Research article

Assessment of plant diversity of endemic species of the Saharo-Arabian region in Egypt

  • Asmaa S ABO HATAB 1 ,
  • Yassin M AL-SODANY 1 ,
  • Kamal H SHALTOUT 2 ,
  • Soliman A HAROUN 1 ,
  • Mohamed M EL-KHALAFY , 1, *
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  • 1Botany and Microbiology Department, Faculty of Science, Kafrelsheikh University, Kafrelsheikh 33511, Egypt
  • 2Botany Department, Faculty of Science, Tanta University, Tanta 31527, Egypt
* Mohamed M EL-KHALAFY (E-mail: )

Received date: 2024-01-25

  Revised date: 2024-05-24

  Accepted date: 2024-06-05

  Online published: 2025-08-14

Abstract

Savanna, semi-deserts, and hot deserts characterize the Saharo-Arabian region, which includes Morocco, Mauretania, Algeria, Tunisia, Libya, Egypt, Palestine, Kuwait, Saudi Arabia, Qatar, Bahrain, the United Arab Emirates, Oman, Yemen, southern Jordan, Syria, Iraq, Iran, Afghanistan, Pakistan, and northern India. Its neighboring regions, the Sudano-Zambezian region belonging to the Paleotropical Kingdom and the Mediterranean and Irano-Turanian regions included in the Holarctic Kingdom, share a large portion of their flora with the Saharo-Arabian region. Despite the widespread acknowledgment of the region's global importance for plant diversity, an up to date list of the Saharo-Arabian endemics is still unavailable. The available data are frequently insufficient or out of date at both the whole global and the national scales. Therefore, the present study aims at screening and verifying the Saharo-Arabian endemic plants and determining the phytogeographical distribution of these taxa in the Egyptian flora. Hence, a preliminary list of 429 Saharo-Arabian endemic plants in Egypt was compiled from the available literature. Indeed, by excluding the species that were recorded in any countries or regions outside the Saharo-Arabian region based on different literature, database reviews, and websites, the present study has reduced this number to 126 taxa belonging to 87 genera and 37 families. Regarding the national geographic distribution, South Sinai is the richest region with 83 endemic species compared with other eight phytogeographic regions in Egypt, followed by the Isthmic Desert (the middle of Sinai Peninsula, 53 taxa). Sahara regional subzone (SS1) distributes all the 126 endemic species, Arabian regional subzone (SS2) owns 79 taxa, and Nubo-Sindian subzone (SS3) distributes only 14 endemics. Seven groups were recognized at the fourth level of classification as a result of the application of the two-way indicator species analysis (TWINSPAN) to the Saharo-Arabian endemic species in Egypt, i.e., I Asphodelus refractus group, II Agathophora alopecuroides var. papillosa group, III Anvillea garcinii group, IV Reseda muricata group, V Agathophora alopecuroides var. alopecuroides group, VI Scrophularia deserti group, and VII Astragalus schimperi group. It's crucial to clearly define the Saharo-Arabian endemics and illustrate an updated verified database of these taxa for a given territory for providing future management plans that support the conservation and sustainable use of these valuable species under current thought-provoking devastating impacts of rapid anthropogenic and climate change in this region.

Cite this article

Asmaa S ABO HATAB , Yassin M AL-SODANY , Kamal H SHALTOUT , Soliman A HAROUN , Mohamed M EL-KHALAFY . Assessment of plant diversity of endemic species of the Saharo-Arabian region in Egypt[J]. Journal of Arid Land, 2024 , 16(7) : 1000 -1021 . DOI: 10.1007/s40333-024-0102-3

1 Introduction

The desert biome includes a number of geographic regions distinguished by a high level of aridity, a significant amount of bare soil, and some organisms that have developed specific adaptations to survive in a very dry climate (Ezcurra et al., 2006). According to Costanza et al. (2014), deserts occupy almost 33.0% of the Earth's surface. Deserts are susceptible to climate variability due to their hot climate and water scarcity issues (Meigs, 1953; El-Nesr et al., 2010). Based on the rate of precipitation, Wickens (1998) divided the world's desert areas into three categories: semi-arid areas, which receive an average of between 200 and 400 mm of precipitation annually; arid areas, which receive less than 200 mm of precipitation annually; and hyper-arid areas, with an average of 12 months without precipitation. There are several different types of deserts in the world, including coastal, tropical and subtropical, rain shadow, continental interior, and polar deserts (Marshak, 2015); usually, these deserts are found in rocky or sandy landscapes (Smith and Pettorelli, 2020). The climates of deserts and semi-deserts are characterized by major continental rain shadows and subtropical high-pressure belts (Keith et al., 2020). On the other hand, the impacts of atmospheric stability can be increased by a combination of geographic, topographic, and oceanographic factors. For example, the location and size of Sahara and Arabian deserts are influenced by the high-pressure belt that lies over the Tropic of Cancer and the effects of continental (Nash, 2012).
As with all types of desert geomorphology, Saharo-Arabian floristic region has deep-rooted plants and shaded therophytes, salt and limestone crusts, and biotic crusts that stabilize sand dunes (Breckle et al., 2001). The Saharo-Arabian region also belongs to shield deserts that formed above very old crystalline floors, which were cracked and reinforced by pressure and heat during many millions of years, resulting in gently sloping swells and basins that retain the best groundwater sources (Ezcurra et al., 2006). Some deserts, such as the Sonoran-Chihuahuan region of the United States and Mexico and the Karoo of South Africa, are key biodiversity hotspots on a global scale (Myers et al., 2000), supporting human cultures and providing a variety of local and international ecosystem services (McClung et al., 2019). The Sahara Desert and the nearby arid Sahel are habitats for high number of endemic species and a patchy distribution region of other species that are limited to small, vulnerable, and humid habitats (Brito et al., 2014). Isolated and remnant bodies of water (lakes, seasonal rivers, and oases) surrounded by sand act as refugia for relict populations and serve as sites for the evolution of rare species (Trape, 2013). Indeed, the presence of water in drylands is a powerful predictor of species richness and community distribution (Cornwell and Ackerly, 2009).
The Saharo-Arabian region, from a phytogeographic perspective, is the vast desert belt that stretches eastward from the Atlantic coast of Morocco and Mauritania, through the Sahara Desert, Sinai Peninsula, extra tropical Arabia, southern Iraq, Iran, and Afghanistan, and terminates in the deserts of India and Pakistan (Sind Provinece, Thar Desert, and Punjab Province) (Zohary, 1973). Its borders are the Mediterranean region in the north and west, the Irano-Turanian region in the east, and the Sudano-Zambezian region and the Tropic of Cancer in the south. It is characterized by extremely dry air, high temperatures, and little precipitation (Heine, 1962). Because of the length of dry season, the low continentality index, and mild winters, the Saharo-Arabian region as a whole differs climatically from its neighbors, particularly the Irano-Turanian neighborhood (Léonard, 1988). Precipitation during the summer in the Saharo-Arabian region varies significantly geographically between the west of Sahara Desert and its east. It is commonly believed that topographic heterogeneity raises the endemism rate (Djamali et al., 2012). According to White and Léonard (1991), they divided the Saharo-Arabian region into three subzones: Sahara regional subzone (SS1), Arabian regional subzone (SS2), and Nubo-Sindian subzone (SS3). White and Léonard (1991) stated that the majority of Egypt excepting for the Mediterranean coastal strip and Elba Mountains is located in the SS1 zone.
White and Léonard (1991) revised the approach of Zohary (1973) and extended the Somalia-Masai regional center of endemism and the Afro-Montane archipelago-like regional center of endemism into southwestern Arabia and divided Saharo-Sindian zone into SS1, SS2, and SS3. The SS1 subzone includes Morocco, Mauritania, Algeria, Tunisia, Libya, and Egypt. The SS2 subzone represents the eastern Saharo-Arabian of Zohary (1973), including Palestine, Kuwait, Saudi Arabia, Qatar, Bahrain, the United Arab Emirates, Oman, Yemen, and south of Jordan, Syria, and Iraq. And the SS3 subzone represents the old Sudano-Decanian tropical vegetation of Zohary (1973), including south of Iran, Afghanistan, Pakistan, and northern India.
Endemic taxa are those that are found only in a certain place and represent an essential feature of spatial distribution (Morrone, 2008). Because they have distinctive genetic materials and are capable of satisfying the fundamental necessities of people, endemic plants are highly valued (Skarbek, 2008). However, they are more vulnerable to changes in the environment and threats caused by humans. Because of this, endemic plants are more likely to become extinct. As a result, conserving endemic plants is regarded as a global concern (El-Khalafy et al., 2021a). Manes et al. (2021) stated that globally, endemic species are three times more likely to become extinct than native species. Endemism is mentioned as a stage before extinction by Hobohm et al. (2014). As a result, endemic plants are the focus of most global conservation efforts, which are mostly carried out in situ through protected areas, ex situ through gardening, and cryo-conservation through the banks of seeds and biotechnological approaches (Coelho et al., 2020).
Few endemic species and genera, as well as none endemic families, are characteristics of the Saharo-Arabian region, according to Wickens (1977) and Boulos (1997). The vegetation of the deserts in Egypt is certainly a unique type of natural plant life (El-Amier and Abdul-Kader, 2015). According to Moustafa (1990) and El-Khalafy et al. (2021a), South Sinai Mountains are a significant habitat for Egypt's endemism because they have larger outcrops of rare species-supporting smooth-faced rocks and a more humid climate than much of the rest of Sinai Peninsula (Danin, 1972, 1978, 1983). Therefore, the mountainous southern Sinai Peninsula has a higher biodiversity as compared with the rest of Egypt (Abd El-Ghani et al., 2017).
Deserts are a significant conservation concern because of their great biodiversity, uniqueness, and rising survival hazards. The future survival of deserts and their species relies on developing strategies for conservation that are capable of dealing with the challenges provided by rapidly shifting climatic conditions. Regional biodiversity knowledge is typically lacking or outdated (Brito et al., 2018). There is no enough data about the list of endemic plant species of the Saharo-Arabian region in Egypt and the vegetation distribution map of these species. Therefore, an accurate list of endemic species is necessary to understand species diversity in regions under heavy anthropogenic impact. It can be utilized as a baseline reference for future monitoring programs as well as to offer data for future in situ and ex situ conservation planning activities. So, this study aims to provide an authenticated database of endemic plant species of the Saharo-Arabian region in Egypt and study the national and global distribution of these species.

2 Materials and methods

2.1 Study area

Less than 25.0% of Egypt's land area is in the south of the Tropic of Cancer, and it is located between latitudes 22° and 32°N, and longitudes 24° and 36°E (Egypt is primarily in the temperate zone). The study area of Egypt includes: the Western Desert except the western Mediterranean coast that extends for an average distance of 45-50 km in north-south direction from the coast (Bidak et al., 2013), the Eastern Desert with Red Sea except the Gebel Elba Mountains, the Sinai Peninsula except the eastern Mediterranean coast that extends for an average distance of 10-15 km in north-south direction from the coast (Zahran et al., 1985), and the Nile Valley and Nile Delta except the coastal land of northern delta that extends for 15-25 km in north-south direction from the coast (Zahran and Willis, 2009). The Saharo-Arabian region in this study includes: Morocco, Mauretania, Algeria, Tunisia, Libya, Egypt, Palestine, Kuwait, Saudi Arabia, Qatar, Bahrain, the United Arab Emirates, Oman, Yemen, southern Iraq, Iran, Afghanistan, and Pakistan, as well as northern India (21 countries in total). Due to lack of data, Western Sahara is not included in the research scope of this paper. Lebanon belongs to the Mediterranean region, so Lebanon is also not included in this paper.

2.2 Data sources and verification literature

2.2.1 Endemic species

The preliminary list of the Saharo-Arabian endemics was compiled from three primary sources that researched the plants of this region. These sources include Bornkamm and Kehl (1990), Boulos (1999-2005), and Abd El-Ghani et al. (2017). Indeed, Boulos (1999-2005) recorded 199 Saharo-Arabian endemic species in the Egyptian flora, while Abd El-Ghani et al. (2017) recorded 187 taxa and Bornkamm and Kehl (1990) recorded 102 Saharo-Arabian endemic species in the Western Desert region of Egypt. All the plants identified in these studies as the Saharo-Arabian endemic plants were included in the preliminary list. Finally, we got a preliminary list with 429 taxa (Table S1).
Table S1 Updated list of the Saharo-Arabian endemic plants in Egypt
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
1 Abutilon pannosum (G.Forst.) Schltdl. Malvaceae Angola, Burkina, Cameroon, Cape Verde, and Chad (SZ) Boulos (1999-2005); GBIF; and POWO × ×
Gebel Elba (SZ) Abutaha et al. (2020)
2 Aeluropus littoralis (Gouan) Parl. Poaceae Baleares, Bulgaria, and France, and Spain (ME) GBIF and POWO × ×
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
3 Aerva javanica (Burm.f.) Juss. ex Schult. Amaranthaceae Botswana, Burkina, Cameroon, Cape Verde, and Chad (SZ) GBIF and POWO × × ×
Gebel Elba (SZ) Boulos (1999-2005)
4 Agathophora alopecuroides var. alopecuroides (Delile) Fenzl ex Bun Amaranthaceae
5 Agathophora alopecuroides var. papillosa (Maire) Boulos Amaranthaceae
6 Allium crameri Asch. & Boiss. Amaryllidaceae Bardawil Lake and Mediterranean coast of Rafah (ME) CAIM (1998); Nada et al. (2013); and El-Khalafy et al. (2021a, b)
7 Allium decaisnei C. Presl Amaryllidaceae
8 Allium papillare Boiss. Amaryllidaceae Bardawil Lake and Rafah (ME) CAIM (1928); Boulos (1999-2005); El-Garf (2000); and El-Bana (2009) ×
9 Allium rothii Zucc. Amaryllidaceae
10 Allium sinaiticum Boiss. Amaryllidaceae
11 Anabasis articulata (Forssk.) Moq. Amaranthaceae Spain (ME) GBIF × × ×
Mediterranen coast in Egypt, Rafah and Sallum (ME) Boulos (1999-2005); Khafagi et al. (2005); Amer et al. (2020); and Gamal et al. (2022)
12 Anarrhinum forskaohlii subsp. pubescens D. A. Sutton Plantaginaceae
13 Anastatica hierochuntica L. Brassicaceae Canary Islands (ME) POWO × ×
Sudan and Eritrea (SZ) POWO
Gebel Elba (SZ) Boulos (1999-2005)
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
14 Anthemis melampodina subsp. deserti (Boiss.) Eig Asteraceae Mediterranen coast in Egypt (ME) Täckholm (1974); Boulos (1999-2005); Fakhry and Al-Anazi (2017); and Osman and Abdein (2019) ×
15 Anthemis melampodina subsp. melampodina Asteraceae Mediterranen coast in Egypt (ME) Täckholm (1974); Boulos (1999-2005); Fakhry and Al-Anazi (2017); and Osman and Abdein (2019) ×
16 Anthemis scrobicularis Yavin Asteraceae
17 Anthemis zoharyana Eig Asteraceae Mediterranen coast in Egypt (ME) Täckholm (1974)
18 Anvillea garcinii (Burm. f.) DC. Asteraceae
19 Argyrolobium saharae Pomel Fabaceae
20 Aristida funiculata Trin. & Rupr. Poaceae Burkina, Cape Verde, Chad, Eritrea, Ethiopia, Niger, Kenya, Senegal, and Somalia (SZ) Boulos (1999-2005); GBIF; and POWO × ×
Mediterranen coast in Lybia (ME) Al-Sghair et al. (2019)
Gebel Elba (SZ) Boulos (1999-2005)
21 Arnebia hispidissima (Sieber ex Lehm.) A.DC. Boraginaceae Cameroon, Chad, Eritrea, Ethiopia, Somalia, Niger, and Sudan (SZ) Boulos (1999-2005); GBIF; and POWO × ×
Gebel Elba (SZ) Boulos (1999-2005)
22 Arnebia tinctoria Forssk. Boraginaceae
23 Artemisia judaica L. Asteraceae Chad (SZ) GBIFand POWO ×
Gebel Elba (SZ) Boulos (1999-2005)
24 Artemisia monosperma Delile Asteraceae Al-Arish (North Sinai), Burg El-Arab, Burullus Wetland, and Mediterranen coast in Egypt (ME) Boulos (1999-2005); Al-Sodany (2009); and Badr et al. (2012) ×
25 Asphodelus refractus Boiss. Asphodelaceae
26 Asphodelus viscidulus Boiss. Asphodelaceae El Burullus and Mediterranen coast in Egypt (ME) Boulos (1999-2005) and Galal and Fawzy (2007) ×
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
27 Asteriscus graveolens (Forssk.) Less. Asteraceae Canary Islands (ME) GBIFand POWO × ×
Chad and Mali (SZ) GBIFand POWO
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
28 Astragalus amalecitanus Boiss. Fabaceae
29 Astragalus annularis Forssk. Fabaceae Lebanon (ME) GBIF × ×
Mediterranen coast in Egypt, Rosetta, and Sallum (ME) Boulos (1999-2005); El-Hawiet et al. (2010); and Amer et al. (2020)
30 Astragalus arpilobus subsp. hauarensis (Boiss.) Podlech Fabaceae North of Iran (IT) GBIF ×
Mediterranen coast in Egypt (ME) Täckholm (1974) and Boulos (1999-2005)
31 Astragalus bombycinus Boiss. Fabaceae North of Iran (IT) Asri (2004) and Osman et al. (2014) ×
32 Astragalus camelorum Barbey Fabaceae El-Arish and Mediterranen coast in Egypt (ME) Täckholm (1974) and CAI (1986)
33 Astragalus caprinus L. Fabaceae Cyprus, and Sicilia (ME) GBIFand POWO × ×
Iran and Turkey (IT) Flora of Libya
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
34 Astragalus dactylocarpus subsp. acinaciferus (Boiss.) E. Ott Fabaceae
35 Astragalus eremophilus Boiss. Fabaceae Chad, Eritrea, and Sudan (SZ) Khdery et al. (2019) and POWO × × ×
Gebel Elba (SZ) Boulos (1999-2005)
36 Astragalus fresenii Decne Fabaceae
37 Astragalus hispidulus DC. Fabaceae Coastal area in Libya (ME) Baayo (2008) × ×
Mediterranen coast in Egypt and Burg el Arab (ME) Boulos (1999-2005) and GBIF
38 Astragalus intercedens Sam. ex Rech.f. Fabaceae
39 Astragalus kahiricus DC. Fabaceae North of Iran, Tadzhikistan, Turkmenistan, and Uzbekistan (IT) Rahimi and Atri (2013); Flora of Libya; GBIF; and POWO ×
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
40 Astragalus palaestinus Eig Fabaceae Mediterranen coast in Egypt (ME) Danin and Fragman-Sapir (2016)
41 Astragalus peregrinus subsp. peregrinus Vahl. Fabaceae Greece and Kriti (ME) Flora of Greece (2023) and POWO ×
El-Arish and Mediterranen coast in Egypt (ME) Boulos (1999-2005) and Badr and Sharawy (2007)
42 Astragalus sanctus Boiss. Fabaceae
43 Astragalus schimperi Boiss. Fabaceae
44 Astragalus sieberi DC. Fabaceae Alexandria El-Alamein desert road, Burullus Wetland, and Matrouh road (ME) Al-Sodany (2009) and Abd El-Ghani et al. (2021)
45 Astragalus sparsus Decne Fabaceae
46 Astragalus trigonus DC Fabaceae Chad, Mali, Niger, and Sudan (SZ) GBIF and POWO × × ×
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
47 Astragalus vogelii (Webb) Bornm. Fabaceae Cape Verde, Eritrea, Ethiopia, Mali, Niger, and Sudan (SZ) Boulos (1999-2005); GBIF; and POWO × × ×
48 Atractylis boulosii Tackh. Asteraceae
49 Atractylis carduus (Forssk.) C. Chr. Asteraceae Mediterranen coast in Egypt and Turkey (ME) Flora of Libya and POWO × ×
Mediterranen coast in Egypt (ME) Täckholm (1974) and Boulos (1999-2005
50 Atractylis mernephthae Asch. & Schweinf. & Letourn. Asteraceae
51 Atractylis prolifera Boiss. Asteraceae Coastal area of Libya and Spain (ME) Flora of Libya and GBIF ×
Mediterranen coast in Egypt, and the northwestern coastal desert of Egypt (ME) Boulos (1999-2005) and Bidak et al. (2015)
52 Atractylis serratuloides Sieber ex Cass. Asteraceae Coastal area of Libya (ME) Flora of Libya
North of Sinai (ME) Täckholm (1974)
53 Atraphaxis spinosa var. sinaica (Jaub. & Spach) Boiss. Polygonaceae
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
54 Atriplex coriacea Forssk. Amaranthaceae Somalia (SZ) GBIF ×
Alexandria, Matrouh, and Mediterranen coast in Egypt (ME) CAI (1959); Boulos (1999-2005)
55 Atriplex dimorphostegia Kar. et Kir. Amaranthaceae Kazakhstan, Kirgizstan, Tadzhikistan, Turkmenistan, Uzbekistan, and Xinjiang Uygur Autonomous Region, China (IT) GBIF and POWO × ×
56 Atriplex farinosa Forssk. Amaranthaceae Eritrea, Kenya, Somalia, and Sudan (SZ) GBIF and POWO ×
57 Atriplex glauca L. Amaranthaceae Canary Islands, Madeira, Portugal, and Spain (ME) Boulos (1999-2005); GBIF; and POWO × ×
58 Atriplex portulacoides L. Amaranthaceae Albania, Baleares, Belgium, Canary Islands, Cyprus, and Denmark (ME) Abd El-Ghani et al. (2017); GBIF; and POWO × × ×
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
59 Atriplex turcomanica (Moq.) Boiss. Amaranthaceae Kazakhstan, Transcaucasus, Turkey, Turkmenistan, and Uzbekistan (IT) Abd El-Ghani et al. (2017); GBIF; and POWO × ×
60 Avicennia marina (Forssk.) Vierh. Acanthaceae Cape Provinces, Eritrea, and Kenya (SZ) GBIF and POWO × ×
61 Ballota kaiseri Tackh. Lamiaceae
62 Bassia arabica (Boiss.) Maire & Weiller Amaranthaceae Mariut and Mediterranen coast in Egypt (ME) Squires and Ayoub (1994); Boulos (1999-2005); and Turki et al. (2006) ×
63 Bellevalia desertorum Eig & Feinbrun Asparagaceae
64 Bellevalia eigii Feinbrun Asparagaceae Burg el Arab, Mariut, Mediterranen coast in Egypt, and Sallum (ME) Boulos (1999-2005); Amer at al. (2020); and GBIF ×
65 Bellevalia flexuosa var. galalensis Tackh. & Drar Asparagaceae
66 Bellevalia zoharyi Feinbrun Asparagaceae
67 Biarum olivieri Blume Araceae Bardawil Lake, Damiette, and Mediterranen coast in Egypt (ME) Boulos (1999-2005); El-Sheikh (2013); and GBIF ×
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
68 Blepharis attenuata Napper Acanthaceae West and central Jordan (IT) Danin and Fragman-Sapir (2016)
69 Blepharis edulis (Forssk.) Pers. Acanthaceae Angola, Chad, Eritrea, Ethiopia, Kenya, Mali, Niger, Somalia, Sudan, Tanzania, and Uganda (SZ) Boulos (1999-2005); GBIF; and POWO × ×
70 Blumea bovei (DC.) Vatke Asteraceae Chad, Eritrea, Ethiopia, Somalia, and Sudan (SZ) Boulos (1999-2005); GBIF; and POWO × ×
71 Brassica deserti Danin & Hedge Brassicaceae El-Arish and north Sinai (ME) Boulos (1999-2005) and Shehata and Kamel (2007)
72 Brocchia cinerea (Delile) Vis. Asteraceae Ethiopia, Mali, and Sudan (SZ) Boulos (1999-2005); GBIF; and POWO × ×
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
73 Bufonia multiceps Decne. Caryophyllaceae
74 Calendula tripterocarpa Rupr. Asteraceae France and Spain (ME) GBIF and POWO ×
75 Calotropis procera (Aiton) W. T. Aiton Apocynaceae Bangladesh, Benin, Burkina, Cambodia, Cameroon, Canary Islands, Cape Verde, Chad, Congo, Eritrea, and Ethiopia (SZ) GBIF and POWO × × ×
Gebel Elba (SZ) Boulos (1999-2005)
76 Campanula dulcis Decne. Campanulaceae
77 Capparis cartilaginea Decne. Capparaceae Chad, Eritrea, Ethiopia, Kenya, Somalia, Sudan, and Tanzania (SZ) GBIF and POWO × ×
Gebel Elba (SZ) Boulos (1999-2005)
78 Capparis spinosa var. aegyptia (Lam.) Boiss. Capparaceae Greece, Kriti, and Turkey (ME) GBIFand POWO × ×
79 Carduus getulus Pomel Asteraceae Mediterranen coast in Egypt (ME) Boulos (1999-2005) ×
80 Caroxylon imbricatum (Forssk.) Moq. Amaranthaceae Chad, Eritrea, Gebel Elbar, Senegal, Somalia, and Sudan (SZ) GBIF and POWO × × ×
Gebel Elba (SZ) Boulos (1999-2005) and Elkordy et al. (2022)
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
81 Caroxylon tetragonum (Delile) Moq. Amaranthaceae Matrouh and Mediterranen coast in Egypt (ME) Turki (1999) and Boulos (1999-2005) ×
82 Caroxylon tetrandrum (Forssk.) Akhani & Roalson Amaranthaceae Canary Islands and Spain (ME) Boulos (1999-2005) and GBIF × ×
83 Caroxylon villosum (Schult.) Akhani & Roalson Amaranthaceae Mediterranen coast in Egypt and north of Syria (ME) Mosallam (2007) and GBIF ×
The northwestern coastal desert of Egypt (ME) Bidak et al. (2015)
84 Caroxylon volkensii (Schweinf. & Asch.) Akhani & Roalson Amaranthaceae Mediterranen coast in Egypt (ME) Boulos (1999-2005); El-Shamy (2016); and Alghanem et al. (2020) ×
85 Carrichtera annua (L.) DC. Brassicaceae Canary Islands, Cyprus, Greece, Kriti, and Portugal (ME) Boulos (1999-2005); GBIF; and POWO × × ×
86 Carthamus eriocephalus (Boiss.) Greuter Asteraceae Matrouh (ME) El-Zanaty et al. (2010) ×
87 Phonus mareoticus (Delile) G.López Asteraceae Alexandria and Sallum (ME) Boulos (1999-2005) and Salama et al. (2005) × ×
88 Caudanthera sinaica (Decne.) Plowes Apocynaceae South of Yemen and Sudan (SZ) GBIF and POWO ×
89 Caylusea hexagyna (Forssk.) M.L. Green Resedaceae Cape Verde, Chad, Eritrea, Mali, Niger, Somalia, and Sudan (SZ) GBIF and POWO × ×
Gebel Elba (SZ) and Mediterranen coast in Egypt (ME) Boulos (1999-2005)
90 Cenchrus divisus (J.F.Gmel.) Verloove, Govaerts & Buttler Poaceae Chad, Eritrea, Niger, and Somalia (SZ) Boulos (1999-2005); GBIF; and POWO × ×
91 Centaurea aegyptiaca L. Asteraceae Sudan (SZ) POWO ×
Gebel Elba (SZ) Boulos (1999-2005)
92 Centaurea ammocyanus Boiss. Asteraceae Lebanon (ME) Boulos (1999-2005) ×
North of Iran (IT) GBIF
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
93 Centaurea glomerata Vahl Asteraceae Italy and Spain (ME) GBIF and POWO × × ×
Mediterranen coast in Egypt (ME) Boulos (1999-2005); Shaltout et al. (2018); and Hamed et al. (2021)
94 Centaurea pallescens Delile Asteraceae Lebanon, France, and Mediterranen coast in Egypt (ME) Boulos (1999-2005); Hamed et al. (2021); GBIF; Euro+Med); Flora of Lebanon; and GBIF × ×
95 Centaurea scoparia Sieber ex Spreng Asteraceae
96 Centaurea sinaica DC. Asteraceae
97 Centaurium malzacianum Maire Gentianaceae
98 Chiliadenus montanus (Vahl) Brullo Asteraceae
99 Citrullus colocynthis (L.) Schrad. Cucurbitaceae Bangladesh, Benin, Burkina, Cape Verde, Chad, Cyprus, Eritrea, and Ethiopia (SZ) and Canary Islands (ME) Boulos (1999-2005); GBIF; and POWO × × ×
Mediterranean coast in Egypt (ME) Abd El-Ghani et al. (2017)
100 Cleome arabica L Cleomaceae Chad, Eritrea, Ethiopia, Niger, and Sudan (SZ) GBIF and POWO ×
101 Cleome brachycarpa Vahl ex DC. Cleomaceae Cape Verde, Chad, Eritrea, Ethiopia, Kenya, Mali, Niger, Somalia, and Sudan (SZ) GBIF and POWO × ×
Gebel Elba (SZ) Boulos (1999-2005)
102 Cleome chrysantha Decne. Cleomaceae Sudan and Ethiopia (SZ) GBIF and POWO × ×
Gebel Elba (SZ) Boulos (1999-2005)
103 Cleome droserifolia (Forssk.) Delile Cleomaceae Eritrea, Somalia, and Sudan (SZ) GBIF and POWO × × ×
Gebel Elba (SZ) Boulos (1999-2005)
104 Cebatha pendula (J. R. Forst. & G. Forst.) Kuntze Menispermaceae Cape Verde, Ethiopia, and Somalia (SZ) GBIF and POWO × × ×
Gebel Elba (SZ) Boulos (1999-2005)
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
105 Colchicum guessfeldtianum Asch. & Schweinf. Colchicaceae
106 Colchicum ritchii R.Br. Colchicaceae Mediterranen coast in Egypt (ME) Boulos (1999-2005) and Flora of Libya ×
107 Colchicum schimperi Janka ex Stef Colchicaceae El Arish (ME) CAI (1965)
108 Cometes surattensis Burm.f. Caryophyllaceae South of Oman (SZ), India, Iran, Iraq, and Pakistan (IT) Al-Sherif et al. (2013) and POWO
109 Commicarpus boissieri (Heimerl) Cufod. Nyctaginaceae South of Oman and south of Yemen (SZ) GBIF ×
110 Convolvulus fatmensis Kunze Convolvulaceae Chad, Mali, and Sudan (SZ) × ×
Lebanon and Mediterranen coast in Egypt (ME) Boulos (1999-2005)
111 Convolvulus hystrix Vahl Convolvulaceae Eritrea, Ethiopia, Somalia, and Sudan (SZ) Boulos (1999-2005); GBIF; and POWO × ×
Gebel Elba (SZ) Boulos (1999-2005)
112 Convolvulus lanatus Vahl Convolvulaceae Mediterranen coast in Egypt, and Tukey (ME) Boulos (1999-2005) and POWO ×
113 Convolvulus schimperi Boiss. Convolvulaceae
114 Convolvulus spicatus Peter ex Hallier f. Convolvulaceae
115 Cornulaca monacantha Delile Amaranthaceae Chad, Mali, and Niger (SZ) GBIF and POWO × × ×
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
116 Cotoneaster orbicularis Schltdl. Rosaceae ×
117 Crataegus sinaica Boiss. Rosaceae Cyprus and Turkey (ME) GBIF and POWO ×
118 Crepis nigricans Viv. Asteraceae
119 Crotalaria aegyptiaca Benth. Fabaceae Somalia (SZ) Boulos (1999-2005); GBIF; and POWO
120 Crucianella ciliata Lam. Rubiaceae ×
121 Crucianella membranacea Boiss. Rubiaceae Lebanon (ME) Boulos (1999-2005) ×
122 Crucihimalaya kneuckeri (Bornm.) Al-Shehbaz, O'Kane & R. A. Price Brassicaceae
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
123 Cucumis prophetarum L Cucurbitaceae Benin, Burkina, Burundi, Cameroon, Chad, Eritrea, and Ethiopia (SZ) Boulos (1999-2005); GBIF; and POWO × ×
124 Cullen plicatum (Delile) C. H. Stirt. Fabaceae Chad, Ethiopia, Mali, Sudan, and Somalia (SZ) Boulos (1999-2005); GBIF; and POWO × ×
125 Cuprella homalocarpa (Fisch. & C. A. Mey.) Salmerón-Sánchez, Mota & Fuertes Brassicaceae North of Sinai (ME) Täckholm (1974) ×
126 Cuscuta pedicellata Ledeb. Convolvulaceae Eritrea, Ethiopia, and Sudan (SZ) GBIF and POWO × ×
Kazakhstan, Kirgizstan, and Russia (IT) and South European (ME) GBIF and POWO
Gebel Elba (SZ) and Mediterranen coast in Egypt (ME) Boulos (1999-2005)
127 Cutandia memphitica (Spreng.) K. Richt. Poaceae Canary Islands, Cyprus, North Caucasus, Spain, Tadzhikistan, and Transcaucasus (ME) GBIF and POWO × × ×
Gebel Elba (SZ) and Mediterranen coast in Egypt (ME) Boulos (1999-2005)
128 Cymbopogon iwarancusa (Jones ex Roxb.) Schult. Poaceae China and Vietnam (IT) GBIF and POWO ×
129 Cynanchum acutum L Apocynaceae South Europe, Mediterranen coast in Egypt, Spain, and Ukraine (ME) Boulos (1999-2005) and GBIF × × ×
130 Cyperus capitatus Vand. Cyperaceae Bulgaria, Canary Islands, Cape Verde, Cyprus, France, Greece, Italy, and Kriti (ME) GBIF and POWO × ×
131 Cyperus conglomeratus Rottb. Cyperaceae Burkina, Central African Republic, Chad, Eritrea, Ethiopia, Gabon, and Kenya (SZ) GBIF and POWO × × ×
Gebel Elba (SZ) and Mediterranen coast in Egypt (ME) Boulos (1999-2005)
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
132 Cyperus laevigatus L. Cyperaceae Angola, Botswana, Burundi, and Cape Verde (SZ) GBIF and POWO × ×
Canary Islands (ME) GBIF and POWO
Gebel Elba (SZ) and Mediterranen coast in Egypt (ME) Boulos (1999-2005)
133 Daucus sahariensis Murb. Apiaceae
134 Delphinium bovei Decne. Ranunculaceae Eastern Mediterranean coastal region (ME) Täckholm (1974)
135 Desmostachya bipinnata (L.) Stapf. Poaceae Cambodia, Central African Republic, Chad, Eritrea, and Ethiopia (SZ) GBIF and POWO ×
136 Deverra tortuosa (Desf.) DC. Apiaceae Coastal area of Libya and Mediterranen coast in Egypt (ME) Boulos (1999-2005) and Flora of Libya ×
137 Deverra triradiata Hochst. ex Boiss. Apiaceae West of Iraq (IT) Abd El-Ghani et al. (2011)
138 Dianthus guessfeldtianus Muschl. Caryophyllaceae
139 Dianthus monadelphus subsp. judaicus (Boiss.) Greuter & Burdet Caryophyllaceae
140 Dianthus sinaicus Boiss. Caryophyllaceae
141 Diplotaxis acris (Forssk.) Boiss. Brassicaceae Turkey and Spain (ME) Boulos (1999-2005) and POWO × ×
Chad (SZ) POWO
142 Diplotaxis harra (Forssk.) Boiss. Brassicaceae Sicilia and Spain (ME) Boulos (1999-2005); GBIF; and POWO × ×
Somalia (SZ) Boulos (1999-2005); GBIF; and POWO
143 Ducrosia ismaelis Asch. Apiaceae
144 Echinops glaberrimus DC. Asteraceae
145 Echinops taeckholmianus Amin Asteraceae Baltim and Rosetta (ME) CAI (1987) and El-Khalafy et al. (2021a, b, 2004) × ×
146 Echiochilon fruticosum Desf. Boraginaceae Mediterranen coast in Egypt (ME) and east of Lebanon (IT) Boulos (1999-2005) and Hamed et al. (2021) × × ×
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
147 Echium angustifolium Mill. Boraginaceae Cyprus, Greece, Kriti, and Turkey (ME) GBIF and POWO × ×
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
148 Echium horridum Batt. Boraginaceae Canary Islands (ME) GBIF and POWO × ×
Mali and Sudan (SZ) GBIF and POWO
Gebel Elba (SZ) Boulos (1999-2005)
149 Echium longifolium Delile Boraginaceae Ethiopia and Sudan (SZ) CAI (1973) and GBIF ×
Gebel Elba (SZ) and Mediterranen coast in Egypt (ME) Täckholm (1974)
150 Echium rauwolfii Delile Boraginaceae Sudan (SZ) POWO × ×
Cyprus (ME) POWO
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
151 Echium rubrum Forssk. Boraginaceae Russia and Spain (ME) GBIF × ×
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
152 Enarthrocarpus strangulatus Boiss. Brassicaceae The western Mediterranean coastal land in Egypt (ME) Boulos (1999-2005) and Turki and El Shayeb (2005) ×
153 Ephedra alata Decne. Ephedraceae Chad and Mali (SZ) GBIF and POWO × × ×
Mediterranen coast in Egypt (ME) Boulos (1999-2005) and Abd El-Ghani et al. (2017)
154 Ephedra aphylla Forssk. Ephedraceae Cyprus (ME) POWO ×
Mediterranen coast in Egypt and Rosetta (ME) Boulos (1999-2005) and Abdel-Kader et al. (2003)
155 Ephedra pachyclada subsp. sinaica (Riedl) Freitag & Maier-St. Ephedraceae
156 Eragrostis aegyptiaca (Willd.) Delile Poaceae Burkina, Chad, Mali, Niger, Senegal, and Sudan (SZ) GBIF and POWO × ×
Gebel Elba (SZ) and Mediterranen coast in Egypt (ME) Boulos (1999-2005)
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
157 Eremobium aegyptiacum subsp. aegyptiacum (Spreng.) Asch. ex Boiss. Brassicaceae Chad (SZ) Khalik (2005) × ×
Arish, Burg El Arab, and Mediterranen coast in Egypt (ME) Boulos (1999-2005); CAI (2004); and TANE (2010)
158 Eremobium aegyptiacum var. lineare (Delile) Zohary Brassicaceae
159 Eremogone sinaica (Boiss.) Dillenb. & Kadereit Caryophyllaceae
160 Erigeron bonariensis L. Asteraceae South of America and Western Europe (ES) Boulos (1999-2005) × × ×
161 Erodium arborescens (Desf.) Willd. Geraniaceae Mediterranen coast in Egypt (ME) Mahklouf (2020)
162 Erodium crassifolium L'Hér. Geraniaceae Cyprus and Kriti (ME) Boulos (1999-2005); GBIF; and POWO ×
163 Erodium glaucophyllum (L.) L'Hér. Geraniaceae Italy (ME) POWO × ×
Sudan (SZ) POWO
Mediterranen coast in Egypt (ME) Täckholm (1974) and Boulos (1999-2005)
Northwestern of Jordan (IT) Flora of Libya
164 Erodium oxyrhinchum M.Bieb. Geraniaceae Turkey, Turkmenistan, Uzbekistan, and Xinjiang Uygur Autonomous region, China (IT) GBIF and POWO × ×
Southwest of Saudi Arabia (SZ) Danin and Fragman-Sapir (2016)
165 Erodium pulverulentum (Boiss.) Batt. Geraniaceae Eritrea (SZ) POWO × ×
Spain (ME) POWO
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
North of Iran (IT) Abd El-Ghani et al. (2017)
166 Erodium touchyanum Delile ex Godr. Geraniaceae Canary Islands and Cyprus (ME) POWO
167 Erucaria pinnata (Viv.) Täckh. & Boulos Brassicaceae Burg Elarab, Matrouh, and Mediterranen coast in Egypt (ME) CAI (1979); Boulos (1999-2005); and TANE (2006) ×
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
168 Euphorbia granulata Forssk Euphorbiaceae Chad, Eritrea, Ethiopia, Kazakhstan, and Kenya (SZ) GBIF and POWO × ×
Gebel Elba (SZ) Boulos (1999-2005)
169 Euphorbia grossheimii (Prokh.) Prokh. Euphorbiaceae Transcaucasus (ME) Boulos (1999-2005); GBIF; and POWO ×
170 Euphorbia obovata Decne. Euphorbiaceae
171 Euphorbia retusa Forssk. Euphorbiaceae France (ME) GBIF ×
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
172 Farsetia aegyptia Turra Brassicaceae Chad, Mali, and Sudan (SZ) Danin and Fragman-Sapir (2016) and POWO × × ×
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
173 Farsetia stylosa R. Br. Brassicaceae Cameroon, Chad, Djibouti, Eritrea, Ethiopia, Kenya, and Mali (SZ) Boulos (1999-2005); Abd El-Ghani et al. (2017); GBIF; and POWO × × ×
174 Ferula sinaica Boiss. Apiacaea Mediterranen coast in Egypt (ME) Täckholm (1974)
175 Ficus palmata Forssk. Moraceae Eritrea, Ethiopia, Somalia, and Sudan (SZ) Boulos (1999-2005); GBIF; and POWO × ×
176 Galium aparine L. Rubiaceae Belgium, Bulgaria, Cyprus, Denmark, Finland, France, Gebel Elbarmany, Greece, and Ireland (ME) Boulos (1999-2005); GBIF; and POWO × ×
177 Galium ceratopodum Boiss. Rubiaceae Tadzhikistan, Turkmenistan, Uzbekistan, Russia, and other places of Central Asia (IT) Boulos (1999-2005); GBIF; and POWO × ×
178 Galium sinaicum (Delile ex Decne) Boiss. Rubiaceae
179 Galium tricornutum Dandy Rubiaceae Albania, Bulgaria, Cyprus, France, Gebel Elbarmany, Greece, and Italy (ME) GBIF and POWO × ×
Central Asia (IT) and Mediterranen coast in Egypt (ME) Boulos (1999-2005)
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
180 Glaucium arabicum Fresen. Papaveraceae Mediterranen coast in Egypt (ME) Boulos (1999-2005) ×
181 Glinus runkewitzii Tackh. & Boulos Molluginaceae
182 Gymnocarpos decander Forssk. Caryophyllaceae Canary Islands (ME) POWO × ×
Mediterranen coast in Egypt (ME) Boulos (1999-2005) and Abd El-Ghani et al. (2017)
183 Gymnocarpos sclerocephalus (Decne.) Dahlgren & Thulin Caryophyllaceae Chad, Djibouti, and Eritrea (SZ) GBIF and POWO × ×
Gebel Elba (SZ) Boulos (1999-2005)
184 Gomphocarpus sinaicus Boiss. Apocynaceae
185 Gypsophila capillaris (Forssk.) C.Chr. Caryophyllaceae North of Syria (IT) Danin and Fragman-Sapir (2016) and GBIF ×
Mediterranen coast in Egypt (ME) Abd El-Ghani et al. (2017)
186 Halopeplis perfoliata (Forssk.) Bunge ex Ung.-Sternb. Amaranthaceae Eritrea and Sudan (SZ) POWO × ×
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
187 Haloxylon negevensis (Iljin & Zohary) L.Boulos Amaranthaceae
188 Haplophyllum poorei C.C.Towns. Rutaceae
189 Haplophyllum tuberculatum (Forssk.) A. Juss. Rutaceae Chad, Somalia, and Sudan (SZ) GBIF and POWO × ×
Gebel Elba (SZ) and Mediterranen coast in Egypt (ME) Boulos (1999-2005)
190 Helianthemum kahiricum Delile Cistaceae Cyprus (ME) POWO × ×
Mediterranen coast in Egypt (ME) Boulos (1999-2005) and Makhlouf (2020)
191 Helianthemum lippii (L.) Dum.Cours. Cistaceae Italy and Kriti (ME) GBIF and POWO × × ×
Chad and Sudan (SZ) GBIF and POWO
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
192 Helianthemum sancti-antoni Schweinf. ex Boiss. Cistaceae
193 Helianthemum schweinfurthii Grosser Cistaceae
194 Helianthemum stipulatum (Forssk.) C. Chr. Cistaceae Djibouti and Somalia (SZ) GBIF and POWO × ×
Cyprus, Greece, and Turkey (ME) GBIF and POWO
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
195 Helianthemum ventosum Boiss. Cistaceae
196 Heliotropium arbainense Fresen. Boraginaceae Ethiopia and Sudan (SZ) Boulos (1999-2005); GBIF; and POWO × ×
197 Heliotropium bacciferum Forssk. Boraginaceae Chad, Comoros, Djibouti, Eritrea, and Ethiopia (SZ) GBIF and POWO × × ×
Gebel Elba (SZ) Boulos (1999-2005)
198 Heliotropium digynum (Forssk.) Asch. ex C.Chr. Boraginaceae The western Mediterranean coastal land (ME) Turki and El Shayeb (2005) × ×
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
West of Iraq (IT) Abd El-Ghani et al. (2017)
199 Herniaria hemistemon J. Gay Caryophyllaceae Mediterranen coast in Egypt and Sallum (ME) Boulos (1999-2005) and Amer at al. (2020) × ×
North of Syria (IT) Makhlouf (2020) and GBIF (2023)
200 Heteroderis pusilla (Boiss.) Boiss. Asteraceae Kazakhstan, Kirgizstan, Tadzhikistan, Turkmenistan, and Uzbekistan (IT) GBIF and POWO ×
201 Hippocrepis areolata Desv. Fabaceae France and Mediterranen coast (ME) Boulos (1999-2005) and POWO ×
202 Hippocrepis constricta Kunze Fabaceae Chad, Sudan, and Tanzania (SZ) GBIF and POWO × ×
203 Hyoscyamus boveanus (Dunal) Asch. & Schweinf. Solanaceae
204 Hyoscyamus desertorum (Asch. ex Boiss.) Täckh. Solanaceae Arish and Mediterranen coast in Egypt (ME) Boulos (1999-2005) and GBIF × ×
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
205 Hyoscyamus pusillus L. Solanaceae Central Asia, and south European Russia, Turkey, Turkmenistan, Uzbekistan, and Xinjiang Uygur Autonomous Region, China (IT) Boulos (1999-2005); GBIF; and POWO × ×
206 Hypecoum aegyptiacum (Forssk.) Asch. & Schweinf. Papaveraceae El Kawm El Akhdar Island and Mediterranen coast in Egypt (ME) Boulos (1999-2005) and TANE (2000) ×
207 Hypecoum aequilobum Viv. Papaveraceae Mediterranen coast in Egypt (ME) Boulos (1999-2005) ×
208 Hypericum sinaicum Hochst. ex Boiss. Hypericaceae
209 Ifloga spicata subsp. hadidii (Fayed & Zareh) Greuter Asteraceae Arish and Mediterranen coast in Egypt (ME) Boulos (1999-2005) and GBIF ×
210 Iphiona mucronata (Forssk.) Asch. & Schweinf. Asteraceae
211 Iphiona scabra DC. ex Decne. Asteraceae Djibouti, Eritrea, Ethiopia, and Sudan (SZ) Boulos (1999-2005); GBIF; and POWO × ×
Gebel Elba (SZ) Boulos (1999-2005)
212 Iris mariae Barbey Iridaceae Bardawil Lake and Rafah (ME) El Banna (2009) and GBIF ×
213 Isatis microcarpa J. Gay ex Boiss. Brassicaceae Mediterranen coast in Egypt (ME) Boulos (1999-2005) ×
214 Kickxia aegyptiaca subsp. aegyptiaca (L.) Nábělek Plantaginaceae Chad and Sudan (SZ) POWO × × ×
Mediterranen coast in Egypt (ME) Boulos (1999-2005) and Danin and Fragman-Sapir (2016)
215 Kickxia floribunda (Boiss.) Täckh. & Boulos Plantaginaceae Arish (ME) Täckholm (1974); El-Hadidi et al. (1999); and GBIF
216 Lasiurus scindicus Henrard Poaceae Chad, Djibouti, Eritrea, Ethiopia, Mali, Niger, Somalia, and Sudan (SZ) GBIF and POWO × ×
Gebel Elba (SZ) and Mediterranen coast in Egypt (ME) Boulos (1999-2005)
217 Lathyrus hirsutus L. Fabaceae France, Greece, Poland, and Portugal (ME) Boulos (1999-2005); GBIF; and POWO × ×
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
218 Lathyrus sphaericus Retz. Fabaceae France, Greece, and Hungary (ME) Boulos (1999-2005); GBIF; and POWO × ×
Eritrea and Ethiopia (SZ) Boulos (1999-2005); GBIF; and POWO
219 Launaea amal-aminiae N. Kilian Asteraceae Gebel Elba and Halayeb Triangle (SZ) Zareh et al. (2016) and Marzouk et al. (2021)
220 Launaea angustifolia (Desf.) Kuntze Asteraceae Mediterranen coast in Egypt (ME) Täckholm (1974)
221 Launaea capitata (Spreng.) Dandy Asteraceae Chad, Eritrea, and Sudan (SZ) Boulos (1999-2005); GBIF; and POWO × ×
Canary Islands (ME) Boulos (1999-2005); GBIF; and POWO
222 Launaea mucronata (Forssk.) Muschl Asteraceae Chad, Eritrea, and Sudan (SZ) Boulos (1999-2005); GBIF; and POWO × ×
223 Launaea mucronata subsp. cassiniana (Jaub. & Spach) N.Kilian Asteraceae Chad and Sudan (SZ) Boulos (1999-2005); GBIF; and POWO × ×
224 Launaea spinosa (Forssk.) Sch. Bip. Ex Kuntze Asteraceae
225 Lavandula atriplicifolia Benth. Lamiaceae
226 Lavandula coronopifolia Poir. Lamiaceae Cape Verde, Chad, and Djibouti (SZ) Boulos (1999-2005); GBIF; and POWO × ×
227 Lepidium niloticum (Delile) Sieber ex Steud. Brassicaceae Chad and Sudan (SZ) GBIF and POWO ×
228 Leptadenia arborea (Forssk.) Schweinf Apocynaceae Burkina, Cameroon, Chad, and Djibouti (SZ) Boulos (1999-2005); GBIF; and POWO × ×
229 Leptadenia pyrotechnica (Forssk.) Decne Apocynaceae Burkina, Cameroon, Chad, Djibouti, and Eritrea (SZ) Boulos (1999-2005); GBIF; and POWO × ×
230 Leobordea platycarpa (Viv.) B. E. van Wyk & Boatwr. Fabaceae Angola, Botswana, Chad, and Central African Republic (SZ) GBIF and POWO × ×
231 Limonium axillare (Forssk.) Kuntze Plumbaginaceae Djibouti, Eritrea, Somalia, and Sudan (SZ) Boulos (1999-2005); GBIF; and POWO × ×
232 Limonium pruinosum (L.) Chaz. Plumbaginaceae El Alamein, north coast of the Nile Delta in Egypt, and Mediterranen coast in Egypt (ME) Boulos (1999-2005); El-Amier et al. (2014); and GBIF × ×
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
233 Lindenbergia indica (L.) Vatke Orobanchaceae Djibouti, Eritrea, and Ethiopia (SZ) Boulos (1999-2005); GBIF; and POWO × ×
234 Lobularia arabica (Boiss.) Muschl. Brassicaceae Greece (ME) El-Amier (2016) and Flora of Greece × ×
Deltaic Mediterranean coast and Mediterranen coast in Egypt (ME) Boulos (1999-2005)
235 Lotus hebranicus Hochst. ex Brand Fabaceae Sudan (SZ) Sokoloff (2001) and POWO ×
236 Lotus glinoides Delile Fabaceae Chad, Eritrea, and Sudan (SZ) Boulos (1999-2005); GBIF; and POWO × ×
237 Lotus lanuginosus Vent. Fabaceae
238 Lupinus digitatus Forssk. Fabaceae Senegal (SZ) GBIF and POWO ×
239 Lycium shawii Roem. & Schult. Solanaceae Botswana, Djibouti, Eritrea, and Ethiopia (SZ) GBIF and POWO × ×
Gebel Elba (SZ) Boulos (1999-2005)
240 Lygeum spartum Loefl. ex L. Poaceae Italy, Kriti, Mediterranen coast in Libya, and Spain (ME) Boulos (1999-2005); GBIF; Flora of Libya); and POWO × × ×
241 Malcolmia pygmaea (DC.) Boiss. Brassicaceae Ale×andria, El Arish, and Mediterranen coast in Egypt (ME) CAI (1992); Boulos (1999-2005); and GBIF × ×
242 Malva ludwigii (L.) Soldano, Banfi & Galasso Malvaceae Mediteranrn coast (ME) Asri (2004) × × ×
Tadzhikistan, Turkmenistan, and Uzbekistan (IT) POWO
243 Matthiola arabica Boiss. Brassicaceae
244 Matthiola longipetala subsp. hirta (Conti) Greuter & Burdet Brassicaceae Mariut (ME) GBIF × ×
Turkey (IT) POWO
Mediterranen coast in Libya (ME) Flora of Libya
245 Matthiola longipetala subsp. livida (Delile) Maire Brassicaceae Canary Islands (ME) POWO × ×
Burg el arab and Mediterranen coast in Egypt (ME) Boulos (1999-2005) and Marzouk et al. (2010)
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
246 Matthiola longipetala subsp. longipetala (Vent.) DC. Brassicaceae Cyprus, Turkey, and Uzbekistan (IT) GBIF and POWO × ×
247 Medicago laciniata (L.) Mill. Fabaceae Botswana, Djibouti, and Ethiopia (SZ) POWO × ×
Canary Islands (ME) POWO
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
248 Melilotus serratifolius Tackh. & Boulos Fabaceae
249 Mesembryanthemum cryptanthum Hook.f. Aizoaceae Angola, Cape Verde, and Namibia (SZ) Danin and Fragman-Sapir (2016) and POWO × × ×
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
250 Micromeria serbaliana Danin & Hedge Lamiaceae
251 Micromeria sinaica Benth. Lamiaceae
252 Microparacaryum intermedium (Fresen.) Hilger & Podlech Boraginaceae Kazakhstan, Tadzhikistan, Uzbekistan, and West Himalaya (IT) GBIF and POWO × ×
Eritrea (SZ) GBIF and POWO
253 Monsonia heliotropioides (Cav.) Boiss. Geraniaceae Chad, NiGebel Elbar, and Sudan (SZ) Boulos (1999-2005); GBIF; and POWO × ×
254 Monsonia nivea (Decne.) Webb Geraniaceae Chad, NiGebel Elbar, and Sudan (SZ) Boulos (1999-2005); Danin and Fragman-Sapir (2016); GBIF; and POWO × ×
255 Morettia canescens Boiss. Brassicaceae Chad, Mali, and Niger (SZ) GBIF and POWO ×
256 Morettia parviflora Boiss. Brassicaceae South of Yemen (SZ) GBIF
257 Morettia philaeana (Delile) DC. Brassicaceae Chad, Mali, NiGebel Elbar, and Sudan (SZ) Boulos (1999-2005); GBIF; and POWO × × ×
258 Moricandia nitens (Viv.) E. A. Durand & Barratte Brassicaceae Alamein, Burg El arab, and Mediterranen coast in Egypt (ME) Boulos (1999-2005); Marzouk et al. (2010); and GBIF × × ×
259 Moricandia sinaica (Boiss.) Boiss. Brassicaceae Somalia (SZ) Abdullah and Al-Dosari (2022); GBIF; and POWO
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
260 Muscari eburneum (Eig & Feinbrun) D.C.Stuart Lamiaceae Mediterranen coast in Egypt (ME) Täckholm (1974)
261 Muscari longipes subsp. negevense (Feinbrun & Danin) Hosni Lamiaceae
262 Najas pectinata (Parl.) Magnus Hydrocharitaceae Chad, Guinea, Mali, and Senegal (SZ) GBIF and POWO ×
263 Nanorrhinum acerbianum (Boiss.) Betsche Plantaginaceae Gebel Elba (SZ) Boulos (1999-2005) ×
264 Nanorrhinum macilentum (Decne.) Betsche Plantaginaceae Chad and Sudan (SZ) Brundu and Camarda (2013) and POWO
265 Nasturtiopsis integrifolia (Boulos) Abdel Khalik & F.T.Bakker Brassicaceae El Arish (ME) GBIF
266 Nepeta septemcrenata Ehrenb. ex Benth. Lamiaceae
267 Neurada procumbens L. Neuradaceae Chad, Eritrea, Mali, Niger, and Sudan (SZ) GBIF and POWO × × ×
Cyprus (ME) GBIF and POWO
Gebel Elba (SZ) and Mediterranen coast in Egypt (ME) Boulos (1999-2005)
268 Nidorella aegyptiaca (L.) J. C. Manning & Goldblatt Asteraceae Benin, Botswana, Burundi, Cambodia, Cameroon, Chad, Congo, Eritrea, Ethiopia, Ghana, and Guinea (SZ) Boulos (1999-2005); GBIF; and POWO × ×
269 Nitraria retusa (Forssk.) Asch. Nitrariaceae Senegal and Sudan (SZ) GBIF and POWO × × ×
Gebel Elba (SZ) and Mediterranen coast in Egypt (ME) Boulos (1999-2005)
270 Ochradenus baccatus Delile Resedaceae Chad, Djibouti, Eritrea, and Ethiopia (SZ) GBIF and POWO × × ×
Gebel Elba (SZ) and Mediterranen coast in Egypt (ME) Boulos (1999-2005)
271 Ogastemma pusillum (Coss. & Durieu ex Bonnet & Barratte) Brummitt Boraginaceae Canary Islands (ME) POWO
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
272 Oligomeris linifolia (Vahl ex Hornem.) J. F. Macbr. Resedaceae Canary Islands and Kriti (ME) Boulos (1999-2005); GBIF; and POWO × ×
Chad (SZ) Boulos (1999-2005); GBIF; and POWO
China (IT) Boulos (1999-2005); GBIF; and POWO
273 Onobrychis ptolemaica (Delile) DC. Fabaceae Turkey (IT) GBIF and TÜBİVES (2023)
274 Onopordum ambiguum Fresen. Asteraceae Centre of Jordan (IT) Rabei et al. (2021) ×
275 Origanum isthmicum Danin Lamiaceae
276 Origanum syriacum subsp. sinaicum (Boiss.) Greuter & Burdet Lamiaceae
277 Pallenis hierichuntica (Michon) Greuter Asteraceae Mediterranen coast in Egypt (ME) Boulos (1999-2005) × × ×
Canary Islands (ME) Boulos (1999-2005) and POWO
East Iran (IT) Flora of Libya
278 Pancratium sickenbergeri Asch. & Schweinf. Amaryllidaceae Bardawil Lake, El Borollos, and Omayed (ME), and Elba Mountain (SZ) TANE (2000) and El-Bana (2009) ×
279 Panicum turgidum Forssk. Poaceae Benin, Burkina, Chad, Congo, Djibouti, Eritrea, and Ethiopia (SZ) Danin and Fragman-Sapir (2016); GBIF; and POWO × × ×
280 Papaver decaisnei (Elkan) Hochst. & Steud. ex Boiss. Papaveraceae Burg El-Arab and Mersa Matruh (ME) Taia (2009) ×
Russia (IT) GBIF
281 Paronychia arabica subsp. arabica (L.) DC. Caryophyllaceae Chad (SZ) POWO × ×
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
282 Paronychia sinaica Fresen. Caryophyllaceae
283 Pergularia tomentosa L. Apocynaceae Benin, Burkina, Chad, Djibouti, Ethiopia, Mali, and Niger (SZ) GBIF and POWO × × ×
Gebel Elba (SZ) Boulos (1999-2005)
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
284 Periploca aphylla subsp. aphylla Decne. Apocynaceae South of Yemen and South of Oman (SZ) Danin and Fragman-Sapir (2016) and GBIF ×
285 Persicaria obtusifolia (Täckh. & Boulos) Greuter & Burdet Polygonaceae
286 Petrorhagia arabica (Boiss.) P. W. Ball & Heywood Caryophyllaceae
287 Phagnalon barbeyanum Asch. & Schweinf. Asteraceae Matrouh and Mediterranen coast in Egypt (ME) Täckholm (1974); Sabri et al. (1992); and Boulos (1999-2005) × ×
288 Phagnalon nitidum Fresen. Asteraceae Ethiopia (SZ) GBIF
289 Phagnalon sinaicum Bornm. & Kneuck. Asteraceae
290 Phlomis aurea Decne. Lamiaceae
291 Phoenix dactylifera L. Arecaceae Canary Islands (ME) Boulos (1999-2005); GBIF; and POWO × × ×
Benin, Cape Verde, and Chad (SZ) Boulos (1999-2005); GBIF; and POWO
China (IT) Boulos (1999-2005); GBIF; and POWO
292 Picris cyanocarpa Boiss. Asteraceae Mediterranen coast in Egypt (ME) Täckholm (1974)
293 Picris sulphurea Delile Asteraceae
294 Pimpinella schweinfurthii Asch. Apiaceae
295 Pistacia khinjuk Stocks Anacardiaceae
296 Plantago amplexicaulis subsp. bauphula (Edgew.) Rech.f. Plantaginaceae Spain (ME) Boulos (1999-2005) and POWO × ×
297 Plantago cylindrica Forssk. Plantaginaceae El Arish, Mediterranen coast in Egypt, and the way between Burg El Arab and El Alamin (ME) CAI (1964) and Boulos (1999-2005) × ×
298 Plantago sinaica (Barneoud) Decne. Plantaginaceae
299 Plicosepalus acaciae (Zucc.) Wiens & Polhill Loranthaceae Chad, Eritrea, and Ethiopia (SZ) Boulos (1999-2005); GBIF; and POWO × ×
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
300 Plicosepalus curviflorus (Benth. exoliv.) Tiegh Loranthaceae Djibouti, Eritrea, Ethiopia, Kenya, Somalia, Sudan, Tanzania, and Uganda (SZ) GBIF and POWO × ×
Gebel Elba (SZ) Boulos (1999-2005)
301 Pluchea dioscoridis (L.) DC. Asteraceae Angola, Botswana, Burundi, Chad, Djibouti, Eritrea, Ethiopia, Kenya, and Sudan (SZ) GBIF and POWO × × ×
Lebanon (ME) Boulos (1999-2005)
302 Podonosma galalensis Schweinf. ex Boiss. Boraginaceae
303 Polycarpaea repens (Forssk.) Asch. & Schweinf. Caryophyllaceae Chad, Niger, and Sudan (SZ) Boulos (1999-2005) and POWO × × ×
Gebel Elba (SZ) Boulos (1999-2005)
304 Polycarpaea robbairea (Kuntze) Greuter & Burdet Caryophyllaceae Chad, Djibouti, Eritrea, Mali, Niger, Somalia, and Sudan (SZ) GBIF and POWO × ×
Gebel Elba (SZ) Boulos (1999-2005)
305 Polycarpon succulentum J.Gay Caryophyllaceae Bardawil Lake and Sallum (ME) El Bana (2009) and Amer et al. (2020) × ×
Lebanon (ME) Boulos (1999-2005)
306 Polygala sinaica var. glabrescens (Zohary) Boulos Polygalaceae
307 Polygala sinaica var. sinaica Botsch. Polygalaceae
308 Evotrochis involucrata Raf. Primulaceae
309 Pseuderucaria clavata (Boiss. & Reut.) O.E. Schulz Brassicaceae Mediterranen coast in Egypt (ME) Boulos (1999-2005) ×
310 Pseuderucaria teretifolia (Desf.) O. E. Schulz Brassicaceae
311 Pteranthus dichotomus Forssk. Caryophyllaceae Canary Islands, Cyprus, and Sicilia (ME) Boulos (1999-2005); GBIF; and POWO × × ×
312 Pterocephalus arabicus Boiss. Caprifoliaceae
313 Pterocephalus sanctus Decne. Caprifoliaceae
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
314 Pulicaria incisa (Lam.) DC. Asteraceae Benin, Burkina, Chad, Djibouti, Eritrea, and Ethiopia (SZ) GBIF and POWO × × ×
Gebel Elba (SZ) Boulos (1999-2005)
315 Pulicaria sicula (L.) Moris Asteraceae Albania, Cyprus, France, Greece, Italy, and Sicilia (ME) GBIF and POWO ×
316 Pulicaria undulata (L.) C. A. Mey. Asteraceae Benin, Burkina, Cameroon, Cape Verde, Chad, Djibouti, Eritrea, and Ethiopia (SZ) Boulos (1999-2005); GBIF; and POWO × ×
317 Pycnocycla tomentosa Decne. Apiaceae
318 Reaumuria hirtella Jaub. & Spach Tamariaceae Mediterranen coast in Egypt and the northern coastal region of Egypt (ME) Boulos (1999-2005) and Abozeed (2014) × × ×
319 Reaumuria hirtella var. palaestina (Boiss.) Zohary & Danin Tamariaceae North of Syria (ME) GBIF
320 Reaumuria negevensis Zohary & Danin Tamariaceae
321 Reseda alba L. Resedaceae Albania, Cyprus, France, and Greece (ME) Boulos (1999-2005); GBIF; and POWO × ×
Eritrea (SZ) Boulos (1999-2005); GBIF; and POWO
322 Reseda arabica Boiss. Resedaceae Burg El Arab, El Arish, and Mediterranen coast in Egypt (ME) Boulos (1999-2005); CAI (2012); and El-Ahmir et al. (2020) × ×
323 Reseda decursiva Forssk. Resedaceae Chile and Spain (ME) GBIF and POWO × ×
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
324 Reseda muricata C. Presl Resedaceae
325 Reseda pruinosa Delile Resedaceae Ethiopia and Sudan (SZ) GBIF and POWO × ×
Gebel Elba (SZ) and Mediterranen coast in Egypt (ME) Boulos (1999-2005)
326 Reseda stenostachya Boiss. Resedaceae
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
327 Retama raetam (Forssk.) Webb & Berthel. Fabaceae Greece and Sicilia (ME) GBIF and POWO × ×
Sudan (SZ) GBIF and POWO
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
328 Rhamnus disperma Ehrenb. ex Boiss. Rhamnaceae ×
329 Rhamphospermum arvense (L.) Besser Brassicaceae lbania, Altay, Cyprus, Denmark, France, Gebel Elbarmany, and Greece (ME) GBIF and POWO × ×
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
330 Rhazya greissii Tackh. & Boulos Apocynaceae
331 Rhazya stricta Decne. Apocynaceae South of Yemen (SZ) GBIF
332 Rhynchosia minima (L.) DC. Fabaceae Angola, Benin, and Botswana (SZ) Boulos (1999-2005); GBIF; and POWO × ×
333 Robeschia schimperi (Boiss.) O. E. Schulz Brassicaceae Central Iran and Syria (IT) GBIF ×
334 Rosa arabica (Crép. ex Boiss.) Déségl. Rosaceae
335 Rostraria rohlfsii (Asch.) Holub Poaceae Chad and Niger (SZ) GBIF and POWO ×
336 Rumex aegyptiacus L. Polygonaceae France (ME) GBIF ×
Burg El Arab and Mediterranen coast in Egypt (ME) El-Khalafy et al. (2021a, b) and Balah and Hozyan (2022)
337 Rumex cyprius Murb Polygonaceae Cyprus (ME) Boulos (1999-2005); GBIF; and POWO × × ×
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
338 Rumex dentatus L. Polygonaceae Albania, Bulgaria, Cyprus, and Greece (ME) GBIF and POWO × × ×
China (IT) GBIF and POWO
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
339 Rumex pictus Forssk. Polygonaceae Mediterranen coast in Egypt and Rosetta (ME) Danin and Fragman-Sapir (2016); Abd El-Aal et al. (2019); and GBIF ×
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
340 Rumex simpliciflorus Murb. Polygonaceae Chad and Sudan (SZ) GBIF and POWO × ×
Gebel Elba (SZ) Boulos (1999-2005)
341 Salvadora persica L. Salvadoraceae Angola, Botswana, Burkina, Chad, Djibouti, Eritrea, Ethiopia, Kenya, Malawi, Mali, Mozambique, Namibia, and Niger (SZ) GBIF and POWO × ×
Gebel Elba (SZ) Boulos (1999-2005) and Abd El-Ghani et al. (2017)
342 Salvia aegyptiaca L. Lamiaceae Chad, Eritrea, Mali, and Niger (SZ) Boulos (1999-2005); GBIF; and POWO × × ×
Canary Islands (ME) Boulos (1999-2005); GBIF; and POWO
Gebel Elba (SZ) and Mediterranen coast in Egypt (ME) Boulos (1999-2005) and Abd El-Ghani et al. (2017)
343 Salvia deserti Decne. Lamiaceae
344 Savignya parviflora (Delile) Webb Brassicaceae El Dabaa (ME) CAI (2021)
East of Iran (IT) Osman et al. (2014) and Flora of Libya
345 Scabiosa eremophila Boiss. Caprifoliaceae Mediterranen coast in Egypt and Omayed (ME) Boulos (1999-2005) and Ahmed et al. (2015) ×
346 Schimpera arabica Hochst. & Steud. Brassicaceae
347 Schouwia purpurea (Forssk.) Schweinf. Brassicaceae Burkina, Chad, Djibouti, Ethiopia, Mali, and Niger (SZ) Alyasi (2020); GBIF; and POWO (2023 × ×
Gebel Elba (SZ) and Mediterranen coast in Egypt (ME) Boulos (1999-2005)
348 Scorzonera drarii V.Tackh. Asteraceae
349 Scorzonera schweinfurthii Boiss. Asteraceae
350 Scrophularia deserti Delile Scrophulariaceae
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
351 Scrophularia hypericifolia Wydler Scrophulariaceae North of Iran (IT) Feyzi et al. (2014); Veiskarami and Sharifi-Tehrani (2017); and GBIF ×
352 Senecio aegyptius L. Asteraceae Ethiopia and Sudan (SZ) Boulos (1999-2005); GBIF; and POWO × ×
Cyprus (ME) Boulos (1999-2005); GBIF; and POWO
353 Senecio belbeysius Delile Asteraceae
354 Senecio flavus (Decne.) Sch. Bip Asteraceae Canary Islands and Spain (ME) Boulos (1999-2005); GBIF; and POWO × ×
Chad, Namibia, and Sudan (SZ) Boulos (1999-2005); GBIF; and POWO
Gebel Elba (SZ) Boulos (1999-2005)
355 Senna holosericea (Freseu) Greuter Fabaceae Chad, Djibouti, Eritrea, Ethiopia, Somalia, and Sudan (SZ) Boulos (1999-2005); GBIF; and POWO × ×
Gebel Elba (SZ) Boulos (1999-2005)
356 Senna italica Mill. Fabaceae Benin (SZ) Boulos (1999-2005); GBIF; and POWO × × ×
Colombia and Jamaica (ES) Boulos (1999-2005); GBIF; and POWO
Gebel Elba (SZ) Boulos (1999-2005) and Abd El-Ghani et al. (2017)
357 Setaria verticillata (L.) P.Beauv. Poaceae Angola, Bangladesh, Benin, Botswana, Bulgaria, Burkina, Burundi, Cambodia, and Cameroon (SZ) Boulos (1999-2005); GBIF; and POWO × × ×
358 Silene arabica Boiss. Caryophyllaceae ×
359 Silene hussonii Boiss. Caryophyllaceae
360 Silene leucophylla Boiss. Caryophyllaceae
361 Silene linearis Decne. Caryophyllaceae
362 Silene odontopetala var. congesta Boiss. Caryophyllaceae
363 Silene oreosinaica Chowdhuri Caryophyllaceae
364 Silene schimperiana Boiss. Caryophyllaceae
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
365 Silene villosa var. graveolens Sickenb. Caryophyllaceae
366 Silene vivianii subsp. viscida (Boiss.) Boulos Caryophyllaceae
367 Silene vivianii subsp. vivianii Steud. Caryophyllaceae Mediterranen coast in Egypt (ME) Boulos (1999-2005) × ×
368 Soda schweinfurthii (Solms) Akhani Amaranthaceae Mediterranen coast in Egypt (ME) Boulos (1999-2005) ×
369 Solenostemma oleifolium (Nectoux) Bullock & E.A. Bruce Apocynaceae Chad, Mali, Niger, and Sudan (SZ) GBIF and POWO × ×
370 Sporobolus spicatus (Vahl) Kunth Poaceae Angola, Botswana, Burundi, Chad, Djibouti, Eritrea, and Ethiopia (SZ) Boulos (1999-2005); GBIF; and POWO × × ×
Mediterranen coast in Egypt (ME) and SZ Abd El-Ghani et al. (2017)
371 Stachys aegyptiaca Pers. Lamiaceae
372 Stipagrostis drarii (Tackh.) de Winter Poaceae
373 Stipagrostis multinerva H. Scholz Poaceae East of Iran (IT) GBIF
374 Stipagrostis plumosa (L.) Munro ex T.Anderson Poaceae Chad, Cyprus, Mali, and Niger (SZ) GBIF and POWO × × ×
Gebel Elba (SZ) and Mediterranen coast in Egypt (ME) Boulos (1999-2005)
375 Stipagrostis vulnerans (Trin. & Rupr.) De Winter Poaceae Chad, Mali, Niger, and Sudan (SZ) Boulos (1999-2005); GBIF; and POWO × ×
376 Suaeda aegyptiaca (Hasselq.) Zohary Amaranthaceae Djibouti and Eritrea (SZ) Boulos (1999-2005); GBIF; and POWO × ×
Cyprus (ME) Boulos (1999-2005); GBIF; and POWO
377 Amaranthaceae Italy (ME) GBIF × ×
Suaeda fruticosa Forssk. ex J. F. Gmel. Somalia (SZ) GBIF
Mexico (ES) GBIF
378 Suaeda monoica Forssk. ex J. Gmelin Amaranthaceae Botswana, Chad, Djibouti, Eritrea, and Ethiopia (SZ) Boulos (1999-2005); GBIF; and POWO × × ×
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
379 Suaeda vermiculata Forssk. ex J.F.Gme Amaranthaceae Chad, Djibouti, and Eritrea (SZ) GBIF and POWO × × ×
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
380 Tamarix amplexicaulis Ehrenb. Tamaricaceae Chad, Mali, and Sudan (SZ) Boulos (1999-2005); GBIF; and POWO × ×
381 Tamarix aphylla (L.) H. Karst Tamaricaceae Chad, Djibouti, Eritrea, Ethiopia, and Niger (SZ) GBIF and POWO × ×
Gebel Elba (SZ) and Mediterranen coast in Egypt (ME) Boulos (1999-2005)
382 Tamarix nilotica (Ehrenb.) Bunge Tamaricaceae Chad, Djibouti, Eritrea, and Ethiopia (SZ) GBIF and POWO × × ×
Greece (ME) GBIF and POWO
Gebel Elba (SZ) and Mediterranen coast in Egypt (ME Boulos (1999-2005)
383 Tamarix passerinoides Delile ex Decne. Tamaricaceae Turkmenistan and Russia (IT) Boulos (1999-2005); GBIF; and POWO × ×
Sudan (SZ) GBIF and POWO
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
384 Tamarix tetragyna Ehrenb. Tamaricaceae Cyperus (ME) POWO × × ×
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
Russia (IT) Abd El-Ghani et al. (2017)
385 Tanacetum sinaicum (Fresen.) Delile ex K.Bremer & Humphries Asteraceae
386 Taverniera aegyptiaca Boiss. Fabaceae Eritrea and Sudan (SZ) GBIF and POWO ×
Mediterranen coast in Egypt (ME) Täckholm (1974)
387 Telephium sphaerospermum Boiss. Caryophyllaceae Somalia, South of Yemen (SZ) GBIF and POWO
388 Tephrosia kassasii Boulos Fabaceae
389 Tephrosia purpurea subsp. apollinea (Delile) Hosni & El-Karemy Fabaceae Chad, Djibouti, Eritrea, and Somalia (SZ) GBIF and POWO × ×
Gebel Elba (SZ) Boulos (1999-2005)
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
390 Tephrosia purpurea subsp. leptostachya (DC.) Brummitt Fabaceae Angola, Benin, Botswana, Burkina, Burundi, Cameroon, Tropical Africa (SZ) Boulos (1999-2005); GBIF; and POWO × ×
391 Teucrium decaisnei C. Presl Lamiaceae
392 Teucrium leucocladum Boiss. Lamiaceae
393 Teucrium polium L. Lamiaceae France, Morocco, and Spain (ME) GBIF and POWO × × ×
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
394 Thymelaea hirsuta (L.) Endl Thymelaeaceae Baleares, Cyprus, France, Greece, and Italy (ME) GBIF and POWO × × ×
Mediterranen coast in Egypt (ME) Abd El-Ghani et al. (2017)
395 Thymus bovei Benth. Lamiaceae
396 Thymus decussatus Benth. Lamiaceae
397 Traganum nudatum Delile Amaranthaceae Matrouh and Mediterranen coast in Egypt (ME) Boulos (1999-2005) and El-Bassossy (2022) ×
398 Tragopogon collinus DC. Asteraceae Armenia and Kazakhstan (IT) GBIF and POWO ×
399 Tribulus kaiseri Hosni Zygophyllaceae
400 Tribulus megistopterus Kralik Zygophyllaceae Chad and Sudan (SZ) GBIF and POWO × ×
Gebel Elba (SZ) Boulos (1999-2005)
401 Trifolium alexandrinum L Fabaceae Bulgaria, Greece, Hungary, and Kriti (ME) and Nepal and China (IT) GBIF and POWO × ×
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
402 Trigonella glabra Thunb. Fabaceae Djibouti and Sudan (SZ) GBIF and POWO × ×
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
403 Trigonella media Delile ex Urb. Fabaceae
404 Trigonella schlumbergeri Boiss. Fabaceae ×
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
405 Tripleurospermum auriculatum (Boiss.) Rech.f. Asteraceae West of Syria and west of Lebanon (IT) Boulos (1999-2005) and GBIF ×
406 Vachellia flava (Forssk.) Kyal. & Boatwr. Fabaceae Benin, Burkina, Cameroon, Central African Republic, and Chad (SZ) Boulos (1999-2005); GBIF; and POWO × × ×
407 Vachellia gerrardi var. najdensis (Chaudhary) Ragup., Seigler, Ebinger & Maslin Fabaceae South of Yemen and south of Oman (SZ) GBIF
408 Vachellia tortilis subsp. raddiana (Savi) Kyal. & Boatwr Fabaceae Burkina, Chad, Eritrea, Ethiopia, and Kenya (SZ) Boulos (1999-2005); GBIF; and POWO × × ×
409 Verbascum decaisneanum Kuntze Scrophulariaceae
410 Verbascum eremobium Murb. Scrophulariaceae Centre of Jordan (IT) Danin and Fragman-Sapir (2016)
411 Verbascum fruticulosum Post & Autran Scrophulariaceae
412 Verbascum schimperianum Boiss. Scrophulariaceae
413 Veronica anagalloides subsp. taeckholmiorum Chrtek & Osb.-Kos. Plantaginaceae El Beheira, El-Mahmudia, and Egypt (ME) CAI (1970)
414 Veronica kaiseri Tackh. Plantaginaceae
415 Vicia sinaica Boulos Fabaceae Arish (ME) El-Khalafy et al. (2021a, b, 2024) and Bedair et al. (2023) ×
416 Volutaria sinaica (DC.) Wagenitz Asteraceae Chad (SZ) GBIF and POWO ×
Gebel Elba (SZ) Boulos (1999-2005)
417 Zilla spinosa subsp. biparmata (O.E.Schulz) Maire & Weiller Brassicaceae Mediterranen coast in Egypt (ME) Greuter et al. (1984, 1986, 1989); Boulos (1999-2005); and Greuter and von Raab-Straube (2008) ×
418 Zilla spinosa subsp. spinosa (L.) Prantl Brassicaceae Gebel Elba (SZ) Boulos (1999-2005) ×
419 Ziziphus nummularia (Burm.f.) Wight & Arn. Rhamnaceae Southwest of Saudi Arabia (SZ) Danin and Fragman-Sapir (2016)
North of Pakistan, centre of Iran (IT) Rauf et al. (2016)
No. Species Family Exclusion region Reference and website The present study Bornkamm and Kehl (1990) Boulos (1999-2005) Abd El-Ghani et al. (2017)
420 Ziziphus spina-christi (L.) Desf. Rhamnaceae Burkina, Cameroon, Chad, Cyprus, Djibouti, Eritrea, Ethiopia, and Ghana (SZ) GBIF and POWO × ×
Gebel Elba (SZ) and Mediterranen coast in Egypt (ME) Boulos (1999-2005)
421 Zygophyllum arabicum (L.) Christenh. & Byng Zygophyllaceae Chad, Mali, Niger, and Sudan (SZ) Baayo (2008); GBIF; and POWO × ×
422 Zygophyllum bruguieri (DC.) Christenh. & Byng Zygophyllaceae Chad, Eritrea, Mali, Niger, Senegal, Somalia, and Sudan (SZ) GBIF and POWO × ×
Gebel Elba (SZ) and Mediterranen coast in Egypt (ME) Boulos (1999-2005)
423 Zygophyllum coccineum L. Zygophyllaceae Eritrea and Sudan (SZ) Boulos (1999-2005); GBIF; and POWO ×
424 Zygophyllum decumbens Delile Zygophyllaceae Namibia, Somalia, south of Oman, Sudan (SZ) Boulos (1999-2005); GBIF; and POWO × ×
425 Zygophyllum dumosum Boiss. Zygophyllaceae
426 Zygophyllum glutinosum (Delile) Christenh. & Byng Zygophyllaceae Chad, Mali, NiGebel Elbar, and Sudan (SZ) Flora of Libya and POWO × ×
Mediterranen coast in Egypt (ME) Boulos (1999-2005)
427 Zygophyllum indicum (Burm.f.) Christenh. & Byng Zygophyllaceae Chad, Djibouti Eritrea, Ethiopia, Kenya, Mali, and Niger (SZ) Boulos (1999-2005); GBIF; and POWO × ×
428 Zygophyllum molle (Delile) Christenh. & Byng Zygophyllaceae
429 Zygophyllum scabrum (Forssk.) Christenh. & Byng Zygophyllaceae

Note: × refers to the excluded species (surely non-Saharo-Arabian endemics), √ refers to the included species (surely Saharo-Arabian endemics), and ● refers to the extended species (species mainly distributed in Saharo-Arabian region but having extensions to neighbouring areas). SA, Saharo-Arabian region; IT, Irano-Turanian region; ME, Mediterranean region; SZ, Sudano-Zambezian region; ES, Euro-Siberian region. The manuscript also referred to Cairo University Herbarium (CAI), Agriculture Museum Herbarium (CAIM), and Tanta University Herbarium (TANE).

Then, the preliminary list of endemic plants of the Saharo-Arabian region in Egypt was assessed to confirm their occurrence by literature analysis. If the taxon was recorded in any countries outside the Saharo-Arabian region, it was excluded from the list. The preliminary list was assessed by the literature listed in Tables 1, 2, and 3. Table 1 shows the articles or books relating to the flora of Egypt; Table 2 shows the literature relating to the floras of other Saharo-Arabian countries; and Table 3 shows websites and global online databases relating to the Saharo-Arabian endemic plants.
Table 1 References of flora of Egypt used for the verification of occurrence of the Saharo-Atabian endemic plants in Egypt
Author Article or book title Journal or publisher Publication year
Abd El-Ghani, Hamdy, and Hamed Habitat diversity and floristic analysis of Wadi El-Natrun depression, Western Desert, Egypt Phytologia Balcanica 2015
Ahmed, Shaltout, Hosni, et al. Flora and vegetation of the different habitats of the western Mediterranean region of Egypt Taeckholmia 2015
Bedair, Shaltout, Ahmed, et al. Characterization of the wild trees and shrubs in the Egyptian flora Egyptian Journal of Botany 2020
Bedair, Ibrahim, Alyamani, et al. Impacts of anthropogenic disturbance on vegetation dynamics: a case study of Wadi Hagul, Eastern Desert, Egypt Plants 2021
Bedair, Shaltout, and Halmy A critical inventory of the Mediterranean endemics in the Egyptian flora Biodiversity and Conservation 2023
El-Khalafy Red list of the endemic and near endemic plant species in Egypt Tanta University 2018
El-Khalafy, Shaltout, and Ahmed Updating and assessing plant endemism in Egypt Phytotaxa 2021
El-Khalafy, Ahmed, Shaltout, et al. Re-assessment of the endemic taxa in the Egyptian Flora African Journal of Ecology 2021
El-Khalafy Biodiversity characteristics of endemic taxa in Egyptian flora Tanta University 2023
El-Khouly and Negm Management and Development of Agricultural and Natural Resources in Egypt's Desert Springer Nature 2021
El-Saied, El-Ghamry, Khafagi, et al. Floristic diversity and vegetation analysis of Siwa Oasis: An ancient agro-ecosystem in Egypt's Western Desert Annals of Agricultural Science 2015
Kamel, Zaghloul, Abd El-Wahab, et al. Current status of the flora of North Sinai: losses and gains Catrina 2008
Salama, Abd El-Ghani, and Gadallah Diversity and Plant FGs of the Vegetation in Eastern Desert of Egypt Lambert Academic Publishing 2016
Shaltout, El-Din, and Ahmed Plant Life in the Nile Delta Tanta University Press 2010
Table 2 References of floras of other countries in the Saharo-Arabian region used for the verification of occurrence of the Saharo-Arabian endemic plants in Egypt
Author Article or book title Publisher or journal name Publication year
Abdullah and Al-Dosari Vegetation of the State of Kuwait IUCN and Environment Public Authority of Kuwait 2022
Ali and Jafri Flora of Libya. Volumes 1-144 University of Tripoli 1976
Almasri, Hmidan, Shater, et al. Syria country report on the identification of important plant areas (IPAs) in Syria (Rapid assessment) National Coordinating Organization Syrian Society for Conservation of Wild Life 2010
Chaudhary Flora of the Kingdom of Saudi Arabia. Volume 1-3 Ministry of Agriculture and Water of Saudi Arabia 1999-2001
El-Gadi Flora of Libya. Volumes 145-147 University of Tripoli 1986
Feinbrun-Dothan Flora Palaestina: Part Three, Ericaceae to Compositae The Israel Academy of Science and Humanities 1978
Feinbrun-Dothan Flora Palaestina: Part Four. Text. Alismataceae to Orchidaceae. The Israel Academy of Science and Humanities 1986
Floc'h, Boulos, and Véla Synonymous Catalog with Commentary of the Flora of Tunisi Ministry of Environment and Sustainable Development, Tunisia 2010
Ghazanfar Status of the flora and plant conservation in the Sultanate of Oman Biological Conservation 1998
Guest Flora of Iraq, Vol. 1 Ministry of Agriculture of Iraq 1966
Kilian, Hein, and Hubaishan New and noteworthy records for the flora of Yemen, chiefly of Hadhramout and Al-Mahra Willdenowia 2002
Meddour, Sahar, and Médail Checklist of the native tree flora of Algeria: diversity, distribution, and conservation Plant Ecology and Evolution 2021
Norton, Böer, Allan, et al. An Illustrated Checklist of the Flora of Qatar United Nations Educational Scientific and Cultural Organization (UNESCO) 2009
Taifour Jordan Plant Red List Volume II Royal Botanic Garden of Jordan 2017
Taifour and El-Oqlah Jordan Plant Red List Volume I Royal Botanic Garden of Jordan 2016
Taleb and Fennane Vascular Plant Communities of Morocco: Phytosociology, Ecology and Geography Springer International Publishing 2019
Yaseen, Ahmad, Sultana, et al. Ethnobotany of medicinal plants in the Thar Desert (Sindh) of Pakistan Journal of Ethnopharmacology 2015
Zohary Flora Palaestina Part I The Israel Academy of Science and Humanities 1968
Zohary Flora Palaestina Part II The Israel Academy of Science and Humanities 1987
Table 3 Global online databases utilized for the verification of occurrence of the Saharo-Arabian endemic plants in Egypt
Database Website
African Plant Database https://africanplantdatabase.ch/en
Algeria Native Plants https://algerianativeplants.net/html-ang/plante-algerie-recherche.php
Catalogue of Life http://www.catalogueoflife.org/annualchecklist/2010
Database of Weed Species in Crops and Countries https://www.fao.org/agriculture/crops/thematic-sitemap/theme/biodiversity/weeds/db-countries/l/en/
Euro+Med Plant Base https://europlusmed.org/
Flora of Greece https://portal.cybertaxonomy.org/flora-greece/intro
Flora of Israel Online http://flora.org.il
Flora of Lebanon http://lebanon-flora.org/
Flora of Libya https://www.floraoflibya.services.ly/
Flora of Morocco https://herbaria.plants.ox.ac.uk/bol/floraofmorocco/Explore
Flora of Pakistan http://www.efloras.org/index.aspx
Flora of Qatar https://www.floraofqatar.com/
Flora of the Kingdom of Saudi Arabia https://floraofksa.myspecies.info/
Global Biodiversity Information Facility (GBIF) https://www.gbif.org/
Global Compositae Database (GCD) https://www.compositae.org/index.php
Global Plants on JSTOR https://plants.jstor.org/
Legume Information System (LIS) https://www.legumeinfo.org/taxa/
National Registry for Egyptian Herbaria http://networks.asrt.sci.eg/Herbarias/Index?searchBy=Species&Search_Data
Old Dominion University Plant Site: Checklist of Plants of the Hashemite Kingdom of Jordan https://ww2.odu.edu/~lmusselm/plant/index.php
Plant Diversity of Saudi Arabia https://www.plantdiversityofsaudiarabia.info/index.htm
Plants of the Middle East https://www.plantsofthemiddleeast.com/
Plants of the World Online (POWO) http://www.plantsoftheworldonline.org
Turkish Plants Data Service (TÜBİVES) http://194.27.225.161/yasin/tubives/index.php
World Plants https://www.worldplants.de/world-plants-complete-list/complete-plant-list
Data were additionally gathered from field trips. Cairo University, Alexandria University, the herbaria of Kafrelsheikh University, Assiut University, Tanta University, Agriculture Museum of Cairo, National Research Centre of Egypt, Desert Research Center of Egypt, and National Registry for Egyptian Herbaria also provided plant data for the endemic species research. The accessible literature (articles, books, master's theses, doctoral dissertations, and scientific reports) in above mentioned institutes was also used for gathering the information. The taxa in the list were organized alphabetically using the Angiosperm Phylogeny Group IV system (Byng et al., 2016). The accepted name of the recorded taxa was assessed using the World Checklist of selected Plant Families (WCSP) and Plants of the World Online (POWO) websites.
The final checklist of the present study includes three categories: the excluded species (surely non-Saharo-Arabian endemics), the included species (surely Saharo-Arabian endemics), and the extended species (species that are primarily found in the Saharo-Arabian region but extend to nearby regions). This study provides a critical and dynamic checklist of the Saharo-Arabian endemic plants in Egypt (Table S1).

2.2.2 Geographical distribution

In the present study, we identified the phytogeographical sectors according to Täckholm (1974). The system of White and Léonard (1991) was used for the assessment of the recorded taxa's global distribution (floristic regions). We defined the distribution of the taxa that were recorded in the countries of the Saharo-Arabian region depending on several databases (Table 3). Floras of countries in the Saharo-Arabian region were also used for the assessment of recorded taxa (Table 4).
Table 4 References used for the assessment of the geographical distribution of the Saharo-Arabian endemic plants in Egypt
Author Article or book title Journal or publisher Publication year
Abdullah and Al-Dosari Vegetation of the State of Kuwait IUCN and Environment Public Authority of Kuwait 2022
Ali and Jafri Flora of Libya. Volumes 1-144 University of Tripoli 1976
Almasri, Hmidan, Shater, et al. Syria country report on the identification of important plant areas (IPAs) in Syria (Rapid assessment) National Coordinating Organization Syrian Society for Conservation of Wild Life 2010
Al-Sodany, El-Khalafy, Ahmed, et al. Plant communities associated with Egyptian endemic flora Egyptian Journal of Botany 2024
Boulos Flora of Egypt. Volumes 1-4 Al-Hadara Publishing 1999-2005
Chaudhary Flora of the Kingdom of Saudi Arabia. Volumes 1-3 Ministry of Agriculture and Water of Saudi Arabia 1999-2001
El-Gadi Flora of Libya. Volumes 145-147 University of Tripoli 1986
El-Hadidi List of Threatened Plants in the Flora of Egypt Kew 1979
El-Hadidi Flora Aegyptiaca Palm Press 2003
Floc'h, Boulos, and Véla Synonymous Catalog with Commentary of the Flora of Tunisi Ministry of Environment and Sustainable Development, Tunisia 2010
Ghazanfar Status of the flora and plant conservation in the Sultanate of Oman Biological Conservation 1998
Kilian, Hein, and Hubaishan New and noteworthy records for the flora of Yemen, chiefly of Hadhramout and Al-Mahra Willdenowia 2002
Meddour, Sahar, and Médail Checklist of the native tree flora of Algeria: diversity, distribution, and conservation Plant Ecology and Evolution 2021
Norton, Böer, Allan, et al. An Illustrated Checklist of the Flora of Qatar UNESCO 2009
Täckholm Students' Flora of Egypt Cairo University Press 1974
Taifour Jordan Plant Red List Volume II Royal Botanic Garden of Jordan 2017
Taifour and El-Oqlah Jordan Plant Red List Volume I Royal Botanic Garden of Jordan 2016
Taleb and Fennane Vascular Plant Communities of Morocco: Phytosociology, Ecology and Geography, Geobotany studies Springer International Publishing 2019
Yaseen, Ahmad, Sultana, et al. Ethnobotany of medicinal plants in the Thar Desert (Sindh) of Pakistan Journal of Ethnopharmacology 2015
Zohary Flora Palaestina Part I The Israel Academy of Science and Humanities 1968
Zohary Flora Palaestina Part II The Israel Academy of Science and Humanities 1987

2.3 Multivariate analysis

The two-way indicator species analysis (TWINSPAN) was used to classify species and countries of occurrence (Hill, 1979). The detrended correspondence analysis (DECORANA) was used for the ordination of countries and species (Hill and Gauch, 1980). We classified the groups of countries using TWINSPAN according to the common species among each group of countries. And we named each group according to its first dominant species. The Community Analysis Package (IRC house, Hampshire, UK) was used for the TWINSPAN and DECORANA analysis (Seaby and Henderson, 2007).

3 Results

3.1 Verification of the list

The prepared prelist of endemic plants of the Saharo-Arabian region in Egypt included 429 taxa. The present study authorized 126 taxa (29.4% of the total 429 taxa) as surely Saharo-Arabian endemics in Egypt (Table S1). The number of the excluded taxa (surely non-Saharo-Arabian endemics) in this study was 209 taxa (48.7%). Three taxa were recognized as the Saharo-Arabian endemics only by the present study, 104 taxa (82.5% of the total Saharo-Arabian endemics) were recognized as the Saharo-Arabian endemics by this study and one previous study—Boulos (1999-2005), 18 taxa (14.3%) were recognized as the Saharo-Arabian endemics by the present study and two previous studies—Boulos (1999-2005) and Abd El-Ghani et al. (2017), only one taxon (0.79%) was recorded as the Saharo-Arabian endemic plant by the present study and three previous studies—Bornkamm and Kehl (1990), Boulos (1999-2005), and Abd El-Ghani et al. (2017). On the other hand, 28 taxa were excluded by the present study only, 122 taxa were excluded by the present study and one previous study (of which 115 taxa were excluded as well as Boulos (1999-2005), while 7 taxa were excluded as well as Abd El-Ghani et al. (2017)), 59 taxa were excluded by the present study and two previous studies (of which 15 taxa were excluded as well as Boulos (1999-2005) and Abd El-Ghani et al. (2017), while 44 taxa were excluded as well as Bornkamm and Kehl (1990) and Boulos (1999-2005)). Besides, 94 taxa (21.9%) were primarily found in the Saharo-Arabian region and extend to nearby regions (i.e., extended species).
The present study verified 126 endemic taxa of the Saharo-Arabian region in Egypt (109 species, 17 subspecies), belonging to 87 genera and 37 families. The major taxonomic divisions revealed that the eudicots had the most diversity including 33 families, 82 genera, 103 species, and 14 subspecies, the monocots included 3 families, 4 genera, 6 species, and 2 subspecies, and the gymonsperm included 1 family (Ephedraceae). The most representative family was Asteraceae (17 species; 13.5%), followed by Lamiaceae (13 species; 10.3%), and Caryophyllaceae and Fabaceae (12 species, respectively; 9.5%) (Table 5).
Table 5 Taxonomic diversity of the Saharo-Arabian endemic plants in Egypt
Family Number of genus Number of species Number of subspecies
Gymnospermae Ephedraceae 1 0 1
Monocotyledon Amaryllidaceae 1 3 0
Asparagaceae 2 2 2
Poaceae 1 1 0
Dicotyledon Amaranthaceae 2 1 2
Anacardiaceae 1 0 1
Apiaceae 4 4 0
Apocynaceae 2 2 0
Asphodelaceae 1 1 0
Asteraceae 14 17 0
Boraginaceae 2 2 0
Brassicaceae 5 4 1
Campanulaceae 1 1 0
Caprifoliaceae 1 2 0
Caryophyllaceae 6 12 4
Cistaceae 1 3 0
Colchicaceae 1 1 0
Convolvulaceae 1 2 0
Euphorbiaceae 1 1 0
Fabaceae 6 12 1
Gentianaceae 1 1 0
Hypericaceae 1 1 0
Lamiaceae 10 13 1
Molluginaceae 1 1 0
Plantaginaceae 3 2 1
Polygalaceae 1 0 2
Polygonaceae 2 1 1
Primulaceae 1 1 0
Resedaceae 1 2 0
Rhamnaceae 1 1 0
Rosaceae 2 2 0
Rubiaceae 2 2 0
Rutaceae 1 1 0
Scrophulariaceae 2 4 0
Solanaceae 1 1 0
Tamariaceae 1 1 0
Zygophyllaceae 2 4 0
Total 87 109 17

3.2 Geographical distribution

3.2.1 National geographic distribution

According to Täckholm (1974), he divided Egypt (except the Mediterranean coast and Red Sea coast) into nine phytogeographic regions. The South Sinai (south of the Sinai Peninsula) was the most rich region, comprising 83 taxa (65.9%) of the total endemic species (e.g., Reseda stenostachya Boiss., Convolvulus spicatus Peter ex Hallier f., and Phagnalon sinaicum Bornm. & Kneuck.), followed by the Isthemic Desert (the middle of Sinai) (53 taxa; 42.1%) (e.g., Pterocephalus arabicus Boiss. Bellevalia zoharyi Feinbrun, and Allium rothii Zucc.), the Arabian Desert (located in the eastern Nile River) (40 taxa; 31.7%) (e.g., Centaurea scoparia Sieb. ex Spreng., Echinops glaberrimus DC., and Launaea spinosa (Forssk.) Sch. Bip. ex Kuntze), the Oases (8 taxa; 6.3%), the Red Sea and Nile Delta (5 taxa; 4.0%), the Nile Valley (4 taxa; 3.2%), the Libyan Desert (3 taxa; 2.4%), and the Nile Faiyum (1 taxon; 0.8%) (Fig. 1).
Fig. 1 Quantitative distribution of the Saharo-Arabian endemic plants in Egypt

3.2.2 Global geographical distribution

The recorded 126 species in this study are endemics to the Saharo-Arabian region and located in Egypt. The SS1 subzone had all the 126 Saharo-Arabian endemic taxa (100.0%), the SS2 subzone had 79 taxa (62.7%), and the SS3 subzone distributed 14 taxa (11.1%) (Table S2). In this study, the Saharo-Arabian region included 21 countries (6 countries in Africa and 15 countries in Asia, excluding Western Sahara due to lack of data). The highest number of the Saharo-Arabian endemic plants in Egypt was recorded in Palestine (79 taxa; 62.7%), followed by Jordan (72 taxa; 57.1%), Saudi Arabia (67 taxa; 53.2%), Iraq (19 taxa; 15.1%), and Iran (14 taxa; 11.1%); while the lowest number of taxa was recorded in Mauritania and India (2 taxa and 1.6% for each other) (Fig. 2).
Table S2 Distribution of the Saharo-Arabian endemic plants in the Saharo-Arabian countries
Species Country Disrtibution area
Eg SA Ku Ba Qa UAE Ye Om Irq Irn Pak Af Jo Pal Sy Li Tu Al Mo Ma In Total
Amaranthaceae
Agathophora alopecuroides var. alopecuroides (Delile) Fenzl ex Bunge 14 SS1, SS2, and SS3
Agathophora alopecuroides var. papillosa (Maire) Boulos 14 SS1, SS2, and SS3
Haloxylon negevensis (Iijin & Zohary) Boulos 2 SS1 and SS2
Amaryllidaceae
Allium decaisnei C.Presl 2 SS1 and SS2
Allium rothii Zucc. 3 SS1 and SS2
Allium sinaiticum Boiss. 4 SS1 and SS2
Anacardiaceae
Pistacia khinjuk Stocks 2 SS1 and SS2
Apiaceae
Daucus sahariensis Murb. 5 SS1
Ducrosia ismaelis Asch. 2 SS1 and SS2
Pimpinella schweinfurthii Asch. 4 SS1 and SS2
Pycnocycla tomentosa Decne. 3 SS1 and SS2
Apocynaceae
Gomphocarpus sinaicus (Boiss.) 5 SS1 and SS2
Rhazya greissii Tackh. & Boulos 1 SS1
Species Country Disrtibution area
Eg SA Ku Ba Qa UAE Ye Om Irq Irn Pak Af Jo Pal Sy Li Tu Al Mo Ma In Total
Asparagaceae
Bellevalia desertorum Eig & Feinbrun 3 SS1 and SS2
Bellevalia flexuosa var. galalensis Tackh. & Drar 1 SS1
Bellevalia zoharyi Feinbrun 3 SS1 and SS2
Muscari longipes subsp. negevense (Feinbrun & Danin) 3 SS1and SS2
Asphodelaceae
Asphodelus refractus Boiss. 9 SS1 and SS2
Asteraceae
Anthemis scrobicularis Yavin 4 SS1 and SS2
Anvillea garcinii (Burm.f.) DC. 13 SS1, SS2, and SS3
Atractylis boulosii Tackh. 2 SS1 and SS2
Atractylis mernephthae Asch. & Schweinf. & Letourn. 7 SS1 and SS2
Centaurea scoparia Sieber ex Spreng 4 SS1 and SS2
Centaurea sinaica DC. 10 SS1 and SS2
Chiliadenus montanus (Vahl) Brullo 1 5 SS1, SS2, and SS3
Crepis nigricans Viv. 4 SS1
Echinops glaberrimus DC. 4 SS1 and SS2
Species Country Disrtibution area
Eg SA Ku Ba Qa UAE Ye Om Irq Irn Pak Af Jo Pal Sy Li Tu Al Mo Ma In Total
Iphiona mucronata (Forssk.) Asch. & Schweinf. 4 SS1 and SS2
Launaea spinosa (Forssk.) Sch. Bip. Ex Kuntze 6 SS1 and SS2
Phagnalon sinaicum Bornm. & Kneuck. 4 SS1 and SS2
Picris sulphurea Delile 2 SS1 and SS2
Scorzonera drarii Tackh. 1 SS1
Scorzonera schweinfurthii Boiss. 5 SS1 and SS2
Senecio belbeysius Delile 1 SS1
Tanacetum sinaicum (Fresen.) Delile ex K. Bremer & Humphries 4 SS1 and SS2
Boraginaceae
Arnebia tinctoria (Forssk.) 7 SS1 and SS2
Podonosma galalensis Schweinf. ex Boiss. 3 SS1 and SS2
Brassicaceae
Crucihimalaya kneuckeri (Bornm.) Al-Shehbaz, O'Kane & R. A. Price 2 SS1 and SS2
Eremobium aegyptiacum var. lineare (Delile) Zohary 4 SS1 and SS2
Matthiola arabica Boiss. 5 SS1 and SS2
Pseuderucaria teretifolia (Desf.) O. E. Schulz 5 SS1
Schimpera arabica Hochst. & Steud. 9 SS1, SS2, and SS3
Species Country Disrtibution area
Eg SA Ku Ba Qa UAE Ye Om Irq Irn Pak Af Jo Pal Sy Li Tu Al Mo Ma In Total
Campanulaceae
Campanula dulcis Decne. 4 SS1 and SS2
Caprifoliaceae
Pterocephalus arabicus Boiss. 2 SS1 and SS2
Pterocephalus sanctus Decne. 4 SS1 and SS2
Caryophyllaceae
Bufonia multiceps Decne. 1 SS1
Dianthus guessfeldtianus Muschl. 1 SS1
Dianthus monadelphus subsp. judaicus (Boiss.) Greuter & Burdet 5 SS1 and SS2
Dianthus sinaicus Boiss. 4 SS1 and SS2
Eremogone sinaica (Boiss.) Dillenb. & Kadereit 2 SS1 and SS2
Paronychia sinaica Fresen. 4 SS1 and SS2
Petrorhagia arabica (Boiss.) P.W. Ball & Heywood 3 SS1 and SS2
Silene arabica Boiss. 11 SS1, SS2, and SS3
Silene hussonii Boiss. 4 SS1 and SS2
Silene leucophylla Boiss. 1 SS1
Silene linearis Decne. 6 SS1 and SS2
Silene odontopetala var. congesta Boiss. 2 SS1 and SS3
Silene oreosinaica Chowdhuri 1 SS1
Species Country Disrtibution area
Eg SA Ku Ba Qa UAE Ye Om Irq Irn Pak Af Jo Pal Sy Li Tu Al Mo Ma In Total
Silene schimperiana Boiss. 1 SS1
Silene villosa var. graveolens Sickenb. 1 SS1
Silene vivianii subsp. viscida (Boiss.) Boulos 3 SS1, SS2, and SS3
Cistaceae
Helianthemum sancti-antoni Schweinf. ex Boiss. 4 SS1 and SS2
Helianthemum schweinfurthii Grosser 1 SS1
Helianthemum ventosum Boiss. 4 SS1 and SS2
Colchicaceae
Colchicum guessfeldtianum Asch. & Schweinf. 2 SS1 and SS2
Convolvulaceae
Convolvulus schimperi Boiss. 2 SS1 and SS2
Convolvulus spicatus Peter ex Hallier f. 5 SS1, SS2, and SS3
Ephedraceae
Ephedra pachyclada subsp. sinaica (Riedl) Freitag & Maier-St 5 SS1 and SS2
Euphorbiaceae
Euphorbia obovata Decne. 4 SS1, SS2, and SS3
Fabaceae
Argyrolobium saharae Pomel 3 SS1
Species Country Disrtibution area
Eg SA Ku Ba Qa UAE Ye Om Irq Irn Pak Af Jo Pal Sy Li Tu Al Mo Ma In Total
Astragalus amalecitanus Boiss. 2 SS1 and SS2
Astragalus dactylocarpus subsp. acinaciferus (Boiss.) E. Ott 8 SS1, SS2, and SS3
Astragalus fresenii Decne 1 SS1
Astragalus intercedens Sam. ex Rech.f. 5 SS1 and SS2
Astragalus sanctus Boiss. 4 SS1 and SS2
Astragalus schimperi Boiss. 15 SS1, SS2, and SS3
Astragalus sparsus Decne 5 SS1 and SS2
Lotus lanuginosus Vent. 6 SS1 and SS2
Melilotus serratifolius Tackh. & Boulos 1 SS1
Tephrosia kassasii Boulos 1 SS1
Trigonella media Delile ex Urb. 1 SS1
Trigonella schlumbergeri Boiss. 1
Gentianaceae
Centaurium malzacianum Maire 2 SS1 and SS2
Hypericaceae
Hypericum sinaicum Hochst. ex Boiss. 4 SS1 and SS2
Lamiaceae
Ballota kaiseri Tackh. 1 SS1
Lavandula atriplicifolia Benth. 3 SS1 and SS2
Species Country Disrtibution area
Eg SA Ku Ba Qa UAE Ye Om Irq Irn Pak Af Jo Pal Sy Li Tu Al Mo Ma In Total
Micromeria serbaliana (Danin & Hedge) 1 SS1
Micromeria sinaica (Benth.) Briq. 3 SS1 and SS2
Nepeta septemcrenata Ehrenb. ex Benth. 4 SS1 and SS2
Origanum isthmicum Danin 2 SS1and SS2
Origanum syriacum subsp. sinaicum (Boiss.) Greuter & Burdet 1 SS1
Phlomis aurea Decne. 2 SS1 and SS2
Salvia deserti Decne. 4 SS1 and SS2
Stachys aegyptiaca Pers. 4 SS1 and SS2
Teucrium decaisnei C. Presl 3 SS1 and SS2
Teucrium leucocladum Boiss. 4 SS1 and SS2
Thymus bovei Benth. 4 SS1 and SS2
Thymus decussatus Benth. 3 SS1 and SS2
Molluginaceae
Glinus runkewitzii Tackh. & Boulos 1 SS1
Plantaginaceae
Anarrhinum forskaohlii subsp. pubescens D.A.Sutton 1 SS1
Plantago sinaica (Barneoud) Decne. 3 SS1 and SS2
Veronica kaiseri Tackh. 2 SS1 and SS2
Species Country Disrtibution area
Eg SA Ku Ba Qa UAE Ye Om Irq Irn Pak Af Jo Pal Sy Li Tu Al Mo Ma In Total
Poaceae
Stipagrostis drarii (Tackh.) De Winter 8 SS1 and SS2
Polygalaceae
Polygala sinaica var. glabrescens (Zohary) Boulos 4 SS1 and SS2
Polygala sinaica var. sinaica Botsch. 1 SS1
Polygonaceae
Atraphaxis spinosa var. sinaica (Jaub. & Spach) Boiss. 3 SS1 and SS2
Persicaria obtusifolia (Täckh. & Boulos) Greuter & Burdet 1 SS1
Primulaceae
Evotrochis involucrata Raf. 1 SS1
Resedaceae
Reseda muricata C. Presl 12 SS1, SS2, and SS3
Reseda stenostachya Boiss. 3 SS1 and SS2
Rhamnaceae
Rhamnus disperma Ehrenb. ex Boiss. 4 SS1 and SS2
Rosaceae
Cotoneaster orbicularis Schltdl. 4 SS1, SS2, and SS3
Rosa arabica (Crép. ex Boiss.) Déségl. 1 SS1
Species Country Disrtibution area
Eg SA Ku Ba Qa UAE Ye Om Irq Irn Pak Af Jo Pal Sy Li Tu Al Mo Ma In Total
Rubiaceae
Crucianella ciliata Lam. 6 SS1, SS2, and SS3
Galium sinaicum (Delile ex Decne) Boiss. 4 SS1 and SS2
Rutaceae
Haplophyllum poorei C. C. Towns. 3 SS1 and SS2
Scrophulariaceae
Scrophularia deserti Delile 16 SS1, SS2, and SS3
Verbascum decaisneanum Kuntze 4 SS1 and SS2
Verbascum fruticulosum Post & Autran 3 SS1and SS2
Verbascum schimperianum Boiss. 4 SS1 and SS2
Solanaceae
Hyoscyamus boveanus (Dunal) Asch. & Schweinf. 1 SS1
Tamaricaceae
Reaumuria negevensis Zohary & Danin 2 SS1and SS2
Zygophyllaceae
Tribulus kaiseri Hosni 3 SS1, SS2, and SS3
Zygophyllum dumosum Boiss. 3 SS1 and SS2
Zygophyllum molle (Delile) Christenh. & Byng 4 SS1 and SS2
Species Country Disrtibution area
Eg SA Ku Ba Qa UAE Ye Om Irq Irn Pak Af Jo Pal Sy Li Tu Al Mo Ma In Total
Zygophyllum scabrum (Forssk.) Christenh. & Byng 6 SS1 and SS2
Total 126 67 13 3 7 9 11 8 19 14 6 5 72 79 9 12 7 10 9 2 2 490

Note: Eg, Egypt; SA, Saudi Arabia; Ku, Kuwait; Ba, Bahrain; Qa, Qatar; UAE, the United Arab Emirates; Ye, Yemen; Om, Oman; Irq, Iraq; Irn, Iran; Pak, Pakistan; Af, Afghanistan; Jo, Jordan; Pal, Palestine; Sy, Syria; Li, Libya; Tu, Tunisia; Al, Algeria; Mo, Morroco; Ma, Mauritania; In, India; SS1, Sahara regional subzone; SS2, Saharo-Arabian subzone; SS3, Nubo-Sindian subzone. Note that the Saharo-Arabian floristic region zoning is based on the White and Léonard (1991), of which SS1 subzone includes Morocco, Mauritania, Algeria, Tunisia, Libya, and Egypt, SS2 subzone includes Palestine, Kuwait, Saudi Arabia, Qatar, Bahrain, the United Arab Emirates, Oman, Yemen, and southern Jordan, Syria, and Iraq, and SS3 subzone includes southern Iran, Afghanistan, and Pakistan and northern India.

Fig. 2 Quantitative distribution of the Saharo-Arabian endemic plants in each Saharo-Arabian country. Note that the Western Sahara is not included in the research scope of this paper due to lack of data.

3.3 Endemic species in Egypt

Remarkably, 25 taxa (19.8%) of the recorded species in this study were exclusive to Egypt (Table 6), and 18 taxa (14.3%) were near endemic (in Egypt and another country). To be specific, 7 taxa were recorded in Egypt and Palestine (Haloxylon negevensis (Iljin & Zohary) L. Boulos, Allium decaisnei C. Presl, Atractylis boulosii Täckh., Eremogone sinaica (Boiss.) Dillenb. & Kadereit, Astragalus amalecitanus Boiss., Origanum isthmicum Danin, and Reaumuria negevensis Zohary & Danin), 6 species in Egypt and Saudi Arabia (Pistacia khinjuk Stocks, Ducrosia ismaelis Asch., Picris sulphurea Delile, Crucihimalaya kneuckeri (Bornm.) Al-Shehbaz, O'Kane & R. A. Price, Colchicum guessfeldtianum Asch. & Schweinf., and Centaurium malzacianum Maire), 4 species in Egypt and Jordan (Pterocephalus arabicus Boiss., Convolvulus schimperi Engl., Phlomis aurea Decne., and Veronica kaiseri Täckh.), and 1 species in Egypt and Iran (Silene odontopetala var. congesta Boiss.). Whereas Scrophularia deserti Delile and Astragalus schimperi Boiss. Were abundantly distributed in the countries of the Saharo-Arabian region.
Table 6 List of the exclusive Saharo-Arabian endemic plants in Egypt
No. Endemic species Family
1 Anarrhinum forskaohlii subsp. pubescens D. A. Sutton Plantaginaceae
2 Astragalus fresenii Decne Fabaceae
3 Ballota kaiseri Tackh. Lamiaceae
4 Bellevalia flexuosa var. galalensis Tackh. & Drar Asparagaceae
5 Bufonia multiceps Decne Caryophyllaceae
6 Dianthus guessfeldtianus Muschl. Caryophyllaceae
7 Glinus runkewitzii Tackh. & Boulos Molluginaceae
8 Helianthemum schweinfurthii Grosser Cistaceae
9 Hyoscyamus boveanus (Dunal) Asch. & Schweinf Solanaceae
10 Melilotus serratifolius Tackh. & Boulos Fabaceae
11 Micromeria serbaliana Danin & Hedge Lamiaceae
12 Origanum syriacum subsp. sinaicum (Boiss.) Greuter & Burde Lamiaceae
13 Persicaria obtusifolia (Täckh. & Boulos) Greuter & Burdet Polygonaceae
14 Polygala sinaica Botsch. var. sinaica Polygalaceae
15 Primula boveana Decne. ex Duby Primulaceae
16 Rhazya greissii Tackh. & Boulos Apocynaceae
17 Rosa arabica (Crép. ex Boiss.) Déségl. Rosaceae
18 Scorzonera drarii Tackh. Asteraceae
19 Senecio belbeysius Delile Asteraceae
20 Silene leucophylla Boiss. Caryophyllaceae
21 Silene oreosinaica Chowdhuri Caryophyllaceae
22 Silene schimperiana Boiss. Caryophyllaceae
23 Silene villosa var. graveolens Sickenb. Caryophyllaceae
24 Tephrosia kassasii Boulos Fabaceae
25 Trigonella media Delile ex Urb. Fabaceae

3.4 Clustering analysis

Seven groups were recognized at the fourth level of classification as a result of the application of TWINSPAN to the Saharo-Arabian endemic plants in Egypt (Fig. 3; Table 7). Figure 4 shows the results of DECORANA analysis. We determined the names of the groups according to the first dominant species that had the highest presence percentage in each group, as follows:
Fig. 3 Dendrogram showing the most representative seven groups produced after using TWINSPAN classification technique at the fourth level. I, Asphodelus refractus group; II, Agathophora alopecuroides var. papillosa group; III, Anvillea garcinii group; IV, Reseda muricata group; V, Agathophora alopecuroides var. alopecuroides; VI, Scrophularia deserti group; VII, Astragalus schimperi group.
Table 7 Vegetation groups of the 126 Saharo-Arabian endemic plants in Egypt
Vegetation group Number of species Distribution country Percentage of African country (%) Percentage of Asian country (%) Dominant species
I Asphodelus refractus group 10 Algeria, Morocco, Mauritania, and Tunisia 100.0 0.0 Asphodelus refractus Boiss., Agathophora alopecuroides (Delile) Fenzl ex Bunge, Anvillea garcinii (Burm. f.) DC., Pseuderucaria teretifolia (Desf.) O. E. Schulz, and Zygophyllum scabrum (Forssk.) Christenh. & Byng.
II Agathophora alopecuroides var. papillosa group 12 Libya 100.0 0.0 Agathophora alopecuroides var. papillosa (Maire) Boulos, Daucus sahariensis Murb., and Anvillea garcinii (Burm. f.) DC.
III Anvillea garcinii group 11 Yemen 0.0 100.0 Anvillea garcinii (Burm. f.) DC., Gomphocarpus sinaicus Boiss., and Launaea spinosa (Forssk.) Sch. Bip. ex Kuntze.
IV Reseda muricata group 126 Egypt, Jordan, Palestine, and Saudi Arabia 25.0 75.0 Reseda muricata C.Presl, Allium sinaiticum Boiss., Agathophora alopecuroides (Delile) Fenzl ex Bunge, Gomphocarpus sinaicus Boiss., and Anthemis scrobicularis Yavin.
V Agathophora alopecuroides var. alopecuroides group 23 Iran, Iraq, Kuwait, and Qatar 0.0 100.0 Agathophora alopecuroides var. alopecuroides, Silene arabica F.N.Williams, Astragalus schimperi Boiss., and Reseda muricata C. Presl
VI Scrophularia desert group 11 India, Pakistan, and Syria 0.0 100.0 Scrophularia deserti Delile, Astragalus schimperi Boiss., Euphorbia obovata Decne., and Agathophora alopecuroides (Delile) Fenzl ex Bunge
VII Astragalus schimperi group 13 Afghanistan, Bahrain, Oman, and the United Arab Emirates 0.0 100.0 Astragalus schimperi Boiss, Reseda muricata C. Presl, Scrophularia deserti Delile, Cotoneaster orbicularis Schltdl., and Silene arabica F. N. Williams
Fig. 4 Detrended correspondence analysis (DECORANA) ordination plot. SA, Saudi Arabia; UAE, the United Arab Emirates.
I Asphodelus refractus group. This group involved Tunisia, Algeria, Morocco, and Mauritania. The most common species among these countries were Asphodelus refractus Boiss., Agathophora alopecuroides (Delile) Fenzl ex Bunge, Anvillea garcinii (Burm.f.) DC., Pseuderucaria teretifolia (Desf.) O.E.Schulz, and Zygophyllum scabrum (Forssk.) Christenh. & Byng.
II Agathophora alopecuroides var. papillosa group. This group included only Libya. The dominant species in Libya include Agathophora alopecuroides var. papillosa (Maire) Boulos, Daucus sahariensis Murb., and Anvillea garcinii (Burm. f.) DC.
III Anvillea garcinii group. This group included only Yemen. The dominant species in Yemen included Anvillea garcinii (Burm. f.) DC., Gomphocarpus sinaicus Boiss., and Launaea spinosa (Forssk.) Sch. Bip. ex Kuntze.
IV Reseda muricata group. This group involved Egypt, Saudi Arabia, Palestine, and Jordan. The most common species among these countries were Reseda muricata C. Presl, Allium sinaiticum Boiss., Agathophora alopecuroides (Delile) Fenzl ex Bunge, Gomphocarpus sinaicus Boiss., and Anthemis scrobicularis Yavin.
V Agathophora alopecuroides var. alopecuroides group. This group included Iran, Iraq, Qatar, and Kuwait. The most common species among these countries were Agathophora alopecuroides var. alopecuroides, Silene arabica F. N. Williams, Astragalus schimperi Boiss., and Reseda muricata C. Presl.
VI Scrophularia deserti group. This group involved Pakistan, India, and Syria. The most common species among these countries were Scrophularia deserti Delile, Astragalus schimperi Boiss., Euphorbia obovata Decne., and Agathophora alopecuroides (Delile) Fenzl ex Bunge.
VII Astragalus schimperi group. This group involved Afghanistan, Oman, the United Arab Emirates, and Bahrain. The most common species among these countries were Astragalus schimperi Boiss, Reseda muricata C. Presl, Scrophularia deserti Delile, Cotoneaster orbicularis Schltdl., and Silene arabica F. N. Williams.

4 Discussion

The results of the current study support the idea that the Saharo-Arabian region is characterized by hot deserts, semi-deserts, and savanna. The Saharo-Arabian countries include Morocco, Mauritania, Western Sahara (not included in this study), Algeria, Tunisia, Libya, Egypt, Palestine, Kuwait, Saudi Arabia, Qatar, Bahrain, the United Arab Emirates, Oman, Yemen, southern Jordon, Syria, Iraq, Iran, Afghanistan, Pakistan, as well as northern India.
A large portion of the flora of the Saharo-Arabian region is shared with the nearby Mediterranean, Irano-Turanian, and Sudano-Zambezian regions of the Paleotropical Kingdom and the Holarctic Kingdom (White and Léonard, 1991). The boundary of the Saharo-Arabian region in the Nile Delta starts after an average distance of 15-25 km in north-south direction from the coast (Zahran and Willis, 2009). This distance includes a transition zone between the Mediterranean and the Saharo-Arabian floristic region that is called Mediterranean-Sahara regional transition zone (El Hadidi, 2003). This transition zone extends for an average width ranging 5-10 km in south direction from the coast. The boundary of the Saharo-Arabian region in the Western Desert starts after an average distance of 45-50 km in north-south direction from the coast in which the Mediterranean-Sahara regional transition zone extends for an average width ranging 5-10 km in south direction from the coast (El Hadidi, 2003). The Sinai Peninsula totally lies within the Saharo-Arabian floristic region except the Mediterranean coastal strip of North Sinai (about 13.0% of Sinai Peninsula) (Zahran et al., 1985; White and Léonard, 1991; Heneidak, 2008). Danin and Plitmann (1987) stated that the highlands of the South Sinai are located in the Irano-Turanian chorotype, while the lowlands of South Sinai are located in the Sudano-Zambezian chorotype. Also, Gebel Elba Mountains that are excluded from the Saharo-Arabian floristic region, are located in the Sudano-Zambezian chorotype (Eig, 1931; White and Léonard, 1991).
Regarding the national distribution of the Saharo-Arabian endemic plants in Egypt, Sinai (especially South Sinai) is the richest region, comprising 83 taxa (65.9%) of the total endemic species. Numerous microhabitats can be found in Sinai Peninsula due to the variety of geology and landforms. Each of them has distinct climatic conditions and a flora that is rich in endemic and medicinal plants (Khedr, 2021). The massive mountains in Sinai Peninsula, which limit the dispersion of seeds over long distances, are likely responsible for the region's abundance of endemic species. In particular, the Saint Katherine Protectorate (SKP) in South Sinai is the most important region for endemism in Egypt (Zohary, 1973). Moreover, in Sinai Peninsula, the wadis serve as drainage channels for gathering rainwater from the surroundings and facilitating the growth of plants. Consequently, these wadis are floristically diverse and provide habitat for many wild species (Khedr, 2021). The highest number of the Saharo-Arabian endemic plants in Egypt also distributes in other Saharo-Arabian countries, especially Palestine, Jordan, Saudi Arabia, Iraq, Iran, and Kuwait, and most of them are distributed in Sinai Peninsula. This can be explained by the separation of the Arabian Peninsula and Sinai subplate away from the larger African landmass as a product of a geological "rifting" process. The physical separation caused many species to become physically separated, and several little springs appeared in the limestone and sandstone hills of western Sinai and southwestern Jordan, respectively (Hegazy and Doust, 2016). Moreover, the microhabitats of the deserts of Sinai Peninsula, Palestine, and Jordan support the rarest plant species (Lavergne, 2003).
Multivariate analysis indicated seven vegetation groups, two of them (groups I and II) distributed in the African countries, while the other five (groups III-VII) mainly distributed the Asian countries. This indicates that the topography governs the vegetation distribution. Friis et al. (2005) showed how topography affected plant diversity and distribution in the Horn of Africa, which in turn increased the number of species and endemism by facilitating evolutionary processes and avoiding extinction. According to Irl et al. (2015) and Méndez-Toribio et al. (2016), complex topography affects the richness, speciation process, diversity, and geographical distribution pattern of plant species, however, its influence is the greatest when determining plant endemic richness patterns. Also, due to enhanced habitat diversity and niche space availability, the varied topography may favor the creation of microrefugia (Badgley et al., 2017; Graae et al., 2018), which may have improved the chances of species survival during periods of shifts in the environment (Dobrowski, 2011; Keppel et al., 2012). In addition, topography promotes isolation, which acts as a dispersal restriction between populations (Janzen, 1967), limiting the flow of genes through isolation, promoting divergence, and creating the conditions necessary for allopatry (Coyne and Orr, 2004). In addition, numerous factors, such as climate, geographic barriers, human influences, biotic interactions, and stochastic occurrences, also have an impact on the distribution of endemic organisms (Morrone, 2018). Extreme weather events, such as drought and flooding, extremely high and low temperatures, wind events, and even changes in salinity or pH, are on the rise as a result of climate change. Therefore, it is likely that most organisms are going to face environmental conditions that are outside of their normal historical tolerance range (Cheng et al., 2017).
Some taxa found in the Saharo-Arabian floristic region extend to neighboring areas in other floristic regions because of some factors such as wind, human activities, and climate change. These taxa were excluded from endemism in the present study because of their extension to adjacent regions. For example, Allium crameri Asch. & Boiss. and Allium papillare Boiss that extend to the Mediterranean coast of Rafah (El-Garf, 2000) and Bardawil Lake (El-Bana, 2009; Nada et al., 2013); and Anthemis melampodina subsp. deserti (Boiss.) Eig, Anthemis zoharyana Eig, Asphodelus viscidulus Boiss., Astragalus camelorum Barbey, Astragalus palaestinus Eig, Bassia arabica (Boiss.) Maire & Weiller, Bellevalia eigii Feinbrun, Carduus getulus Pomel, Caroxylon tetrandrum (Forssk.) Akhani & Roalson, Carthamus eriocephalus (Boiss.) Greuter, Colchicum ritchii R. Br., Ferula sinaica Boiss., and Glaucium arabicum extend to the Mediterranean region including different adjacent countries (Täckholm, 1974; Squires and Ayoub, 1994; Boulos, 1999-2005; Turk and El Shayeb, 2005; Turki et al., 2006; Galal and Fawzy, 2007; El-Zanaty et al., 2010; Danin and Fragman-Sapir, 2016; Fakhry and Al-Anazi, 2017; Osman and Abdein, 2019; Mahklouf, 2020). In addition, some species extend to the Irano-Turanian region such as Caroxylon villosum (Schult.) Akhani & Roalson (Mosallam, 2007), Onopordum ambiguum Fresen. (Rabei and Elgamal, 2021), Verbascum eremobium Murb. and Blepharis attenuata Napper (Danin and Fragman-Sapir, 2016), Astragalus bombycinus Boiss. (Asri, 2004; Osman et al., 2014), and Stipagrostis multinerva H. Scholz (GBIF, 2023). Moreover, some species extend to the Sudano-Zambezian region like Ziziphus nummularia (Burm. f.) Wight & Arn. (Danin and Fragman-Sapir, 2016), Cometes surattensis Burm. f. (Al-Sherif et al., 2013), and Deverra triradiata Hochst. ex Boiss. (Abd El-Ghani et al., 2011).
The distribution of species, genetic diversity, and effective population size can all be impacted by human activity (Zhao et al., 2018, 2019). In many parts of the world, human activities have overtaken climate change in recent centuries as the primary factor influencing species distributions (Li et al., 2015). Anthropogenic disturbances such as livestock grazing and logging have an impact in changing the richness, diversity, distribution pattern, and vegetation structure in an ecosystem. The magnitude of their influence depends on the severity of disturbance (Hogan et al., 2016). Changes in climatic conditions are impacting the distribution and occurrence of natural resources in deserts. Both the timing and the routes of species migrations are predicted to change as a result of altered food distribution and the spatiotemporal unpredictability of water availability, which could adversely affect the dynamics of migratory ungulate populations (Duncan et al., 2012). It is widely acknowledged that human activities play a significant role in dryland climate change. Salinization, desertification, loss of vegetative cover, biodiversity loss, and other forms of environmental degradation are all caused in part by human activities. With rapid economic development, the modern lifestyle has become increasingly reliant on the use of fossil fuels (Barnett and O'Neill, 2010). In addition, the spatial patterns of energy consumption have strongly influenced temperature trends in recent years (Li and Zhao, 2012), especially in developing countries.

5 Conclusions

The current study is considered the pioneer in providing recent and verified checklist of the Saharo-Arabian endemic species in Egypt. A preliminary list of 429 Saharo-Arabian endemics in Egypt was compiled from three available literature. By comparing with three previous studies, this study excluded the species that were recorded in any countries or region outside the Saharo-Arabian region based on different literature, database reviews, and websites in Egypt and other countries, and finally reduced this number to 126 endemics belonging to 87 genera and 37 families. The eudicots had the highest plant diversity including 33 families, 82 genera, and 103 species, the monocots included 3 families, 4 genera, and 6 species, and there was also 1 species of gymnosperm. The most representative family was Asteraceae (17 species). Among the nine phytogeographic regions in Egypt, the South Sinai was the richest region with the Saharo-Arabian endemic species. Moreover, the SS1 subzone had all the 126 endemics, the SS2 subzone owned 79 taxa, and the SS3 subzone distributed 14 endemic plants. Seven groups were recognized at the fourth level of classification as a result of the application of TWINSPAN to the Saharo-Arabian endemics in Egypt, including Asphodelus refractus group that involved Tunisia, Algeria, Morocco, and Mauritania; Agathophora alopecuroides var. papillosa group that included only Libya; Anvillea garcinii group that included only Yemen; Reseda muricata group that involved Egypt, Saudi Arabia, Palestine, and Jordan; Agathophora alopecuroides var. alopecuroides group that included Iran, Iraq, Qatar, and Kuwait; Scrophularia deserti group that involved Pakistan, India, and Syria; and Astragalus schimperi group that involved Afghanistan, Oman, the United Arab Emirates, and Bahrain. The majority of the endemic plants identified in this study are very rare, emphasizing the need for comprehensive planning and conservation efforts, both in situ and ex situ.

Conflict of interest

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

Acknowledgements

We are grateful to the anonymous reviewers and editors for their insightful comments and suggestions in improving this manuscript.

Author contributions

Conceptualization and design: Asmaa S ABO HATAB, Kamal H SHALTOUT, Yassin M AL-SODANY, Mohamed M EL-KHALAFY; Material preparation, data collection, and analysis: Asmaa S ABO HATAB, Mohamed M EL-KHALAFY; Writing - original draft preparation: Asmaa S ABO HATAB, Kamal H SHALTOUT, Yassin M AL-SODANY, Mohamed M EL-KHALAFY; Writing - review and editing: Asmaa S ABO HATAB, Kamal H SHALTOUT, Yassin M AL-SODANY, Soliman A HAROUN, Mohamed M EL-KHALAFY; Supervision: Kamal H SHALTOUT, Yassin M AL-SODANY, Soliman A HAROUN. All authors approved the manuscript.

Appendix

Table S1 Updated list of the Saharo-Arabian endemic plants in Egypt
Table S2 Distribution of the Saharo-Arabian endemic plants in the Saharo-Arabian countries
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