exine sculpture Microreticulate 1 0 1 1 0 1 Reticulate, lumina with single baculum 0 1 0 0 0 0 Perforate or microreticulate 0 0 0 0 1 0 38- P/E ratio > 1 1 1 1 0 1 1 < 1 0 0 0 1 0 0 39- Epidermal anticlinal wall Undulate 0 1 1 0 1 0 Straight 1 0 0 1 0 1 40- Epidermal anticlinal wall thickness Thick 0 0 0 0 0 1 Thin 1 1 1 1 1 0 41- Adaxial epidermal periclinal wall Swollen 0 0 0 1 0 0 Semiswollen 1 1 1 0 1 0 Papillate 0 0 0 0 0 1 42- Abaxial epidermal periclinal wall Swollen 0 1 0 1 0 1 Semiswollen 1 0 1 0 1 0 43- Epidermal cell surface sculpture Reticulate- foveolate 0 0 1 0 0 1 Smooth 0 0 0 1 1 0 Glebulate 1 1 0 0 0 0 44- Epicuticular wax on epidermis Present 0 1 1 1 1 1 Absent 1 0 0 0 0 0 45- Stomatal position Hypostomatic 1 1 0 0 0 1 Amphistomatic 0 0 1 1 1 0 46- Stomatal types Anomocytic 0 0 1 1 1 0 Actinocytic 0 0 0 1 0 1 Anisocytic 1 1 1 1 1 1 Unusual 4-celled anisocytic 1 1 1 1 1 1 47- Stomatal shape Oblong 1 0 1 0 0 0 Silt 0 0 0 1 0 0 Elliptic 0 1 0 0 1 1 48- Trichome type Unicellular with tapering apex 0 1 1 1 1 0 Multicellular 0 0 0 0 1 0 Clavate unicellular 1 0 0 0 1 1 89 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Shamso et al. Bull. Iraq nat. Hist. Mus. (2024) 18 (1): 65-90. Oxalis Linnaeus, 1753ألوراق جنس حبوب اللقاح والتشريح السطحي ةدراس ( في مصرOxalidales، رتبة Oxalidaceae عائلة) إيمان محمود شمسو*، حسناء أحمد حسني*، أمل عبد البصير دراز**، *ينسامح رضا حس و سلوى علي القواتش ي** الجيزة ،جامعة القاهرة /العلومكلية /قسم النبات وامليكروبيولوجي /*املعشبة .مصر ،12613ص ب الجيزة ،الدقي ،املركز القومى للبحوث/** قسم الكيمياء و تصنيف النباتات .مصر ،12622ص ب 20/6/2024، النشر: 3/12/2023القبول: ،26/11/2023املراجعة: ، 28/9/2023االستالم: الخالصة ُدِرَست(، SEMمليكروسكوب اإللكتروني املاسح )باستخدام امليكروسكوب الضوئي وا فين من يحبوب اللقاح و مورفولوجيا البشرة الخارجية لألوراق ألربعة أنواع وصن التابعة لثالثة أقسام في لجنيس Oxalis Linnaeus, 1753األوكساليس املصرية Oxalis Linnaeus, 1753 وب تحديد أهمية خصائص حب الهدف من الدراسة، كان اللقاح و البشرة الخارجية لألوراق كأدلة تصنيفية. أظهرت الدراسة الحالية أن حبوب اللقاح في جميع األنواع املدروسة هي أحادية ومتماثلة، و من الحجم املتوسط، و يتراوح الكروي إلى تحت البيضوي. كما كانت فتحات حبوب اللقاح ثالثية الشكل من تحت الجدار الخارجي كان الثقوب مع أغشية مزخرفة )بنمط مموج، أو نتوءات متجمعة(. ,Kunth شبكي إلى شبكي، ومثقب في-لحبوب اللقاح شبه مغطى، مع تزخرف ميكرو 1822 O. debilisمن الثغور: . كما كشفت الدراسة عن وجود أربعة أنواعactinocytic, anisocytic, anomocytic and an unusual 4-celled anisocytic . كانت خاليا البشرة العليا والسفلية غير منتظمة أو مضلعة الشكل، مع جدران متموجة أو مستقيمة، وجدران داخلية شبه منتفخة إلى منتفخة مع وجود طبقة من الشعيرات غير الغددية وأحادية الخلية: شعر عانلوحظ نو شمعية خارجية منتشرة. 90 Bull. Iraq nat. Hist. Mus. 18 (1): 65-90. Palynological and foliar epidermal studies قصير بشكل مدبب وشعر طويل ذو طرف مدبب مع سطح محبب. ناقشت الدراسةأيضا املجمعة وخصائص حبوب اللقاح املظهريةالنتائج العددية استناًدا إلى الخصائص لتصنيف والبشرة الخارجية لألوراق. باإلضافة إلى ذلك، تم تصميم مفتاح اصطناعي األنواع املدروسة. تعتبر هذه الدراسة أول تقرير يتعامل مع الخصائص التشريحية جنس في مصر. هذا الوحبوب اللقاح ألنواع