35 Key words: phytosociology, correspondence factorial analysis, cartography, syntaxa, Morocco. Ključne besede: fitocenologija, korespondenčna faktorska analiza, kartografija, sintaksoni, Maroko. Corresponding author: Nadia Belahbib E-mail: nadia.belahbib@uit.ac.ma Received: 26. 1. 2023 Accepted: 15. 8. 2023 Phytosociological analysis of the Sidi Boughaba Biological Reserve, Kénitra, Morocco Abstract The biological reserve of Sidi Boughaba has a unique plant biodiversity, with many animals, including birds, finding refuge there. Data on this biodiversity is old and incomplete. This phytosociological study was carried out to define the vegetation structure of the area. Stratified sampling was carried out and 124 relevés were recorded. Multivariate statistical analyses (correspondence factorial analysis and hierarchical ascending classification) used to process the relevés revealed the presence of 9 plant groups, including 4 plant associations and 5 subassociations, which belong to 3 phytosociological classes (Ammophiletea, Quercetea ilicis and Querco roboris-Fagetea sylvaticae). Quercetea ilicis is the most widespread in the area, showing several successional stages on grey and consolidated dunes. Vegetation of Ammophiletea can only be found on mobile sand. Vegetation of the class Querco roboris-Fagetea sylvaticae occupies an inter-dunar depression on hydromorphic soil. The association Pterido aquilinum-Populetum albae is newly described as five subassociations of Clematido cirrhosae-Juniperetum lyciae. Izvleček V rezervatu Sidi Boughaba se nahaja edinstvena diverziteta rastlin s številnimi živalskimi vrstami, predvsem pticami, ki v njem najdejo zatičišče. Podatki o tej raznolikosti so stari in nepopolni. Da bi opredelili strukturo vegetacije v tem območju, smo izvedli fitocenološko raziskavo s stratificiranim vzorčenjem in naredili 124 vegetacijskih popisov. Z multivariatno statistiko (korespondenčna faktorialna analiza in hierarhična klasifikacija) smo popise razdelili v 9 skupin, od tega v 4 asociacije in 5 subasociacij, ki jih uvrščamo v tri razrede (Ammophiletea, Quercetea ilicis in Querco roboris-Fagetea sylvaticae). V rezervatu je najbolj razširjena vegetacija razreda Quercetea ilicis s številnimi sukcesijskimi stadiji na sivih in utrjenih sipinah. Vegetacijo razreda Ammophiletea najdemo na mobilnem pesku, razred Querco roboris-Fagetea sylvaticae pa se pojavlja v depresijah med dinami na hidromorfnih tleh. Opisali smo novo asociacijo Pterido aquilinum-Populetum albae ter 5 subasociacij asociacije Clematido cirrhosae-Juniperetum lyciae. Dounia Challi1, Jamila Dahmani2, El Habib Jdi3 & Nadia Belahbib4 1 Laboratory of Plant and Animal Production and the Agro-industry, Faculty of Sciences, Ibn Tofail University, Kénitra, Morocco 2 Laboratory of Plants, Animals Productions and Agro-industry, Faculty of Sciences, Ibn Tofail University, Kénitra, Morocco. 3 High Commission for Water and Forests and the Fight against Desertification, Provincial Directorate, Kénitra, Morocco. 4 Laboratory of Plants, Animals Productions and Agro-industry, Faculty of Sciences, Ibn Tofail University, Kénitra, Morocco. DOI: 10.2478/hacq-2023-000223/1 • 2024, 35–68 23/1 • 2024, 35–68 36 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco Introduction The Sidi Boughaba Biological Reserve is a wetland lo- cated on the Atlantic coast in western Morocco. It is a refuge for fauna, in particular birds, which find a welcom- ing environment in the humid atmosphere and the rich and varied vegetation. The faunal diversity is estimated at 205 species, including 37 sedentary species, 34 migra- tory species that winter regularly, 13 nesting species and 8 non-breeding species (Cherkaoui et al., 2013). Vegetation of the region was studied by Atbib (1983) and Jdi et al. (1987), while the most recent study on floristic diversity mentions 221 taxa (Bathaoui, 2001); since then, no veg- etation investigations have been carried out in the area. A synthesis of climatic data for a period of 34 years (1982– 2016) compared to syntheses of the periods 1948–1970 (Atbib, 1983) and 1957–1981 (Jdi et al., 1987) showed that the bioclimatic stage of the area has migrated from subhumid with temperate winters to semi-arid with cool winters (Challi et al., 2021). Following this migration, the area is now at significant risk of erosion, which would have an impact on the reserve's habitats. These habitats of- fer a high level of biodiversity, sheltering animal and plant species, including rare or endangered species. Studying this vegetation is important in that it helps managers to draw up appropriate management programs. Indeed, the landscape around the reserve has undergone major chang- es, with urban growth and coastal development. There is a lot of tourist traffic within the reserve, and visitor ac- tivities are not always environmentally friendly. That said, red juniper forest is well developed on the consolidated dunes and seems to harbor a considerable range of flora. Similarly, the hygrophilous vegetation around the “merja” (=lake) is remarkable for its diversity. It is therefore useful to study this biodiversity to see to what extent the change in climate, protection measures and the change in the sur- rounding landscape have impacted on the vegetation. The aim of this study was to define the plant groups found in the reserve and to compare them with previous descriptions and summaries (Atbib, 1983; Jdi et al., 1987). Materials and Methods Study area The Sidi Boughaba Biological Reserve is a protected area, included in the Ramsar list since 1980. It is therefore part of a complex of wetlands in Morocco, giving it interna- tional importance as an ornithological site. It is located in the south of the province of Kenitra and is oriented NNE-SSW. It extends between 34°12' N and 6°42' W on the Atlantic coast in northwest Morocco (Figure 1) and is bounded to the north by the mouth of the Oued Sebou. The area covers 650 ha, 450 of which are covered by veg- etation and 200 by “merja” (= lake). To the west, white dunes are made up of loose limestone sand; grey dunes of the Quaternary run parallel to them to the east and then, further inland, are slightly higher consolidated sandstone dunes of the Middle Quaternary. The climate of the region is Mediterranean with an oceanic influence. The average annual rainfall in Kéni- tra is 544.5 mm, with an average maximum temperature of 38.9°C in the hottest month (August) and an aver- age minimum temperature of 1.5°C in the coldest month (January) (Challi et al., 2021). The main soil types are sandy in the majority of the study area. They are classified as (i) raw mineral soils on unconsolidated white dunes, (ii) calco-mesimorphic soils of eolian origin and sandy texture on deep sand and dune sandstone, (iii) sandy-clay Mediterranean red soils, not very humus-rich, on slopes, superficial on dune sandstone and very deep in the inter- dune depression and (iv) hygromorphic soils correspond- ing to the water table, with sandy horizons of varying depths (Aubert, 1965; Duchauffour, 1982). Sampling In the spring of 2017, 20 homogeneous habitat types were determined on the basis of different physiogno- mies. A total of 124 relevés were taken from these habi- tat types, considering the different plant layers (Table 1). The number of relevés per habitat type was variable and depended on the floristic heterogeneity and the area ex- plored (Figure 2). The phytosociological study was conducted according to the Braun-Blanquet (1964) approach. The relevés were re- corded with a standard plot size of 100 m², considered suf- ficiently representative in Mediterranean plant formations (Fennane, 2003). For each relevé, GPS coordinates with latitude and longitude, altitude, slope and exposure were noted. Sampling was carried out during three consecutive years: June 2017, April and May 2018 and June 2019. A summary table presents the relative frequencies (F %) of the different species inventoried in the reserve (Table 2), with F % = (number of relevés in which the species is present / Total relevés)*100; this information was used in the numerical analyses (Meinard & Thébaud, 2019). The nomenclature of the plant communities iden- tified follows Benabid (2000) and new communities are described according to Theurillat et al. (2021). The plant material collected was identified in the labo- ratory based on the three volumes of the Practical Flora of Morocco (Fennane et al., 1999; Fennane et al., 2007; Fennane et al., 2014). 23/1 • 2024, 35–68 37 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco Habitat types (Ht) Number of relevés Area (ha) Lambert coordinates (Km) Geographical coordinates X (Km) Y (Km) Longitude Latitude Ht1 29 110.30 381.08 403.24 -6.6909 W 34.2245 N Ht2 9 38.29 381.74 404.56 -6.6839 W 34.2365 N Ht3 7 35.94 381.27 402.99 -6.6888 W 34.2222 N Ht4 20 172.59 382.30 405.12 -6.6779 W 34.2416 N Ht5 4 22.37 383.26 406.62 -6.6677 W 34.2552 N Ht6 9 21.33 380.81 401.38 -6.6936 W 34.2077 N Ht7 1 9.95 380.98 401.30 -6.6917 W 34.2069 N Ht8 3 10.80 381.13 401.31 -6.6902 W 34.2071 N Ht9 3 20.70 381.50 402.58 -6.6863 W 34.2186 N Ht10 1 10.03 381.36 402.08 -6.6877 W 34.2141 N Ht11 19 71.99 382.67 404.29 -6.6738 W 34.2341 N Ht12 1 4.63 381.64 402.65 -6.6847 W 34.2192 N Ht13 4 24.53 382.38 404.12 -6.6769 W 34.2326 N Ht14 2 2.88 383.71 406.63 -6.6628 W 34.2553 N Ht15 2 3.80 382.88 404.96 -6.6716 W 34.2402 N Ht16 1 0.62 383.31 406.32 -6.6671 W 34.2525 N Ht17 2 4.81 381.43 401.91 -6.6869 W 34.2125 N Ht18 5 9.73 383.17 405.66 -6.6686 W 34.2465 N Ht19 1 2.70 383.53 406.03 -6.6647 W 34.2499 N Ht20 1 2.95 383.81 406.65 -6.6618 W 34.2556 N Figure 1: Location map of the Sidi Boughaba Biological Reserve. Slika 1: Lokacija rezervata Sidi Boughaba. Table 1: Geographical location and area of the 20 habitat types studied. Tabela 1: Lokacije in površina 20 preučevanih habitatnih tipov.                                                                                                             23/1 • 2024, 35–68 38 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco Data analysis A Digital Terrain Model (DTM) provides complete topo- graphic and altimetric information (values of slopes, alti- tudes). It is therefore possible to draw up thematic maps based on the processing of relief information alone, or its combination with data of a different nature. The data collected and ordered in altitude classes thus allowed the elaboration of a hypsometric map of the study area (Fig- ure 3). This gave a synoptic view of the terrain and infor- mation on its geomorphology. The various maps (hypso- metric, slopes) were produced by ArcGIS 10.3 software (ESRI, 2015) using the DTM of the Sidi Boughaba Figure 2: Location of the habitat types (black line limits) based on the homogeneous physiognomy of the plant communities and location of the relevés (red points). Slika 2: Karta habitatnih tipov (omejeni s črno črto) na osnovi homogene fizionomije rastlinskih združb in lokacije vegetacijskih popisov (rdeče točke). Biological Reserve. The DTM was also used to establish a slope map using the (create slope & aspect) function; each point on the slope map corresponds to the value of the slope on the ground. The map thus produced allowed us to identify slope classes that provide information on the slope of the area (Figure 4). Using IBM-SPSS (IBM Corp., 2010) statistics soft- ware, statistical analyses of the collected floristic data were then performed. The processing of all the data was carried out after transforming the Braun-Blanquet abun- dance-dominance coefficient index into ordinal values in accordance with the suggestion of van der Maarel (1979), by replacing the + code with the value 0.5 and the r code with the value 0.1.                                                                                              23/1 • 2024, 35–68 39 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco We used a four-letter code for the species (for example: DJPH where D=Dounia and JPH = Juniperus phoenicea). Correspondence factor analysis was followed systemati- cally, with a hierarchical ascending classification whereby the data processed corresponded to a matrix of 124 rel- evés and 270 species. Species occurring only once were excluded from the analyses. Results and Discussion Data on the elevation and slope of the study area were collected and processed by ArcGis 10.3 software and used to produce the maps (Figure 3). The altitude varied from 6 to 77 m and the slope from 0 to almost 48%. During vegetation sampling, we recorded 270 species belonging to 199 genera and 58 families, the most repre- sented of which were Fabaceae and Asteraceae. Only 38 taxa had a frequency higher than 16% (Table 2) whereby Juniperus phoenicea and Retama monosperma were the most frequent, with a frequency of nearly 65% and 56%, respectively (Table 2). Species P F (%) Species P F (%) Juniperus phoenicea ssp. lycia 80 64.5 Arctotheca calendula 29 23.4 Retama monosperma 69 55.6 Lotus corniculatus 28 22.6 Galactites elegans 52 41.9 Silene gallica 26 20.9 Ammi majus 45 36.3 Ononis spinosa 25 20.2 Rumex bucephalophorus 44 35.5 Solanum sodomaeum 25 20.2 Ephedra fragilis 42 33.9 Prasium majus 25 20.2 Anagallis arvensis 42 33.9 Ammophila arenaria 24 19.3 Fumaria ouazzanensis 41 33.1 Torilis nodosa 23 18.5 Clematis cirrhosa 38 30.6 Mercurialis annua 23 18.5 Olea europaea 35 28.2 Achyranthes sicula 23 18.5 Malva sylvestris 35 28.2 Plantago lagopus 23 18.5 Phillyrea angustifolia 33 26.6 Urtica membranacea 23 18.5 Cerinthe major 33 26.6 Arisarum vulgare 22 17.7 Hordeum murinum 32 25.8 Papaver rhoeas 21 16.9 Emex spinosa 32 25.8 Parietaria mauritanica 20 16.1 Daucus carota 32 25.8 Asparagus aphyllus 20 16.1 Lagurus ovatus 30 24.2 Paronychia argentea 20 16.1 Pistacia lentiscus 29 23.4 Smyrnium olusatrum 20 16.1 Chamaerops humilis 29 23.4 Sonchus oleraceus 19 15.3 Figure 3: Maps of slope (a) and elevation (b) classes. Slika 3: Karta razredov naklona (a) in nadmorske višine (b). Table 2: List of the most frequent species in the Sidi Boughaba Biological Reserve (P: number of relevés in which the species was present; F (%): Frequency). Tabela 2: Seznam najbolj pogostih vrst v rezervatu Sidi Boughaba (P: število popisov, v katerih je bila vrsta prisotna; F (%): frekvenca).                                                                                                                                     23/1 • 2024, 35–68 40 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco Floristic analysis by correspondence factorial analysis The results of correspondence factorial analysis of all the relevés and the most important species recorded across the study area are shown in Table 3, in which we have taken the first two factorial axes, 1 and 2. Ammophila arenaria clearly stands out. This group belongs to the plant formations of the white sand dunes. Another group, characterized by Populus alba and Pteridium aquili- num, appears to be strongly linked to axis 1. These relevés were recorded on unconsolidated and consolidated fixed sand dunes. The third group that stands out is found in the habitat delimited on the left side of axis 1 and axis 2; it was observed on unconsolidated and consolidated fixed sand dunes. The last group, dominated by Solanum sodo- maeum and Retama monosperma, was developed on the edges of the juniper forest, near cultivated areas. C om po ne nt Initial eigenvalues Extraction sum of squares of selected factors Total % of variance % cumulated Total % of variance % cumulated 1 11.169 7.300 7.300 11.169 7.300 7.300 2 9.165 5.990 13.290 9.165 5.990 13.290 Table 3: Eigenvalues and percentages of inertia of the first factorial axes of correspondence factorial analysis. Tabela 3: Graf lastnih vrednosti in odstotki inercije prvih faktorskih osi v korespodnenčni faktorski analizi. Eigenvalues The eigenvalues and inertia rates are relatively high for the first axis and become low and almost constant from the eighth axis onwards (Figure 4). The first two axes explain 13.29% of the variation. Figure 4: Eigenvalue plot (152 species/124 relevés) at the Sidi Boughaba Biological Reserve. Slika 4: Lastne vrednosti (152 vrst/124 popisov) v rezervatu Sidi Boughaba. Meaning of the factorial axes The correspondence factorial analysis allowed four clusters to be distinguished (Figure 5). In the habitat delimited by the right side of axis 1, a group of Euphorbia paralias and Figure 5: Factorial plan of the F1 and F2 axes of the correspondence factorial analysis of the relevés in the Sidi Boughaba Biological Reserve (152 species/124 relevés). Slika 5: Faktorski diagram osi F1 in F2 korespondenčne faktorske analize popisov iz rezervata Sidi Boughaba (152 vrst/124 popisov). Legend (Legenda): EpAa Euphorbio paralie-Ammophiletum arenariae, CcJl- Clematido cirrhosae-Juniperetum lyciae, SsRm Solano sodomaeum- Retametum monospermae, PaPa- Pteridio aquilinum-Populetum albae. In relation to axis 1, the species on the right side form two closely related groups: group I formed by Euphorbia paralias and Ammophila arenaria occupying the mobile white dunes and group II characterised by Populus alba and Pteridium aquilinum from the bottom of the inter- dunal depression. On the negative side, group III was characterised by Juniperus phoenicea ssp. lycia and Clema- tis cirrhosa and was located on consolidated grey dunes, while group IV, characterised by Solanum sodomaeum and Retama monosperma, is far from this axis and is much more linked to axis 2. It seems that axis 1 represents the soil type factor, since group I is made up of psammophilous species, group II of hydrophilous species and group III of matorral species. In relation to axis 2, group IV, dominated by Retama mono- sperma and nitrophilous species, indicates that factor 2 corresponds to anthropization.                                                             23/1 • 2024, 35–68 41 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco Determination of phytosociological groups The bottom-up hierarchical clustering method applied to the 124 relevés provided a hierarchical tree (Figure 6). The shape of the graph allowed us to divide the relevés into nine groups. The nine groups are also presented in the phytoso- ciological tables (Tables 4–12). The factorial study of the vegetation of the Sidi Boughaba Biological Reserve allowed us to distinguish four plant communities (Fig- ure 5), which we further divided into 9 units or plant groups by hierarchical classification (Figure 6). panied by Pteridium aquilinum and other hygrophilous species. The two main factors that determine the distribu- tion of vegetation in the study area are the type of soil and anthropization; the first factor is predominant, being the result of zonation from the sea, while the second factor has only recently begun to take hold. Our synthesis thus allowed the distinction of 9 plant groups. Group 1 (Ammophila arenaria, Euphorbia paralias) (Table 4) This plant community corresponds to the first vegeta- tion belt (Habitat type 1), which appears on the mobile dunes. It contains perennial pioneer vegetation of the up- per beach to Polygonum maritimum, Medicago marina, Otanthus maritimus and Ipomoea stolonifera. It is subject to the effects of strong winds and settles on sandy min- eral substrates. A few specimens of Acacia cyanophylla and Acacia cyclops can still be observed at the eastern limit of the territory of this phytosociological group. These two species were originally introduced to fix the mobile dunes and are now gradually being eliminated by the ex- panding native vegetation. The stands are dominated by Ammophila arenaria and Euphorbia paralias. This made it possible to define the association Euphorbio paraliae- Ammophiletum arenariae, which belongs to Euphorbio paraliae-Ammophiletea australis Géhu & Géhu-Franck 1988 corr. Géhu in Bardat et al. 2004, Ammophiletalia australis BR.-Bl. 1933, Ammophilion arenariae (Tüxen in Br.-Bl. & Tüxen 1952) Géhu 1988. This association is found across Morocco in the same geographical areas as coastal red juniper formations (Benabid, 2000). Group 2 (Juniperus phoenicea, Clematis cirrhosa) (Table 5) The landscape in the study area is dominated by Junipe- rus phoenicea, which is the most widespread tree species. It occupies various topographic positions and is mainly associated with Clematis cirrhosa, Ephedra fragilis, Philly- rea angustifolia, Olea europaea, Retama monosperma and Chamaerops humilis. The plant community is located on the western slopes of the two consolidated dunes and on the northern parts of the second unconsolidated grey dune and can be found in places in the form of practically impenetrable groves. It is a climax community forming a plant association called Clematido cirrhosae-Juniperetum lyciae by Barbera, Quézel & Rivas-Martinez (1981), which belongs to the following higher phytosociological units: Quercetea ilicis Br.-Bl., 1947, Pistacio-Rhamnetalia alaterni Rivas-Martinez 1975 and Juniperion-lyciae Bar- bera, Quézel & Rivas-Martinez, 1981. Benabid (2000) describes this association as characteristic of maritime dunes in subhumid bioclimates and the thermo-mediter- ranean vegetation stage. Figure 6: Dendrogram of the hierarchical relevé classification. Slika 6: Dendrogram hierarhične klasifikacije popisov. Discussion The vegetation of the study area consists of three main groups of plant communities. Above the tidal zone, on the mobile sand dunes, the vegetation of Ammophila arenarea, Euphorbia paralias and other psammophilous species, is sparse. The dune ridges that follow have in- creasingly consolidated substrates eastwards; they are col- onized by communities of species adapted to each type of environment, with the ultimate evolution of these com- munities leading to juniper forest. This is generally found in a matorralized state but it can tend towards a state of equilibrium on stabilized gray dunes where the soil is well developed. The interdune depressions are occupied by typical hygrophilous vegetation, Populus alba accom-                                                        23/1 • 2024, 35–68 42 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco Group 3 (Juniperus phoenicea, Clematis cirrhosa, Ephedra fragilis) (Table 6) It is a sparse tree formation with low cover. The species that characterize it are Juniperus phoenicea and Ephedra fra- gilis. It occupies the fixed sandy dunes forming the second vegetation belt after the tidal zone. The substrate is sandy on deep sand enriched with organic matter. This commu- nity can be described as a subassociation Clematido cir- rhosae-Juniperetum lyciae ephedretosum fragilis subass. nov.; it can only be found on completely fixed and stabilized dunes and can be considered to be close to a climax com- munity. Indeed, a balance would have been established between the vegetation and the environmental conditions because of a fence installed during the last 50 years. Group 4 (Juniperus phoenicea, Clematis cirrhosa, Phillyrea angustifolia) (Table 7) This phytosociological group, characterized by three species, is the subassociation Clematido cirrhosae-Junipere- tum lyciae phillyreetosum angustifoliae subass. nov. It oc- cupies fixed sandy dunes, forming the second vegetation belt, on substrates with a sandy texture and deep sand enriched with organic matter. Phillyrea angustifolia, being less demanding for organic matter and water than Pistacia lentiscus, has been able to occupy these moderately moist sandy textured soils, where it is relatively abundant in asso- ciation with red juniper, Retama monosperma and Ephedra fragilis. It is a species that is resistant to soil water deficit. The plant community is the result of a fire in 1973 that destroyed part of the juniper climax community that was developing on the fixed and unconsolidated grey dune. Thanks to its great resistance to fire, Phillyrea angustifolia was able to establish itself together with the remaining sparse stands of juniper and of some species characteristic of the climax community. The presence of Clematis cir- rhosa, even in the form of rare plants, testifies to the forest atmosphere of the Clematido cirrhosae-Juniperetum lyciae association. We propose the newly described subassocia- tion Clematido cirrhosae-Juniperetum lyciae phillyreetosum angustifoliae subass. nov. It is a regeneration community developed on dunes. Observation showed a weak pres- ence of Pistacia lentiscus with an abundance of less than 5% in the territory of the subassociation thus defined; before the fire, this species formed a degradation com- munity according to Benabid (1982). Group 5 (Juniperus phoenicea, Clematis cirrhosa, Olea europea) (Table 8) In this group, stands of oleaster are largely located on the west-facing slopes of the first consolidated dune and on the less humid mid-western slopes of the second con- solidated dune, on superficial soils and dune sandstone. Oleaster is more frequently mixed with red juniper on dune sandstones, while Phillyrea angustifolia is mixed with red juniper on deep sands. Olea europaea is a very plastic species, indifferent to the nature of the substrate (Atbib, 1983). This is the third subassociation, for which we propose the name Clematido cirrhosae-Juniperetum lyciae oleetosum europeae subass. nov. Olea europaea is relatively abundant, especially in the eastern zone (con- solidated dune) of the reserve. Large, well-developed and physiologically vital individuals of Olea europaea were ob- served, associated with Juniperus phoenicea, Retama mono- sperma, Pistacia lentiscus and Clematis cirrhosa, which is the characteristic climax species. Group 6 (Juniperus phoenicea, Clematis cirrhosa, Retama monosperma) (Table 9) This is the fourth subassociation, which we would call Clematido cirrhosae-Juniperetum lyciae retametosum mono- spermae subass. nov.. Retama monosperma is a substrate- independent species, occurring on deep calcareous sands, deep sandy soils and superficial dune sandstones. This subassociation occupies the peripheral parts of the north- ern zone and the central part of the association Clematido cirrhosae-Juniperetum lyciae. It is an old plant community with aged and decaying specimens of Juniperus phoenicea. Clematis cirrhosa is still present with a high abundance- dominance ranging from 20% to 45%. It is a plant com- munity strongly invaded by Retama monosperma, whose cover can reach 100%. This phytosociological group therefore represents the beginning of matorralisation of plant formation. This is an evolutionary state of the vege- tation marked by sparsely-spaced, poorly-developed trees, and is found in areas with a Mediterranean climate. Group 7 (Juniperus phoenicea, Retama monosperma, Chamaerops humilis) (Table 10) This plant community is presented as the fifth subas- sociation, with the name Clematido cirrhosae-Juniperetum lyciae chamaeropetosum humilis subass. nov. It is a highly degraded phytosociological group that is beginning to be infiltrated by Chamaerops humilis, with a tree cover of no more than 25% and a shrub cover of no more than 50%. It derives from the Juniperus phoenicea, Clematis cirrhosa and Retama monosperma phytosociological group, and occu- pies the top of the dune system in a continuous strip from north to south, marking the transition and limit between the Juniperus phoenicea formation and the crops to the east. It is also found throughout most of the interdune furrow, on deep, dry, sandy soils. The species observed in the study area are characteristic of Quercetea ilicis; thermophilic ele- ments such as Olea europaea, Chamaerops humilis, Pistacia lentiscus, Juniperus phoenicea and Retama monosperma. 23/1 • 2024, 35–68 43 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco                                                               ­  € ­                                                                                                          Group 8 (Retama monosperma, Chamaerops humilis, Solanum sodomeum) (Table 11) This plant community is a highly degraded state of Solano-Retametum, with a total absence of Juniperus phoen- icea. This matorral, with Retama monosperma, can therefore be considered to be a plant association for which the name Solano sodomaeum-Retametum monospermae is proposed. In fact, this association has already been described in an internal report by Jdi et al. (1987) and is validly published for the first time in this work. It mainly occupies the wide furrow separating the two consolidated dune systems, the northern part of the “merja”, the transition zone between the juniper formation and the cultivated land to the east and an extensive area to the south of the reserve. It belongs to the same syntaxa as Clematido cirrhosae-Juniperetum ly- ciae, since it occupies its territory and derives from it by extensive degradation. The absence of Juniperus phoenicea and the low representation of the characteristic climax spe- cies, replaced by annual and nitrophilous plants, are the result of a strong anthropization of the environment. Figure 7: Vegetation map of plant communities in the Sidi Boughaba Biological Reserve. Slika 7: Vegetacijska karta rastlinskih združb v rezervatu Sidi Boughaba. 23/1 • 2024, 35–68 44 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco Group 9 (Populus alba, Pteridium aquilinum) (Table 12) This is a hygrophilous plant community and is linked to the hydromorphic soil type at the south-eastern end of the “merja”. This arboreal vegetation can be distinguished in the study area by its color and density of leaves, which varies according to the season, thus bringing variability to the landscape. The phytosociological group observed in the study area would therefore be a plant association designated Pteridio aquilinum-Populetum albae ass. nov. Populetalia albae represents hygrophilic communities, the most characteristic of which are Phillyrea angusti- folia, Populus alba, Juncus acutus, Pteridium aquilinum, Adiantum capillus-veneris and other subordinate taxa. The nine plant communities are classified as follows in the syntaxonomical system: Ammophiletea Br.-Bl. et Tx. ex Westhoff et al. 1946 Ammophiletalia Ammophilion arenariae (Tüxen in Br.-Bl. & Tüxen 1952) Géhu 1988 1. Euphorbio paraliae-Ammophiletum arenariae Géhu & Géhu-Franck 1988 corr. Géhu 2004 Quercetea ilicis Br.-Bl. 1947 Pistacio-Rhamnetalia alaterni Rivas-Martinez 1975 Juniperion-lyciae, Barbero, Ouézel and Rivas-Mar- tinez 1981 2. Clematido cirrhosae-Juniperetum lyciae Barbero, Ouézel & Rivas-Martinez 1981 2a. Clematido cirrhosae-Juniperetum lyciae typicum 2b. Clematido cirrhosae-Juniperetum lyciae ephedretosum fragilis subass. nov. hoc loco Holotypus: Table 6, relevé 96 2c. Clematido cirrhosae-Juniperetum lyciae phillyreetosum angustifoliae subass. nov. hoc loco Holotypus: Table 7, relevé 84 2d. Clematido cirrhosae-Juniperetum lyciae oleeto- sum europeae subass. nov. hoc loco Holotypus: Table 8, relevé 34 2e. Clematido cirrhosae-Juniperetum lyciae reta- metosum monospermae subass. nov. hoc loco Holotypus: Table 9, relevé 89 2f. Clematido cirrhosae-Juniperetum lyciae chamaeropetosum humilis subass. nov. hoc loco Holotypus: Table 10, relevé 38 3. Solano sodomaeum-Retametum monospermae Jdi in Challi et al. ass. nov. Holotypus: Table 11, relevé 33 Querco roboris-Fagetea sylvaticae Br.-Bl. & J. Vlieger in J. Vlieger 1937 Populetalia albae Br.-Bl. ex Tchou 1948 Populion albae Braun-Blanq. ex Tchou 1949 4. Pterido aquilinum-Populetum albae ass. nov. hoc loco Holotypus: Table 12, relevé 74 The study carried out in the Sidi Bougnaba biological reserve made it possible to define plant groups, to specify their phytosociological affiliation and to delimit their area of distribution thanks to the geographical coordinates taken for each relevé. This allowed the creation of a veg- etation map (Figure 7) that reflects the ecological condi- tions of the environment, and several plant communities may belong to the same plant formation (physiognomy) and the same vegetation series. Conclusions The Sidi Boughaba Biological Reserve is a wetland that is distinguished by both its fauna and flora. Statistical analy- ses of the data collected by the numerical method based on correspondence factorial analysis and then on a hier- archical ascending classification, allowed the definition of nine phytosociological groups, including four associations and five subassociations. The association considered to be climactic in the area is Clematido cirrhosae-Juniperetum ly- ciae Barbero et al. 1981. Depending on the environmental conditions and the impact of anthropization, stages of deg- radation derived from the climax were established. They range from matorralised structures such as the subassocia- tions: Clematido cirrhosae-Juniperetum lyciae ephedretosum fragilis, Clematido cirrhosae-Juniperetum lyciae phillyreeto- sum angustifoliae, Clematido cirrhosae-Juniperetum lyciae oleetosum europeae, Clematido cirrhosae-Juniperetum lyciae retametosum monospermae and Clematido cirrhosae-Juni- peretum lyciae chamaeropetosum humilis to very degraded or even completely therophytised structures that are heav- ily infiltrated by nitrophilous species. The present study thus made it possible to propose five subassociations and one new association. The phytosociological groups thus defined belong to three phytosociological classes: Ammo- philetea, Quercetea ilicis and Querco roboris-Fagetea sylvati- cae. The results obtained show that the rich and remark- able biodiversity of the study area is strongly impacted by anthropozoic factors, even within the reserve. It is there- fore essential to safeguard this heritage. Dounia Challi  https://orcid.org/0009-0003-5173-1815 Jamila Dahmani  https://orcid.org/0000-0003-2716-1106 El Habib Jdi  https://orcid.org/0000-0001-5769-8292 Nadia Belahbib  https://orcid.org/0009-0001-9938-9479 23/1 • 2024, 35–68 45 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco References Atbib, M. (1977). Etude phyto-écologique de la réserve biologique de Mehdia (littoral atlantique, Maroc). [Thèse de 3éme cycle, Université des Sciences et Techniques de Montpellier]. Atbib, M. (1983). Phytoecological study of the biological reserve of Mehdia (Atlantic coast of Morocco) -2- The vegetation of the dune environment. Bulletin de l'Institut Scientifique, Rabat, suppl., 5, 73–133. Aubert, G. (1965). Classification des sols. Tableaux des classes, sous-classes, groups et sous-groupes de sols utilisés par la section de pédologie de l’O.R.S.T.O.M. Cahiers O.R.S.T.O.M. Pédol, Paris, 3(4), 89–90. Bathaoui, F. (2001). 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N or th Lo ng .W es t -6.6988 34.2084 -6.6841 34.2428 -6.6851 34.2401 -6.6830 34.2428 -6.6820 34.2473 -6.6817 34.2482 -6.6820 34.2464 -6.6776 34.2410 -6.6819 34.2437 -6.6830 34.2446 -6.6914 34.2238 -6.6882 34.2301 -6.6851 34.2392 -6.6893 34.2283 -6.6872 34.2319 -6.6872 34.2337 -6.6860 34.2356 Pr St at io n 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 El ev at io n (m ) 20 20 26 18 71 33 31 26 54 22 36 61 22 24 20 66 20 O rie nt at io n N W W W S N N N N N N N N N N N N N Sl op e (% ) 0 3 5 0 1 14 6 13 5 26 39 5 1 7 7 5 0 So il i i i i i i i i i i i i i i i i i Re le vé si ze (m ²) 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 D ia gn os ti c sp ec ie s o f t he a ss oc ia ti on   Am m op hi la a re na ria 3 3 4 4 1 2 3 3 4 3 4 2 3 3 3 4 4 17 Eu ph or bi a pa ra lia s 2 3 2 1 2 2 1 2 2 3 4 3 3 3 14 O ta nt hu s m ar iti m us 2 3 2 2 2 2 3 2 8 M ed ica go m ar in a 1 2 2 2 3 2 3 6 Po lyg on um m ar iti m um 4 3 3 2 2 2 6 Ip om oe a sto lo ni fer a 4 2 3 4 2 4 6 Lo tu s c re tic us 3 3 2 3 C ak ile m ar iti m a 3 1 2 3 Sa lso la k al i 1 2 2 Er yn gi um m ar iti m um 2 1 D iff er en ti al sp ec ie s Ly ciu m eu ro pe au m 1 1 Ep he dr a fra gi lis 2 1 Ta m ar ix g al lic a 2 1 H ier ac iu m to m en to su m + 1 G al ac tit es ele ga ns 3 1 2 Ri ec ha rd ia ti ng ita na 3 4 2 Ps eu do rla ya p um ila 1 1 2 So la nu m so do m eu m + 1 Ac ac ia cy an op hy lla 2 1 C ar po br ot us ed ul is                   2               1 23/1 • 2024, 35–68 47 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco Ta bl e 5 (T ab el a 5) : C lem at id o cir rh os ae -Ju ni pe re tu m ly cia e B ar be ro , O ué ze l a nd R iv as -M ar tin ez 1 98 1 R EL EV É N U M BE R R 80 R 82 R 65 R 10 0 R 59 R 21 R 61 R 47 R 48 R 70 R 58 R 66 R 69 R 63 R 64 R 68 R 51 R 52 Pr C oo rd in at e La t. N or th Lo ng . W es t -6.6911 34.2201 -6.6892 34.2229 -6.6743 34.2438 -6.6819 34.2446 -6.6701 34.2537 -6.6734 34.2358 -6.6712 34.2510 -6.6696 34.2429 -6.6700 34.2438 -6.6814 34.2358 -6.6701 34.2528 -6.6776 34.2428 -6.6760 34.2428 -6.6722 34.2492 -6.6744 34.2456 -6.6835 34.2322 -6.6711 34.2483 -6.6664 34.2502 Pr St at io n 3 3 4 1 4 13 4 11 18 4 4 4 4 4 4 4 4 4 El ev at io n (m ) 20 22 67 48 32 11 72 18 53 56 25 70 20 28 50 22 27 32 O rie nt at io n S W N W S S N E N S S S E N S N N N S S Sl op e (% ) 2 6 6 36 17 4 6 0 5 8 6 5 1 2 27 20 8 13 So il ii ii ii ii ii ii ii ii ii ii ii ii ii ii ii ii ii ii Re le vé si ze (m ²) 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 D ia gn os ti c sp ec ie s o f t he a ss oc ia ti on   Ju ni pe ru s p ho en ice a 2 2 4 2 2 3 3 2 2 3 3 3 2 3 1 3 2 4 18 C lem at is cir rh os a 3 2 3 2 2 3 3 3 4 4 3 3 4 4 4 3 3 4 18 Ru bi a pe re gr in a 1 1 2 2 4 3 7 Fu m ar ia o ua zz an en sis 2 3 2 3 5 O no ni s n at rix 3 2 3 3 4 Br yo ni a di oi ca 4 + 2 Sm ila x as pe ra 2 2 2 Li m on iu m si nu at um 1 1 C lem at is fla m m ul a 2 1 D iff er en ti al sp ec ie s o f t he a ss oc ia ti on   Ph ill yr ea a ng us tif ol ia 1 + + 1 2 1 2 + 2 3 10 Ep he dr a fra gi lis 1 1 1 + 2 1 + + + 9 Pr as iu m m aj us 2 4 3 3 3 4 3 3 9 O lea eu ro pa ea + 1 + 1 1 + + 1 8 An ag al is ar ve ns is 4 3 3 2 3 3 3 8 O no ni s s pi no sa 3 2 2 4 4 5 4 8 Re ta m a m on os pe rm a 2 2 2 + 2 2 1 7 Ru m ex b uc ep ha lo ph or us 4 2 5 1 5 2 5 7 Pa pa ve r r ho ea s 3 + 2 2 2 7 La th yr us cl ym en um 1 4 3 3 4 4 3 7 Ar isa ru m v ul ga re 3 3 3 2 3 3 6 Ja sio ne m on ta na 3 3 3 3 3 3 6 Pi sta cia le nt isc us + 2 1 + 1 5 Si len e g al lic a 1 3 3 3 5 C ha m ae ro ps h um ili s 2 1 2 2 4 23/1 • 2024, 35–68 48 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco R EL EV É N U M BE R R 80 R 82 R 65 R 10 0 R 59 R 21 R 61 R 47 R 48 R 70 R 58 R 66 R 69 R 63 R 64 R 68 R 51 R 52 Pr As pa ra gu s a lb us 2 2 1 2 4 G er an iu m p ur pu re um 1 2 3 2 4 Fe di a pa lle sce ns 1 2 2 2 4 G al iu m a pa rin e 2 1 2 3 4 La nt an a ca m ar a 2 4 2 As pa ra gu s a ph yll us 3 1 Ju nc us a cu tu s 3 1 Ve rb as cu m v irg at um 4 1 Ly ciu m eu ro pe au m + 1 Tu be ra ria g ut ta ta 2 1 Lo bu la ria m ar iti m a 4 1 Ta m ar ix g al lic a + 1 C or on ill a re pa nd a 2 1 G la di ol us co m m un is 3 1 Ar en ar ia m au rit an ica 3 1 O th er sp ec ie s C er in th e m aj or 3 1 3 3 3 3 3 3 3 9 La gu ru s o va tu s 5 1 4 3 2 3 5 3 9 G al ac tit es ele ga ns 3 1 2 3 3 1 3 3 8 H or de um m ur in um 5 5 5 5 4 5 7 Lo tu s c or ni cu la tu s 1 3 4 4 3 6 Pa rie ta ria m au rit an ica 5 2 3 3 1 6 Am m i m aj us 3 1 2 4 3 6 M al va sy lv est ris 3 3 4 1 2 6 D au cu s c ar ot a 3 2 2 3 3 2 6 Tr ifo liu m ca m pe str e 2 3 1 4 2 5 Ac hy ra nt he s s icu la 1 2 4 2 5 An isa nt ha st er ili s 5 4 5 3 5 Pl an ta go la go pu s 2 1 1 2 4 Ar cto th ec a ca len du la 3 3 3 4 4 H yp oc ha er is gl ab ra 3 4 4 4 C en tra nt hu s c al cit ra pa e 1 2 3 4 Sm yr ni um o lu sa tr um 1 2 + 2 4 C al en du la a rv en sis 2 2 4 4 4 Tr ise ta ria p an ice a 5 3 3 2 4 Lo liu m p er en ne 3 5 5 4 23/1 • 2024, 35–68 49 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco R EL EV É N U M BE R R 80 R 82 R 65 R 10 0 R 59 R 21 R 61 R 47 R 48 R 70 R 58 R 66 R 69 R 63 R 64 R 68 R 51 R 52 Pr To ril is no do sa 3 4 4 3 G er an iu m m ol le 3 2 3 Vi cia lu te a 1 1 2 3 C en ta ur ea p ul la ta 3 4 3 3 Vi cia sa tiv a 4 2 4 3 M ed ica go p ol ym or ph a 3 1 2 3 U rt ica m em br an ac ea 3 4 2 3 Pa re nt uc ell ia v isc os a 2 4 1 3 D ip ca di se ro tin um 3 4 2 3 Eu ph or bi a te rr ac in a 2 3 4 3 Ps eu do rla ya p um ila 1 1 1 3 Ph ag na lo n sa xa til e 1 1 4 3 M iso pa te s c al yc in um 1 3 2 Em ex sp in os a 2 4 2 Ev ax p yg m ae a 3 1 2 Pa ro ny ch ia a rg en te a 2 2 2 Vi cia v ill os a 3 2 2 Sc ro ph ul ar ia a ur icu la ta 2 3 2 U ro sp er m um p icr oi de s 2 2 2 Er ig er on fl or ib un du s 2 2 2 Li nu m st ric tu m 3 5 2 O xa lis p es- ca pr ae 3 1 2 As tra ga lu s b oi tic us 3 1 2 U rt ica u re ns 2 1 Bi scu te lla la ev ig at a 2 1 An ac yc lu s c la va tu s 2 1 U rt ica d io ica 2 1 Av en a ste ril is 1 1 M er cu ria lis a nn ua 1 1 Lu pi nu s a ng us tif ol iu s 3 1 As tra ga lu s h am os us 3 1 H yp oc ha er is ra di ca ta 4 1 D elp hi ni um p ub esc en s 4 1 Er od iu m ci co ni um 4 1 Tr ise tu m fl av esc en s 4 1 Ag ro py ro n ju nc eu m 3 1 23/1 • 2024, 35–68 50 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco R EL EV É N U M BE R R 80 R 82 R 65 R 10 0 R 59 R 21 R 61 R 47 R 48 R 70 R 58 R 66 R 69 R 63 R 64 R 68 R 51 R 52 Pr Ph leu m p ra te ns is 1 1 So nc hu s o ler ac eu s 3 1 Ec hi um p la nt ag in eu m 2 1 Ru m ex p ul ch er + 1 H ym en oc ar po s c irc in na tu s 2 1 U m bi lic us h or iz on ta lis + 1 Li na ria su pi na 3 1 Pl an ta go m ac ro rh iz a 3 1 Tr ifo liu m a ng us tif ol iu m 3 1 So nc hu s a sp er 2 1 So la nu m n ig ru m + 1 Pl an ta go la nc eo la ta 3 1 Ba rt sia tr ix ag o 2 1 As pa ra gu s s et ac eu s 2 1 Ro str ar ia cr ist at a 1 1 Po lyc ar po n te tra ph yll um 3 1 N on ea ca lyc in a 2 1 Fe ru la co m m un is 1 1 Lo tu s c re tic us 2 1 Ag av e a m er ica na + + 2 Ac ac ia cy an op hy lla + + 2 Eu ca lyp tu s g om ph oc ep ha la + + 2 O pu nt ia fi cu s i nd ica 3 1 Pi nu s s tro bu s + 1 Ac ac ia cy an op hy lla + + 1 Ac ac ia cy clo ps       +                             1 23/1 • 2024, 35–68 51 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco Table 6 (Tabela 6): Clematido cirrhosae-Juniperetum lyciae ephedretosum fragilis subass. nov. RELEVÉ NUMBER R103 R107 R41 R62 R112 R72 R96 R109 R123 R87 R113 R124 R104 Pr Coordinate Lat. North Long. West -6 .6 79 7 34 .2 47 2 -6 .6 84 3 34 .2 35 4 -6 .6 64 7 34 .2 50 2 -6 .6 72 3 34 .2 50 1 -6 .6 81 9 34 .2 42 8 -6 .6 75 6 34 .2 47 7 -6 .6 84 0 34 .2 41 0 -6 .6 80 9 34 .2 43 7 -6 .6 80 9 34 .2 44 6 -6 .6 89 7 34 .2 17 4 -6 .6 80 8 34 .2 42 8 -6 .6 86 1 34 .2 37 4 -6 .6 80 9 34 .2 45 5 Pr Station 2 2 11 4 2 4 1 2 2 4 2 1 2 Elevation (m) 61 21 27 72 57 60 18 24 32 29 62 18 20 Orientation N N N N N WS W N N W N N N Slope (%) 6 21 15 6 1 31 0 6 31 3,4 1 4 0 Soil ii ii ii ii ii Ii ii ii ii ii ii ii ii Relevé size (m²) 100 100 100 100 100 100 100 100 100 100 100 100 100 Diagnostic species of the association Clematido cirrhosae-Juniperetum lyciae   Juniperus phoenicea 2 3 1 1 2 3 1 3 1 1 2 1 3 13 Phillyrea angustifolia 1 2 1 + 1 3 2 8 Prasium majus 2 2 1 3 + 1 7 Retama monosperma + + 1 1 2 1 6 Fumaria ouazzanensis 1 4 1 2 5 Pistacia lentiscus 1 + + + 4 Lycium europeaum 1 2 1 3 Clematis cirrhosa 1 1 2 Bryonia dioica 1 2 Limonium sinuatum + 1 Osyris quadripartita 2 1 Differential species of the subassociation Clematido cirrhosae-Juniperetum lyciae ephedretosum fragilis subass. Ephedra fragilis 3 4 2 2 4 2 2 2 4 2 3 1 2 14 Ammophila arenaria + 1 Polygonum maritimum 2 1 Lotus creticus 2 1 Otanthus maritimus 2 1 Cakile maritima 2 1 Fedia pallescens 2 1 Other species Daucus carota 1 3 1 3 1 1 6 Galactites elegans 3 + 2 2 4 Anagalis arvensis 1 3 2 3 Ononis spinosa 3 4 1 3 Emex spinosa 4 1 2 3 Malva sylvestris 1 1 2 3 Cerinthe major 3 1 2 Arctotheca calendula 1 2 2 Plantago lagopus 2 3 2 Anacyclus radiatus 3 3 2 Lolium perenne 3 4 2 Papaver rhoeas 1 1 2 Paronychia argentea + 2 2 Chamaerops humilis + 1 2 Erodium cicutarium 3 2 2 Agropyron junceum 1 1 2 Urtica membranacea 3 3 2 Calendula arvensis 4 1 2 23/1 • 2024, 35–68 52 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco RELEVÉ NUMBER R103 R107 R41 R62 R112 R72 R96 R109 R123 R87 R113 R124 R104 Pr Trisetaria panicea 1 1 2 Euphorbia terracina 3 2 2 Arisarum vulgare 3 1 Sonchus asper 1 1 Plantago lanceolata 2 1 Pseudorlaya pumila 1 1 Oxalis pes-caprae 1 1 Salvia verbenaca + 1 Urtica dioica 1 1 Rumex bucephalophorus 1 1 Parietaria mauritanica 1 1 Silybum marianum 2 1 Centaurea pullata 2 1 Hordeum murinum 1 1 Silene gallica 3 1 Echium vulgare 2 1 Lupinus angustifolius + 1 Vicia sativa 1 1 Lotus corniculatus 1 1 Tolpis barbata 1 1 Hypochaeris radicata 1 1 Smyrnium olusatrum 1 1 Melilotus indicus 3 1 Trisetum flavescens 1 1 Parentucellia viscosa 2 1 Ammi majus 2 1 Linaria maroccana + 1 Sonchus tenerrimus 2 1 Anacyclus clavatus 2 1 Lantana camara 1 1 Jasione montana 1 1 Lathyrus clymenum 2 1 Armeria mauritanica 3 1 Anisantha tectorum 1 1 Logfia gallica 1 1 Asparagus setaceus 3 1 Phagnalon sexatile 2 1 Nonea calycina 2 1 Opuntia ficus indica 1 1 2 Agave americana 2 1 23/1 • 2024, 35–68 53 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco Table 7 (Tabela 7): Clematido cirrhosae-Juniperetum lyciae phillyreetosum angustifoliae subass. nov. RELEVÉ NUMBER R79 R83 R86 R39 R57 R67 R81 R72 R84 R85 R90 Pr Coordinate Lat. North Long. West -6 .6 93 5 34 .2 16 5 -6 .6 88 2 34 .2 23 8 -6 .6 88 6 34 .2 21 1 -6 .6 63 6 34 .2 52 0 -6 .6 73 4 34 .2 53 7 -6 .6 76 1 34 .2 41 0 -6 .6 90 3 34 .2 21 1 -6 .6 75 6 34 .2 47 7 -6 .6 88 2 34 .2 24 7 -6 .6 87 6 34 .2 22 5 -6 .6 80 9 34 .2 47 3 Station 1 3 3 11 4 4 3 4 3 3 1 Elevation (m) 29 21 20 24 33 46 26 60 20 20 18 Orientation S W W N N N W WS W W W Pr Slope (%) 12 7 0 25 11 5 16 31 0 1 0 Soil ii ii ii ii ii ii ii ii Ii ii ii Relevé size (m²) 100 100 100 100 100 100 100 100 100 100 100 Diagnostic species of the association Clematido cirrhosae-Juniperetum lyciae Juniperus phoenicea 2 2 2 2 3 3 2 2 + 2 1 11 Ephedra fragilis 1 + + + 1 1 1 2 + 9 Fumaria ouazzanensis 1 3 2 2 2 4 2 8 Bryonia dioica 2 2 2 3 1 + 7 Ononis natrix 2 2 3 2 3 3 7 Retama monosperma 1 1 + 2 4 Olea europaea 2 1 + 1 4 Pistacia lentiscus + 1 + 3 Daucus carota 1 2 1 3 Clematis cirrhosa 2 2 2 Limonium sinuatum + 1 Smilax aspera 2 1 Differential species of the subassociation phillyreetosum angustifoliae Phillyrea angustifolia 3 2 2 + 2 2 2 3 3 2 + 11 Prasium majus + 1 2 + 1 5 Lotus creticus 3 1 Polygonum maritimum 2 1 Ammophila arenaria + 1 Other species Galactites elegans 3 2 3 2 2 2 2 3 2 2 10 Cerinthe major 2 2 1 3 2 2 3 3 8 Lotus corniculatus 1 2 2 3 2 3 2 7 Emex spinosa 3 3 3 1 2 2 1 7 Rumex bucephalophorus 3 3 2 4 3 4 6 Ammi majus 2 2 1 4 2 2 6 Calendula arvensis 3 1 2 1 3 5 Plantago lagopus 2 1 1 1 2 5 Silene gallica 1 1 3 3 + 5 Medicago polymorpha 2 3 1 3 2 5 Arctotheca calendula 3 1 2 2 4 Ononis spinosa 2 2 4 3 4 Mercurialis annua 1 1 1 1 4 Asparagus albus 1 1 + 1 4 Hypochaeris glabra 1 2 3 3 4 Polypogon monspeliensis 2 1 4 5 4 Dipcadi serotinum 1 3 2 2 4 Oxalis pes-caprae 3 1 1 3 Arisarum vulgare 1 1 2 3 23/1 • 2024, 35–68 54 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco RELEVÉ NUMBER R79 R83 R86 R39 R57 R67 R81 R72 R84 R85 R90 Pr Hordeum murinum 1 1 4 3 Malva sylvestris 2 1 1 3 Lolium perenne 4 1 1 3 Urtica membranacea 3 1 2 3 Fedia pallescens 1 2 1 3 Pseudorlaya pumila 2 1 1 3 Vicia villosa 1 + 2 3 Anacyclus radiatus 1 3 1 3 Jasione montana + 1 1 3 Solanum sodomeum + + 2 Anagalis arvensis + 2 2 Smyrnium olusatrum + 2 2 Chamaerops humilis + 1 2 Asparagus aphyllus + + 2 Lathyrus clymenum 4 1 2 Evax pygmaea 1 1 2 Erodium cicutarium 1 2 2 Astragalus hamosus 2 1 2 Sonchus oleraceus 3 3 2 Rubia peregrina 1 3 2 Sonchus asper 3 3 2 Plantago lanceolata 3 3 2 Lagurus ovatus 1 1 Torilis nodosa 2 1 Parietaria mauritanica 2 1 Trifolium campestre 3 1 Achyranthes sicula 1 1 Lobularia maritima 1 1 Juncus acutus 3 1 Rostraria cristata + 1 Misopates calycinum 3 1 Scolymus hispanicus 1 1 Papaver rhoeas + 1 Geranium molle 3 1 Silybum marianum 2 1 Centaurea pullata + 1 Echium vulgare 2 1 Lycium europeaum 1 1 Hypochaeris radicata 1 1 Melilotus indicus 3 1 Trisetum flavescens 1 1 Echium plantagineum 3 1 Umbilicus horizontalis + 1 Sonchus tenerrimus 1 1 Ruscus hypophyllum + 1 Verbascum boerhavii + 1 Sedum rupestre 1 1 Urospermum picroides 1 1 Solanum nigrum 1 1 Ononis reclinata 2 1 23/1 • 2024, 35–68 55 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco RELEVÉ NUMBER R79 R83 R86 R39 R57 R67 R81 R72 R84 R85 R90 Pr Euphorbia terracina 2 1 Helianthemum salicifolium + 1 Gleditsia triacanthos 1 1 Phagnalon sexatile 2 1 Bromus hordeaceus 2 1 Nonea calycina 2 1 Ferula communis 1 1 Agave american 2 2 2 Opuntia ficus indica               1     1 2 Table 8 (Tabela 8): Clematido cirrhosae-Juniperetum lyciae oleetosum europeae subass. nov. RELEVE NUMBER R19 R20 R18 R22 R23 R34 R44 R45 R29 R50 Pr Coordinate Lat. North Long. West -6 .6 72 1 34 .2 37 5 -6 .6 72 0 34 .2 36 6 -6 .6 76 1 34 .2 35 2 -6 .6 74 8 34 .2 32 0 -6 .6 76 7 34 .2 29 2 -6 .6 73 2 34 .2 39 3 -6 .6 68 9 34 .2 42 9 -6 .6 69 1 34 .2 44 6 -6 .6 69 7 34 .2 40 2 -6 .6 69 0 34 .2 47 4 Station 13 11 11 11 11 11 11 11 11 18 Elevation (m) 17 13 30 16 17 15 20 20 63 68 Pr Orientation NW NW NW NW NW NW N N S S Slope (%) 5 1 0 13 13 6 9 1 8 6 Soil ii ii ii ii ii ii ii ii ii ii Relevé size (m²) 100 100 100 100 100 100 100 100 100 100 Diagnostic species of the association Clematido cirrhosae-Juniperetum lyciae   Juniperus phoenicea 3 3 1 3 3 3 2 2 1 2 10 Fumaria ouazzanensis 3 + 2 3 3 3 3 3 9 Retama monosperma 1 1 + 1 1 1 + 1 8 Clematis cirrhosa 1 3 3 3 2 3 2 7 Pistacia lentiscus 1 + + 3 + 4 1 7 Fedia pallescens + 1 2 1 4 Prasium majus 2 1 1 3 Limonium sinuatum + + 2 3 Ephedra fragilis + 1 Rubia peregrina 2 1 Bryonia dioica 1 1 Differential species of the subassociation oleetosum europeae   Olea europaea 3 3 5 3 4 4 3 4 3 2 10 Ammi majus 2 1 + 3 2 3 3 4 2 9 Cerinthe major 3 2 3 3 2 1 2 7 Asparagus albus 1 1 + + 1 1 6 Chamaerops humilis + 1 1 1 + 1 6 Asparagus aphyllus 2 3 1 3 Other species Achyranthes sicula + 2 2 1 2 + 2 2 1 9 Sonchus oleraceus + 3 1 4 2 3 3 2 8 Geranium molle + 1 3 2 2 1 6 Agropyron junceum 4 5 + 3 4 5 6 Lagurus ovatus + + 2 5 3 5 Malva sylvestris 1 1 2 3 2 5 23/1 • 2024, 35–68 56 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco RELEVE NUMBER R19 R20 R18 R22 R23 R34 R44 R45 R29 R50 Pr Torilis nodosa + + 2 2 4 Anagalis arvensis 3 3 3 3 4 Parietaria mauritanica 2 3 2 2 4 Ornithopus sativus + 1 + 2 4 Tolpis barbata + 2 3 1 4 Smyrnium olusatrum + 1 1 2 4 Urtica membranacea 3 3 2 3 4 Rumex bucephalophorus + 4 3 3 Misopates calycinum 3 1 1 3 Emex spinosa 2 3 3 3 Arisarum vulgare 1 2 2 3 Hordeum murinum 2 3 3 3 Cerastium glomeratum + + 3 3 Mercurialis annua 4 3 2 3 Vicia sativa + 2 2 3 Silene gallica + + 2 3 Phleum pratensis 3 r 3 3 Medicago orbicularis 1 1 2 3 Daucus carota 3 1 4 3 Ononis natrix 1 3 2 Avena sterilis 3 3 2 Galactites elegans 3 1 2 Tuberaria guttata 2 1 2 Campanula lusitanica + 2 2 Ruscus hypophyllum + 1 2 Briza major (maxima) 3 3 2 Geranium purpureum 3 1 2 Parentucellia viscosa + + 2 Calendula arvensis 3 1 Anacyclus radiatus 1 1 Urginea maritima + 1 Solanum sodomeum 3 1 Verbascum sinuatum 1 1 Evax pygmaea 1 1 Vicia lutea 3 1 Paronychia argentea 2 1 Erodium cicutarium 2 1 Hypochaeris radicata 2 1 Medicago polymorpha 3 1 Echium plantagineum + 1 Hypochaeris glabra 1 1 Sonchus arvensis 2 1 Ononis spinosa 3 1 Delphinium pubescens 1 1 Erodium ciconium 2 1 Vicia villosa 3 1 Umbilicus horizontalis 1 1 Lactuca virosa 1 1 Allium subhirsutum + 1 Lipunus micranthus + 1 23/1 • 2024, 35–68 57 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco RELEVE NUMBER R19 R20 R18 R22 R23 R34 R44 R45 R29 R50 Pr Vulpia geniculata 1 1 Anthoxanthum odorutum 2 1 Convolvulus sp. 1 1 Briza minor 2 1 Narcissus tazetta 1 1 Anacyclus clavatus 1 1 Heteranthemis viscidehirta 2 1 Lolium perenne 2 1 Sonchus asper 2 1 Bromus diandrus 3 1 Dactylis glomerata 1 1 Poa pratensis 2 1 Gladiolus italicus 2 1 Anisantha sterilis         2 1 Table 9 (Tabela 9): Clematido cirrhosae-Juniperetum lyciae retametosum monospermae subass. nov. RELEVÉ NUMBER R27 R34 R43 R44 R89 R35 R42 R54 R61 Pr Coordinate Lat. North Long. West -6 .6 71 0 34 .2 37 5 -6 .6 73 2 34 .2 39 3 -6 .6 66 8 34 .2 47 5 -6 .6 68 9 34 .2 42 9 -6 .6 66 9 34 .2 56 5 -6 .6 62 6 34 .2 56 5 -6 .6 65 7 34 .2 49 3 -6 .6 69 1 34 .2 53 7 -6 .6 71 2 34 .2 51 0 Station 11 11 11 11 5 14 11 4 4 Elevation (m) 16 15 31 20 29 17 29 64 72 Pr Orientation S NW N N E N N S N Slope (%) 11 6 7 9 9 6 13 2 6 Soil ii ii ii ii Ii ii ii ii ii Relevé size (m²) 100 100 100 100 100 100 100 100 100 Diagnostic species of the association Clematido cirrhosae-Juniperetum lyciae Juniperus phoenicea 2 2 1 2 + 1 2 1 1 9 Clematis cirrhosa 3 3 3 2 3 3 2 + 4 9 Fumaria ouazzanensis 3 3 3 2 3 1 1 2 8 Pistacia lentiscus 1 2 + + + 5 Olea europaea 1 2 1 1 4 Limonium sinuatum 1 + + 1 4 Fedia pallescens 1 2 2 Clematis flammula 2 1 Phillyrea angustifolia + 1 Differential species of the subassociation retametosum monospermae Retama monosperma 3 4 4 3 4 4 2 3 4 9 Chamaerops humilis 1 + 1 3 + 5 Asparagus aphyllus 1 1 2 Other species Daucus carota 4 4 2 2 3 2 3 7 Rumex bucephalophorus 4 3 1 4 1 5 6 Galactites elegans 1 2 2 1 3 5 Anagalis arvensis 1 2 3 1 3 5 Mercurialis annua 4 1 3 1 1 5 Silene gallica 2 2 1 1 3 5 Emex spinosa 3 2 4 3 4 23/1 • 2024, 35–68 58 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco RELEVÉ NUMBER R27 R34 R43 R44 R89 R35 R42 R54 R61 Pr Papaver rhoeas 1 2 3 1 4 Geranium molle 4 3 3 1 4 Solanum sodomeum + 1 1 + 4 Paronychia argentea 1 2 2 2 4 Malva sylvestris 3 1 3 2 4 Cerinthe major 3 3 2 3 4 Linaria maroccana 1 1 + 1 4 Lolium perenne 1 1 2 3 4 Misopates calycinum 1 2 2 3 Plantago lagopus 3 2 2 3 Verbascum sinuatum 1 1 + 3 Parietaria mauritanica 2 1 2 3 Centaurea pullata 2 3 1 3 Delphinium pubescens 1 2 2 3 Smyrnium olusatrum 1 + 2 3 Sonchus oleraceus 2 3 3 3 Medicago orbicularis 2 3 2 3 Ononis natrix 2 3 3 3 Anacyclus clavatus 1 1 1 3 Reichardia tingitana 1 2 3 3 Cynodon dactylon 2 3 1 3 Bromus diandrus 2 4 1 3 Plantago lanceolata 3 3 3 3 Lotus corniculatus 3 2 2 Vicia lutea 3 2 2 Arisarum vulgare 2 2 2 Erodium cicutarium 3 3 2 Hordeum murinum 1 1 2 Echium vulgare 2 1 2 Hypochaeris glabra 2 3 2 Ammi majus 3 3 2 Lupinus angustifolius + + 2 Tolpis barbata 3 1 2 Hypochaeris radicata 2 3 2 Agropyron junceum 2 2 2 Achyranthes sicula + 2 2 Parentucellia viscosa + 1 2 Dipcadi serotinum 1 1 2 Anthemis arvensis 1 2 2 Anthyllis vulneraria 1 + 2 Erucastrum gallicum + + 2 Medicago truncatula 2 2 2 Amaranthus blitoides + + 2 Diplotaxis tenuifolia 2 1 2 Urospermum picroides + + 2 Dactylis glomerata 1 + 2 Verbascum virgatum 3 2 2 Ajuga iva + 2 2 Jasione montana + 3 2 Torilis nodosa 1 1 Riechardia tingitana 3 1 23/1 • 2024, 35–68 59 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco RELEVÉ NUMBER R27 R34 R43 R44 R89 R35 R42 R54 R61 Pr Arctotheca calendula 3 1 Evax pygmaea 3 1 Cerastium glomeratum + 1 Ornithopus sativus brot 2 1 Hedypnois rhagadioloides 2 1 Tuberaria guttata 1 1 Sherardia arvensis 3 1 Glebionis coronaria 3 1 Asparagus albus + 1 Vicia sativa 3 1 Lagurus ovatus 2 1 Astragalus hamosus 1 1 Ononis spinosa 2 1 Campanula rapunculus + 1 Umbilicus rupestris 3 1 Trifolium campestre 3 1 Erodium ciconium 2 1 Medicago polymorpha 3 1 Urtica membranacea 2 1 Umbilicus horizontalis 1 1 Calendula arvensis 1 1 Vulpia geniculata 3 1 Polypogon monspeliensis 2 1 Asphodelus microcarpus 2 1 Bromopsis inermis 2 1 Euphorbia segetalis 2 1 Anacyclus radiatus 1 1 Juncus acutus 3 1 Heteranthemis viscidehirta 3 1 Lantana camara 2 1 Verbascum boerhavii 1 1 Scille maritime + 1 Herniaria glabra + 1 Trisetaria panicea 3 1 Poa pratensis 2 1 Nigella arvensis + 1 Anchusa officinalis 1 1 Ononis reclinata 3 1 Prangos trifida 1 1 Orobanche sanguinea + 1 Pseudorlaya pumila 2 1 Arenaria mauritanica 3 1 Bartsia trixago 2 1 Gladiolus communis 3 1 Allium ampeloprasum 2 1 Lium strictum 1 1 Evolvulus arizonicus + 1 Thapsia garganica 1 1 Opuntia ficus indica + 2 Eucalyptus gomphocephala + 1 23/1 • 2024, 35–68 60 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco Ta bl e 10 (T ab el a 10 ): C lem at id o cir rh os ae -Ju ni pe re tu m ly cia e c ha m ae ro pe to su m h um ili s s ub as s. no v. R EL EV É N U M BE R R 7 R 10 R 11 R 24 R 25 R 26 R 37 R 38 R 39 R 40 R 55 R 60 R 71 R 75 R 78 Pr C oo rd in at e La t. N or th Lo ng . W es t -6.6890 34.2094 -6.6858 34.2170 -6.6882 34.2148 -6.6789 34.2258 -6.6720 34.2357 -6.6752 34.2302 -6.6630 34.2548 -6.6629 34.2530 -6.6636 34.2520 -6.6637 34.2514 -6.6669 34.2556 -6.6712 34.2519 -6.6790 34.2407 -6.6966 34.2093 -6.6945 34.2147 St at io n 8 12 9 11 11 11 14 11 11 19 5 4 4 7 1 El ev at io n (m ) 20 18 42 17 16 16 19 34 24 31 20 25 43 20 57 Pr O rie nt at io n S SE S W S S W S N N N N S N W S N S Sl op e (% ) 0 0 19 ,8 3, 7 4, 4 2, 2 3, 9 2, 8 25 5, 3 4, 4 28 ,2 18 ,2 0 23 So il iii iii iii ii ii ii ii Ii ii iii ii ii ii i i Re le vé si ze (m ²) 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 D ia gn os ti c sp ec ie s o f t he a ss oc ia ti on C le m at id o ci rr ho sa e- Ju ni pe re tu m ly ci ae Ju ni pe ru s p ho en ice a 3 3 1 2 1 + 3 1 2 3 2 2 1 2 1 15 Am m i m aj us 2 1 1 2 1 + 1 2 2 + + 2 12 Fu m ar ia o ua zz an en sis 2 1 2 2 3 1 6 Li m on iu m si nu at um 1 1 + 2 5 As pa ra gu s a ph yll us 1 1 + 2 1 5 O lea eu ro pa ea 1 1 2 1 4 Fe di a pa lle sce ns 1 2 2 Ep he dr a fra gi lis + + 2 Ly ciu m eu ro pe au m + + 2 D iff er en ti al sp ec ie s o f t he su ba ss oc ia ti on c ha m ae ro pe to su m h um ili s Re ta m a m on os pe rm a + 4 4 3 3 4 3 4 2 4 2 3 2 3 2 15 So la nu m so do m eu m + + 1 + 1 1 1 + 8 C ha m ae ro ps h um ili s + 2 + + 1 + 6 Ac hy ra nt he s s icu la 3 1 2 1 2 5 M ar ru bi um v ul ga re 1 1 1 1 4 As pa ra gu s a lb us 3 3 1 1 4 C er in th e m aj or 2 1 1 3 O th er sp ec ie s Em ex sp in os a 3 2 3 2 3 2 2 3 3 2 10 G al ac tit es ele ga ns 3 2 3 3 2 1 1 2 3 9 An ag al is ar ve ns is 2 2 2 3 2 3 2 3 8 Ru m ex b uc ep ha lo ph or us 3 2 2 2 3 2 3 1 8 Si len e g al lic a 1 2 2 2 2 1 1 2 8 La gu ru s o va tu s 2 3 2 2 2 2 3 2 8 Pl an ta go la go pu s 2 2 2 2 2 3 2 7 23/1 • 2024, 35–68 61 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco R EL EV É N U M BE R R 7 R 10 R 11 R 24 R 25 R 26 R 37 R 38 R 39 R 40 R 55 R 60 R 71 R 75 R 78 Pr Lo tu s c or ni cu la tu s 1 3 3 3 3 3 3 7 To lp is ba rb at a 2 3 2 3 1 2 1 7 H or de um m ur in um 3 3 2 2 1 3 3 7 U rt ica m em br an ac ea 2 2 2 2 2 2 6 Pa ro ny ch ia a rg en te a 2 1 1 2 3 5 Ar cto th ec a ca len du la 2 1 1 2 3 5 C en ta ur ea p ul la ta 1 2 + 2 2 5 Er od iu m ci cu ta riu m + 1 3 2 3 5 M al va sy lv est ris 3 3 2 2 1 5 D au cu s c ar ot a 3 2 2 1 4 Pa pa ve r r ho ea s 1 + 1 1 4 O no ni s s pi no sa 2 3 3 3 4 C al en du la a rv en sis 3 2 2 2 4 M er cu ria lis a nn ua 3 2 2 2 4 Ar isa ru m v ul ga re 2 3 2 2 4 M iso pa te s c al yc in um 2 + 1 2 4 Ve rb as cu m b oe rh av ii 2 + + + 4 So nc hu s o ler ac eu s 2 3 3 1 4 Pa re nt uc ell ia v isc os a 1 + 3 3 Sm yr ni um o lu sa tr um 2 3 3 3 Vi cia sa tiv a 2 2 3 3 Li na ria m ar oc ca na + 1 1 3 As tra ga lu s h am os us 1 1 2 3 Lo liu m p er en ne 2 3 3 3 Sc ol ym us h isp an icu s 1 1 1 3 Sa lv ia v er be na ca + 2 2 Ec hi um v ul ga re 2 2 2 Ag ro py ro n ju nc eu m 2 2 2 Eu ph or bi a te rr ac in a 1 3 2 To ril is no do sa 3 3 2 Ve rb as cu m si nu at um + + 2 Ev ax p yg m ae a 1 2 2 G er an iu m m ol le 2 3 2 C er as tiu m g lo m er at um 1 + 2 G er an iu m ro tu nd ifo liu m 3 1 2 O rn ith op us sa tiv us 1 2 2 23/1 • 2024, 35–68 62 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco R EL EV É N U M BE R R 7 R 10 R 11 R 24 R 25 R 26 R 37 R 38 R 39 R 40 R 55 R 60 R 71 R 75 R 78 Pr Tu be ra ria g ut ta ta 2 3 2 H yp oc ha er is gl ab ra 2 2 2 Tr ifo liu m ca m pe str e 2 3 2 M ed ica go p ol ym or ph a 2 3 2 M ed ica go o rb icu la ris 3 3 2 Br iz a m aj or (m ax im a) 3 2 2 O no ni s n at rix 2 2 2 H ym en oc ar po s c irc in na tu s + 1 2 Bi scu te lla la ev ig at a 2 3 2 Pl an ta go m ac ro rh iz a 2 1 2 So nc hu s t en er rim us 3 2 2 D ip ca di se ro tin um 2 3 2 C irs iu m v ul ga re 3 2 2 Ri cin us co m m un is 3 1 2 So nc hu s a sp er 3 1 2 Ps eu do rla ya p um ila 1 2 2 U rg in ea m ar iti m a 1 1 Av en a ste ril is 2 1 N ico tia na g la uc a 1 1 Pa rie ta ria m au rit an ica 3 1 H yp oc ha er is ra di ca ta 1 1 Sh er ar di a ar ve ns is 1 1 G leb io ni s c or on ar ia + 1 Tr ifo liu m to m en to su m 3 1 M or ae a sis yr in ch iu m + 1 So nc hu s a rv en sis 2 1 Li na ria p ur pu re a 1 1 Tr ifo liu m n ig re sce ns 3 1 U rt ica u re ns 3 1 M ed ica go m in im a 1 1 C re pi s c ap ill ar is 2 1 D elp hi ni um p ub esc en s + 1 O rn ith op us co m pr ess us 1 1 Lu pi nu s m icr an th us + 1 Er od iu m ci co ni um 2 1 Ec hi um p la nt ag in eu m 1 1 23/1 • 2024, 35–68 63 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco R EL EV É N U M BE R R 7 R 10 R 11 R 24 R 25 R 26 R 37 R 38 R 39 R 40 R 55 R 60 R 71 R 75 R 78 Pr Ru bi a pe re gr in a 1 1 Vi cia v ill os a 1 1 G er an iu m p ur pu re um 2 1 Po a an nu a 3 1 Lo bu la ria m ar iti m a 3 1 Tr ifo liu m ch er ler i 4 1 Tr ifo liu m su bt er ra ne um 2 1 Tr ifo liu m a rv en se 2 1 Po lyp og on m on sp eli en sis 2 1 M iso pa te s o ro nt iu m 2 1 Sc he do no ru s p ra te ns is 2 1 H yp ar rh en ia h irt a 5 1 As ph od elu s m icr oc ar pu s 3 1 Ba scu te lla la ev ig at a 4 1 An ac yc lu s c la va tu s 2 1 Ju nc us a cu tu s 2 1 Re ich ar di a tin gi ta na 3 1 La nt an a ca m ar a 2 1 H eli ot ro pi um eu ro pa eu m 2 1 Ro rip pa sy lv est ris 2 1 Se du m ru pe str e 1 1 Sp er ul a ru pi co la 1 1 Ve rb as cu m v irg at um 1 1 G al iu m a pa rin e 2 1 Tr ifo liu m h yb rid um 3 1 Sc ab io sa a tro pu rp ur ea 1 1 M eli lo tu s o ffi cin al is 2 1 Fi cu s c ar ica + 1 Ag av e a m er ica na 2 1 O pu nt ia fi cu s i nd ica 2 1 Ac ac ia cy an op hy lla + 2 2 Eu ca lyp tu s g om ph oc ep ha la + 1 23/1 • 2024, 35–68 64 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco Ta bl e 11 (T ab el a 11 ): So la no so do m ae um -R et am et um m on os pe rm ae (J di e t a l., 1 98 7) R EL EV É N U M BE R R 4 R 5 R 28 R 9 R 12 R 3 R 77 R 1 R 31 R 36 R 73 R 88 R 2 R 33 R 56 R 6 R 13 R 14 R 15 R 16 R 17 R 30 R 10 8 Pr C oo rd in at e La t. N or th Lo ng . W es t -6.6934 34.2062 -6.6916 34.2048 -6.6699 34.2393 -6.6869 34.2130 -6.6891 34.2130 -6.6950 34.2053 -6.6945 34.2120 -6.6958 34.2067 -6.6697 34.2420 -6.6615 34.2565 -6.6913 34.2129 -6.6661 34.2564 -6.6967 34.2058 -6.6732 34.2402 -6.6712 34.2555 -6.6901 34.2075 -6.6913 34.2118 -6.6923 34.2118 -6.6923 34.2111 -6.6923 34.2102 -6.6934 34.2084 -6.6696 34.2412 -6.6803 34.2446 St at io n 6 8 11 17 10 6 1 6 11 20 9 5 6 15 7 8 6 6 6 6 6 11 2 El ev at io n (m ) 20 20 33 40 30 20 30 20 20 25 24 19 20 68 20 20 29 39 22 31 20 27 54 Pr O rie nt at io n S S S S N W S N SE N E W S N N N N S N E E W N S N Sl op e (% ) 0 0 8 6 9 0 7 0 0 16 12 15 0 7 0 0 15 9 6 4 3 15 51 So il iii iii ii iii ii iii i iii ii ii iii ii iii Ii ii iii iii iii iii iii iii ii ii Re le vé si ze (m ²) 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 10 0 D ia gn os ti c sp ec ie s o f t he a ss oc ia ti on S ol an o so do m ae um -R et am et um m on os pe rm ae Re ta m a m on os pe rm a 4 4 5 2 5 3 4 4 4 5 4 2 5 5 5 5 5 4 3 3 5 5 2 23 C ha m ae ro ps h um ili s + + + + + 1 + 7 So la nu m so do m eu m + + + + + 3 2 7 As pa ra gu s a ph yll us 2 1 2 1 2 5 O lea eu ro pa ea 1 1 1 + 4 Ep he dr a fra gi lis + 3 + 3 Ar isa ru m v ul ga re 1 1 2 3 C lem at is cir rh os a 3 2 2 Pi sta cia le nt isc us 1 3 2 Ph ill yr ea a ng us tif ol ia + 1 Ru bi a pe re gr in a 1 1 O th er sp ec ie s Ru m ex b uc ep ha lo ph or us 3 2 1 2 2 1 3 3 4 4 4 2 2 13 An ag al is ar ve ns is 2 2 1 1 1 2 2 1 1 2 1 11 G al ac tit es ele ga ns 1 1 2 2 2 4 + 2 2 2 10 To ril is no do sa 2 1 + 1 3 1 2 2 1 1 10 Ar cto th ec a ca len du la 4 3 1 2 3 3 1 1 3 2 10 Am m i m aj us 2 2 2 3 2 5 + 1 1 + 10 M al va sy lv est ris 3 2 3 3 5 3 3 1 8 M ar ru bi um v ul ga re 1 + 3 + 1 1 1 7 Pa ro ny ch ia a rg en te a 2 2 2 + 1 2 1 7 H or de um m ur in um 3 1 1 1 1 4 3 7 Fu m ar ia o ua zz an en sis + 1 3 1 1 + 2 7 M iso pa te s c al yc in um 1 2 + 2 1 + 6 Lo tu s c or ni cu la tu s 2 1 3 3 + 1 6 23/1 • 2024, 35–68 65 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco R EL EV É N U M BE R R 4 R 5 R 28 R 9 R 12 R 3 R 77 R 1 R 31 R 36 R 73 R 88 R 2 R 33 R 56 R 6 R 13 R 14 R 15 R 16 R 17 R 30 R 10 8 Pr Em ex sp in os a 2 1 3 4 1 1 6 Ev ax p yg m ae a 1 2 + 2 2 + 6 M er cu ria lis a nn ua 2 + 3 4 3 4 6 Pa rie ta ria m au rit an ica 3 1 2 3 1 5 H yp oc ha er is gl ab ra 1 3 1 2 + 5 La gu ru s o va tu s 4 2 3 1 1 5 Sc ol ym us h isp an icu s + + 2 1 4 Ve rb as cu m si nu at um 1 + + + 4 N ico tia na g la uc a + 3 1 + 4 Pa pa ve r r ho ea s + 1 4 + 4 G er an iu m m ol le 3 1 4 2 4 C en ta ur ea p ul la ta 2 2 1 3 4 O no ni s s pi no sa 2 2 3 3 4 D elp hi ni um p ub esc en s + 1 + + 4 Li m on iu m si nu at um + 2 + + 4 U rt ica m em -b ra na ce a 3 1 2 3 4 Ac hy ra nt he s s icu la 1 2 2 1 4 D au cu s c ar ot a 2 3 1 1 4 Ju nc us a cu tu s + 1 2 4 4 Pl an ta go la go pu s 1 1 1 3 Sa lv ia v er be na ca + + 1 3 U rt ica d io ica 2 1 1 3 Vi cia lu te a 1 2 1 3 C er as tiu m g lo m er at um + + 1 3 G er an iu m ro tu nd ifo liu m 1 2 1 3 Si lyb um m ar ia nu m + + + 3 Sh er ar di a ar ve ns is 3 + 2 3 Er od iu m ci cu ta riu m + 3 2 3 G leb io ni s c or on ar ia 3 + 1 3 Lu pi nu s a ng us tif ol iu s + + 1 3 Vi cia sa tiv a 2 3 2 3 G al ac tit es to m en to su s 1 2 1 3 To lp is ba rb at a 2 1 3 3 Sm yr ni um o lu sa tr um 2 3 1 3 M eli lo tu s i nd icu s 1 2 1 3 So nc hu s o ler ac eu s 3 3 3 3 U rg in ea m ar iti m a + 1 2 23/1 • 2024, 35–68 66 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco R EL EV É N U M BE R R 4 R 5 R 28 R 9 R 12 R 3 R 77 R 1 R 31 R 36 R 73 R 88 R 2 R 33 R 56 R 6 R 13 R 14 R 15 R 16 R 17 R 30 R 10 8 Pr Ri ec ha rd ia ti ng ita na + 2 2 Pl an ta go co ro no pu s 2 1 2 O rn ith op us sa tiv us 3 1 2 Ec hi um v ul ga re 1 3 2 As tra ga lu s h am os us 2 2 2 U rt ica u re ns 3 3 2 Sc or pi ur us m ur ica tu s + + 2 Tr ise tu m fl av esc en s 4 4 2 Ag ro py ro n ju nc eu m 3 3 2 M ed ica go p ol ym or ph a 3 1 2 C er in th e m aj or 2 + 2 C al en du la a rv en sis 1 4 2 Eu ph or bi a seg et al is 3 2 2 An ac yc lu s c la va tu s + 2 2 Ly th ru m ju nc eu m 3 1 2 Tr ise ta ria p an ice a 1 3 2 Ri cin us co m m un is 1 + 2 Tr ifo liu m a ur eu m 2 2 2 G al iu m a pa rin e 2 + 2 M icr op yr um te ne llu m 3 1 An isa nt ha m ad rit en sis 3 1 Av en a ste ril is 3 1 H ed yp no is rh ag ad io lo id es + 1 Tu be ra ria g ut ta ta 1 1 Tr ifo liu m to m en to su m 3 1 As pa ra gu s a lb us 2 1 Er yn gi um p la nu m 2 1 M or ae a sis yr in ch iu m + 1 Si len e g al lic a 2 1 Li na ria p ur pu re a + 1 Tr ifo liu m n ig re sce ns 2 1 C en tra nt hu s c al cit ra pa e 2 1 C am pa nu la ra pu nc ul us 1 1 As ph od elu s a lb us 1 1 M ed ica go m in im a 3 1 U m bi lic us ru pe str is + 1 23/1 • 2024, 35–68 67 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco R EL EV É N U M BE R R 4 R 5 R 28 R 9 R 12 R 3 R 77 R 1 R 31 R 36 R 73 R 88 R 2 R 33 R 56 R 6 R 13 R 14 R 15 R 16 R 17 R 30 R 10 8 Pr Lu pi nu s m icr an th us + 1 As tra ga lu s s esa m en s 1 1 Tr ifo liu m ca m pe str e 2 1 Er od iu m ci co ni um 1 1 M ed ica go o rb icu la ris 4 1 C am pa nu la lu sit an ica 3 1 O no ni s n at rix 1 1 Bi scu te lla la ev ig at a 2 1 Fe di a pa lle sce ns 1 1 Pr as iu m m aj us 2 1 Li na ria m ar oc ca na 2 1 Tr ifo liu m a rv en se 1 1 Tr ifo liu m a ng us tif ol iu m 3 1 La ctu ca se rr io la 1 1 Tr ifo liu m re pe ns 2 1 Lu pi nu s c os en tin ii 1 1 Ju nc us m ar iti m us 4 1 Be ru la er ec ta 3 1 La nt an a ca m ar a 2 1 Sc ill e m ar iti m e 2 1 C he no po di um a lb um + 1 Lo liu m p er en ne 4 1 An isa nt ha st er ili s 2 1 Ve rb as cu m v irg at um 2 1 Eu ph or bi a te rr ac in a 1 1 Vu lp ia b ro m oi de s + 1 Sc ab io sa a tro pu rp ur ea     1 1 23/1 • 2024, 35–68 68 Challi et al. Phytosociological analysis of the Sidi Boughaba Biological Reserve, Morocco Table 12 (Tabela 12): Pterido aquilinum-Populetum albae ass.nov. RELEVÉ NUMBER R32 R49 R74 Pr Papaver rhoeas + 1 Anagalis arvensis 4 1 Parietaria mauritanica 3 1 2 Cerastium glomeratum 1 1 Geranium rotundifolium l 2 1 Arisarum vulgare 4 1 Hordeum murinum 3 2 2 Ammi majus 2 1 Malva sylvestris 4 2 2 Lagurus ovatus 2 2 2 Ononis spinosa 2 1 Urtica urens 2 1 Fumaria ouazzanensis 1 4 2 Trifolium campestre 2 1 Agropyron junceum 3 1 Achyranthes sicula 2 1 Cerinthe major 2 3 2 Daucus carota 4 1 Biscutella laevigata 3 1 Fedia pallescens 3 1 Lobularia maritima 3 1 Anacyclus radiatus 2 1 Anacyclus clavatus 1 3 2 Scrophularia auriculata 3 1 Ricinus communis 1 1 Lolium perenne 2 3 2 Anisantha sterilis 3 1 Verbascum virgatum 1 1 Adonis aestivalis 1 1 Galium aparine 2 1 Oxalis pes-caprae 2 1 Brachypodium distachyum   2 1 RELEVÉ NUMBER R32 R49 R74 Pr Coordinate Lat. North Long. West -6 .6 72 0 34 .2 40 0 -6 .6 70 0 34 .2 45 6 -6 .6 90 2 34 .2 13 9 Pr Station 15 18 9 Elevation (m) 69 21 20 Orientation N S WS Slope (%) 11 12 1 Soil iv iv Iv Relevé size (m²) 100 100 100 Diagnostic species of the association   Populus alba + 2 5 3 Pteridium aquilinum + + 3 3 Adiantum capillus-veneris + + 2 3 Smyrnium olusatrum 3 1 Bryonia dioica 3 1 Smilax aspera 2 1 Espèces compagnes Galactites elegans 3 1 Rubus ulmifolius 2 1 Rumex bucephalophorus 3 1 Lotus corniculatus 2 1 Torilis nodosa 1 3 2 Juncus acutus 2 2 2 Rubia peregrina 1 2 2 Solanum sodomeum 1 1 Clematis cirrhosa 2 1 Retama monosperma + 1 Marrubium vulgare 2 1 Verbascum sinuatum 2 1 Phragmites australis 4 1 Arctotheca calendula 3 1 Nicotiana glauca 2 1 Legend Pr: Presence, (i): raw mineral soils on unconsolidated white dunes, (ii): calco-magnesimorphic soils of eolian origin and sandy texture on deep sand and dune sandstone, (iii): Mediterranean red sandy-clay soils, with little moisture on the slopes, superficial on dune sandstone and very deep in the inter-dune depression, (iv): Hygromorphic soils corresponding to the water table, with sandy horizons of varying depths.