119
Key words: alien species, 
Chorology, Coniferae, Cryptomeria 
D. Don, Laricoideae, naturalization, 
Pinopsida, Taxodium Rich.
Ključne besede: tujerodne vrste, 
horologija, Coniferae, Cryptomeria 
D. Don, Laricoideae, naturalizacija, 
Pinopsida, Taxodium Rich.
Corresponding author:
Ridha El Mokni
E-mail:  
ridhaelmokni@yahoo.fr
Received: 5. 8. 2022
Accepted: 14. 11. 2023
New records of coniferous species 
(Gymnospermae, Pinidae) for the non-native 
woody flora of Tunisia and North Africa
 
Abstract
Ten new taxa of gymnosperms are added to the non-indigenous woody flora of 
Tunisia, after almost two decades of botanical surveys. Four of them (Cryptomeria 
japonica, Pinus brutia var. pityusa, Pseudotsuga menziesii var. glauca, Taxodium 
distichum var. distichum) are also new for the flora of North Africa. Pinus coulteri 
is here first reported for the Mediterranean Basin. Three genera (Cryptomeria, 
Pseudotsuga and Taxodium) are firstly recorded for the non-native vascular flora 
of Africa whereas two genera (Cedrus and Platycladus) are reported for the first 
time only for the woody flora of Tunisia. Brief descriptions together with filed 
photographs, global distributions and habitats at national scale are provided. Some 
taxonomic comments on their distinguishable features as well as updated keys are 
also presented. 
Izvleček
Po skoraj dveh desetletjih botaničnih raziskav smo neavtohtoni lesnati flori Tunizije 
dodali deset novih taksonov golosemenk. Štiri med njimi (Cryptomeria japonica, 
Pinus brutia var. pityusa, Pseudotsuga menziesii var. glauca, Taxodium distichum var. 
distichum) so novi tudi za floro Severne Afrike. Pinus coulteri je prvič omenjen za 
sredozemski bazen. Trije rodovi (Cryptomeria, Pseudotsuga in Taxodium) so bili 
prvič zabeleženi za tujerodno floro cevnic Afrike, medtem ko sta dva roda (Cedrus 
in Platycladus) prvič zabeležena le za floro lesnatih rastlin Tunizije. Podani so kratki 
opisi s fotografijami, globalna razširjenost in habitati v Tuniziji. Predstavljeni so 
tudi nekateri taksonomski komentarji o njihovih značilnostih in posodobljeni 
določevalni ključi.
Ridha El Mokni1,2
1 Department of Pharmaceutical Sciences ‘A’, Laboratory of Botany, Cryptogamy and Plant Biology, Faculty of Pharmacy of Monastir, Avenue Avicenna, 
Monastir-5000, Monastir University, Tunisia.
2 Department of Forestry, Laboratory of Forest Ecology, National Research Institute of Rural Engineering, Water and Forests, Ariana-2080, IRESA, Carthage 
University, Tunisia.
DOI: 10.2478/hacq-2023-000423/1 • 2024, 119–136
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New records of coniferous species for the non-native woody flora of Tunisia and 
North Africa
Introduction
The introduction, during the last few centuries and in re-
cent decades, of thousands of woody plant species in al-
most all over of the world is increasingly intensified. These 
species used mainly in ornamental gardening, silviculture, 
erosion control, for fruit sources or other purposes, can 
easily escape and resulting in the naturalization (some-
times even as invasive) of many trees and shrubby taxa 
(Richardson & Rejmánek, 2011). Woody plants selected 
for introduction usually originate from regions with simi-
lar climate conditions; therefore, after certain time lag, 
they start to spread outside places of cultivation, become 
naturalized or even invasive (Gudžinskas et al., 2017). 
Such introductions contribute in the majority of cases to 
the increase of alien woody species in Europe as well as in 
other regions of the world (se e.g. Pyšek et al., 2009; Rich-
ardson & Rejmánek, 2011; Pergl et al., 2016; Mayer et al., 
2017). Regions with the largest number of woody invasive 
alien species are Australia (183 species), Southern Africa 
(170 species), North America (163 species), and Pacific 
Islands (147 species) whereas Europe with 107 woody in-
vasive alien species (59 species of trees and 48 species of 
shrubs) is in the fifth place together with New Zealand 
(cf. Richardson & Rejmánek, 2011). In Northern Africa, 
there is no exact updated number for woody (incl. gym-
nosperms) alien species since the percentage (ca. 13 %) 
published by Vilà et al. (1999) despite many recent publi-
cations dealing with new shrubby and woody alien species 
(see e.g. El Mokni 2019a, 2019b, 2019c, 2019d, 2019e, 
2019f, 2020a, 2020b, 2021a, 2021b, 2021c, 2021d; El 
Mokni & De Bélair, 2020; El Mokni & Domina, 2020a, 
2020b; El Mokni & Iamonico, 2020; El Mokni & Ver-
loove, 2020a, 2020b; Iamonico & El Mokni, 2021; El 
Mokni et al., 2022). Thus, high probability of new records 
of escaped woody species exists and needs more and more 
field investigations. Providing data about the distribution 
and status of naturalization of escaped alien woody plants 
remains useful and essential to prevent or reduce their im-
pacts/threats on native communities and ecosystems (see 
e.g. Pyšek et al. 2020; Gallardo et al. 2019).
As part of an ongoing investigation on the Tunisian 
coniferous diversity, extended botanical surveys were 
undertaken, over the last twelve years in some Northern 
and Central Tunisian areas (mainly Kroumirian, Mogods’ 
and Tunisian Dorsale mountains) and had led to the col-
lection of specimens of 10 conifer species that represent 
new national/North African records or new distributions 
at national scale. This paper details those new records, 
updates dichotomous keys for each new reported taxon 
and gives an updated checklist of the Pinidae Subclass in 
Tunisia. 
Material and methods
This work is based on extensive field botanical surveys car-
ried out in various regions of Tunisia mainly within the 
governorate of Jendouba for more a decade (2010 to 2022) 
and the revision of relevant literature. Collected specimens 
are stored at the personal herbarium of the author (Her-
barium El Mokni!) deposited in the Faculty of Pharmacy 
of Monastir (not yet listed in Index Herbariorum, Thiers, 
2023+), Monastir University. Reported taxa are presented 
in alphabetical order (the nomenclature and taxonomic 
treatment are following mainly Raab-Straube, 2014+ in 
Euro+Med PlantBase; POWO, 2022; APD, 2022), mor-
phological description (incl. author personnel observa-
tion) and global distribution are mainly according to Earle 
(2007+), Eckenwalder (2009), Farjon & Filer (2013) and 
Farjon (2017), distribution and habitat in Tunisia are re-
ported from personal observations. When other genera are 
newly cited within the same family or also other species 
and/or subspecies within the same genus are newly re-
ported for the Tunisian coniferous flora, an updated (di-
chotomous) key is provided (cf. Schulz et al., 2005; Farjon, 
2017). Taxonomic notes are also given, once necessary.
A list of updated and hierarchical taxonomic position 
of different accepted gymnosperm taxa (native and aliens) 
in Tunisia with their actual distributions is provided in 
Appendix 1.
Results
Ten new taxa of gymnosperms that represent new Tuni-
sian/North African records are here arranged alphabetical-
ly according to their Subclasses, Orders, Families, Subfam-
ilies, Genera, Subgenera, Sections and Subsections. Four 
of them (Cryptomeria japonica, Pinus brutia var. pityusa, 
Pseudotsuga menziesii var. glauca, Taxodium distichum var. 
distichum) are new for the flora of North Africa. Pinus 
coulteri is here first reported for the Mediterranean Basin. 
Three genera (Cryptomeria, Pseudotsuga and Taxodium) are 
firstly recorded for the non-native vascular flora of Africa 
while the genera Cedrus and Platycladus are reported for 
the first time only for the woody flora of Tunisia.
SUBCLASS PINIDAE1 Cronquist, Takht. & Zim-
merm., Taxon 15: 134 (1966).
ORDER 1. PINALES Gorozh., Lekts. Morf. Sist. Ar-
chegon.: 88 (1904).
1 Within this Subclass, we find also the genus Araucaria Juss. [Araucariace-
ae Henkel & W. Hochst., Araucariales Gorozh. (1904)] with commonly 
A. araucana (Molina) K. Koch, A. heterophylla (Salisb.) Franco planted in 
many regions of Tunisia where they are able to produce a lot of seeds each 
year without any note for juvenile individuals from seedlings! 
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New records of coniferous species for the non-native woody flora of Tunisia and 
North Africa
Family. 1. Pinaceae Spreng. ex F.Rudolphi, Syst. Orb. 
Veg.: 35 (1830), nom. cons.
Subfam. 1. Abietoideae (Bercht. & J.Presl, 1820) Rich. 
ex Sweet, 1826.
Genus 1. Cedrus Trew, 1757
Cedrus atlantica (Endl.) Manetti ex Carrière, Traité 
Gén. Conif.: 285. 1855. 
Syn. C. libani A. Rich. var. atlantica (Endl.) Hook. f., 
[Cedars Lebanon] Nat. Hist. Rev., ser. 2, 2: 15. 1862; 
C. libani A. Rich. subsp. atlantica (Endl.) Batt. & Trab., 
Fl. Algérie Tunisie: 397. 1905. 
First official report as an established taxon for the flora 
of Tunisia.
Trees up to 20 m tall and ca. 30 cm in diameter at 
breast height, usually with a single columnar trunk, often 
forked. Crown broad-conical, composed of massive pri-
mary branches, with foliar units often arranged in hori-
zontal planar groups; bark rough, reddish brown weather-
ing dark gray. Twigs short, firm, gray-brown. Vegetative 
buds’ ovoid or globose; bud scales broad-ovate, reddish 
to very dark brown, deciduous; leaves on long shoots spi-
rally arranged, more crowded near base of shoot; on short 
shoots densely crowded; apex acute to acuminate; seed 
cones also terminal on short shoots, erect, sessile, woody 
in the second year, ovoid to cylindrical with an obtuse 
to inset apex, 5–8 × 3–5 cm, light green ripening to pale 
brown, dehiscent with a persistent rachis; seed scales thin, 
coriaceous, 2–3 cm long and 2.5–2.5 cm wide, glabrous, 
pubescent at base, orange-brown, margin entire; seeds 
ovoid-conical, brown, 8–13 mm long. 
Distribution: The species has a North African na-
tive range (Algeria and Morocco). It was introduced 
and cultivated in many European countries (Portugal, 
Spain, France, Italy), reported as naturalized in Great 
Britain and as casual in Corse (as Cedrus libani subsp. at-
lantica  (Endl.) Batt. & Trab.) (Raab-Straube, 2014+ in 
Euro+Med PlantBase; POWO, 2022a; APD, 2022a). 
Habitat in Tunisia (Figure 1): Cedrus atlantica is 
found (more than 100 individuals) within mixed oak for-
ests (Quercus canariensis Willd. and Q. suber L.) in the 
region of Aîn Draham (Aîn Boulehya) and its surround-
Figure 1: Cedrus atlantica in Tunisia. A: habit of some trees in their habitat within Kroumirian forests; B: part of the bark trunk; C: leaves; 
D: immature female cones; E: mature seed cone. (Ghar Dimaou, NW Tunisia), Photographs R. El Mokni.
Slika 1: Cedrus atlantica v Tuniziji. A: habitus dreves na njihovem rastišču v Kroumirskih gozdovih; B: del lubja; C: listi; D: nezreli ženski storži; 
E: zrel semenski storž. (Ghar Dimaou, SZ Tunizija), fotografije R. El Mokni. 
D
E
C
AB
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New records of coniferous species for the non-native woody flora of Tunisia and 
North Africa
ings (El Ghorra, Djebel Bir, Majen Essaf ), in the region 
of Ghar Dimaou (El Mouadjen), in the surroundings of 
Tabarka region within Jendouba governorate (NW Tuni-
sia) where it was introduced since colonization. Popula-
tions of El Mouadjen, Majen Essaf and El Ghorra show 
many individuals of different ages and diameter at breast 
height thus many are from seedlings. 
Notes: Compared to Cedrus libani A. Rich. (an indig-
enous to E Mediterranean area), C. atlantica shows seed 
cones up to 8 cm long when fully mature (vs. up to 10 
(-12) cm long). Moreover, pollen cones are shorter than 
5 cm for both C. atlantica and C. libani whereas they are 
usually longer than 5 cm in C. deodara (Lamb.) G. Don. 
Subfam. 2. Pinoideae Link, 1831.
Genus 2. Pinus L., Sp. Pl. 2: 1000 (1753).
Subg. 1. Pinus
Sect. 1.1. Trifolius2 Duhamel
Subsect. 1.1.1. Ponderosae Loudon 
Pinus coulteri D. Don, Trans. Linn. Soc. London 17: 
440. 1836. 
Syn. Pinus ponderosa Douglas ex C. Lawson subsp. coul-
teri (D. Don) E. Murray, Kalmia 12: 23.1982. 
First official report as a casual species for the coniferous 
flora of Africa and the Mediterranean area.
Trees to 15–25 m tall; trunk monopodial, straight or 
curved at base; bark on trunk rough, dark brown with 
black fissures; branches spreading horizontally to more or 
less ascending, forming a broad, irregular pyramidal and 
open crown; leaves in fascicles of 3, very rigid, straight 
or curved, slightly twisted, 15–25(–30) cm long, 1.9–
2.2 mm wide; margins serrulate; apex acute pungent to 
subulate; leaf colour light green to grey-green; seed cones 
solitary (occasionally in pairs), or in whorls of 3–4(–5) on 
bole of young trees; mature cones ovoid, massive, heavy, 
usually slightly oblique or curved, moderately serotinous, 
20–35 × 15–20 cm when open, extremely resinous; seed 
scales flat, thick woody with thin, curved margins, widest 
towards the apophysis; apophysis very strongly developed, 
merging into a long, uncinate umbo, up to 30 mm wide, 
light yellowish brown or light caramel coloured; dorsal 
umbo, elongate, curved, with keeled sides, 25–35 mm 
long, 10–15 mm wide at base, ending in a sharp unci-
nate claw (Figure 2.A); seeds obliquely obovoid, slightly 
flattened, 10–18 × 7–10 mm, smooth, lustrous dark 
brown, turning blackish; seed wings dolabriform to semi-
2 Within this section, many species (Pinus contorta Douglas; Subsect. Con-
tortae Little et Critchfield and P. radiata D. Don; Subsect. Attenuatae Van 
Der Burgh) were introduced and cultivated with hundreds of individuals 
in many regions mainly of Northern Tunisia and could more probably 
naturalizing from seedlings!
ovate, 18–30 × 12–16 mm, orange-brown to dark reddish 
brown.
Distribution: The species is native to the USA: Cali-
fornia (Coast Ranges); Mexico: Baja California Norte. In 
North Africa, the taxon is reported as introduced in Mo-
rocco and as cultivated in Algeria (Raab-Straube, 2014+ 
in Euro+Med PlantBase).
Habitat in Tunisia: Pinus coulteri is found within some 
clayey slopes in the region of Aîn Draham and its sur-
roundings, within Jendouba governorate (NW Tunisia) 
where it seems to be introduced since colonization. Juve-
nile individuals (ca. 17) are reported from seedlings but 
usually misidentified due to the absence of their typical 
seed cones! 
Notes: Despite that Pinus coulteri shows similarities 
with P. sabiniana Douglas ex D. Don (with which shares 
in part the same biotope) in having almost fasciculate 
leaves; persistent seed cones, opening slowly; scales thick 
woody; apophyses extremely raised and elongated into a 
strongly curved spine; it is different by having thick and 
rigid leaves, 1.9–2.2 mm wide (vs. slender leaves, pliant, 
1.5 mm wide), seed cones solitary or in whorls of 2–5 (vs. 
seed cones solitary, rarely in pairs for P. sabiniana). 
Sect. 1. 2. Pinus
Subsect. 1. 2.1. Pinaster Mayr ex Koehne
Pinus brutia Ten., Fl. Napol. 1, Prodr.: lxxii. 1811–15.
Pinus brutia Ten. var. pityusa (Steven) Silba, Phytologia 
58: 367. 1985. 
Syn. P. pityusa Steven, Bull. Soc. Imp. Naturalistes Mos-
cou 11: 49. 1838.
First official report as a naturalizing taxon for the flora 
of North Africa.
Trees up to 30 m tall; trunk usually a straight bole, 
or slightly sinuous, sometimes forked; bark thin, orange-
brown, becoming thick only on lower trunk of large 
trees, then deeply fissured longitudinally, scaly, breaking 
into elongated plates, pale brown to red-brown; branches 
long, spreading and ascending, forming a broad pyrami-
dal or rounded, open crown; buds ovoid-conical, acute, 
10–15 mm long, not resinous; cataphylls with recurved 
apices, reddish brown fringed with white hairs; leaves in 
fascicles of 2 (sometimes 3), held by a 5–8 cm long, rigid 
and spreading, persistent sheath, retained on branchlet 
2–3 years, straight and rigid, spreading, rarely lax and 
pendulous, (5–)10–18(–29) cm long, 1–1.5 mm wide; 
pollen cones spirally arranged, short cylindrical, 1–2 cm 
long, yellow; seed cones solitary or in whorls of 2–3(–4), 
short pedunculate to nearly sessile, persistent, spreading 
forward or at right angle to shoot when full grown, nar-
rowly or broadly ovoid-conical or ovoid to rarely sub-
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New records of coniferous species for the non-native woody flora of Tunisia and 
North Africa
wide, broadly rhombic in outline, tan or grey-brown, 
unarmed; seeds obovoid, slightly flattened, 6–7(–8) mm 
long, grey-brown, sometimes dark mottled; wing 14–
20 mm long, 8–11 mm wide, oblique, grey-brown with 
darker streaks.
Distribution: the variety pityusa (Steven) Silba is native 
to North Caucasus: Krasnodar, Transcaucasia (Abkhazia), 
Georgia, Ukraine: Krym (POWO, 2022f ).
Habitat in Tunisia (Figure 2): 
Pinus brutia  var. pityusa is found 
within mixed oak forests pre-
dominantly by cork oak (Quercus 
suber L.) in the region of Aîn Dra-
ham (Babouch) and the region of 
Fernana (Aîn Debba) within the 
governorate of Jendouba (NW 
Tunisia) where it was introduced 
and cultivated since many decades 
in clayey slopes. Both populations, 
mainly that of Aîn Debba show 
many individuals of different ages 
and diameter at breast height thus 
many are from seedlings. 
Notes: the variety pityusa is 
easily distinguished for the other 
known varieties (var. brutia, var. 
eldarica (Medw.) Silba, and var. 
pendulifolia Frankis) by its short 
leaves with only 5–8 cm long (vs. 
10–18 cm long in var. brutia and 
18–29 cm long in var. pendulifo-
lia) and its red-brown apophyses 
(vs. whitish grey in var. eldarica).
globose when closed, (4–)6–11(–13) cm long, variously 
serotinous, 3–5 cm wide when closed, 5–8 cm wide 
when opened; seed scales thick woody, rigid, straight, 
oblong; apophyses nearly flat or slightly raised, (sharply) 
transversely keeled and with thin rays radiating from the 
centre, more or less rhombic or often with a rounded up-
per margin, to 20 mm wide at mid-cone, lustrous red-
brown weathering grey; umbo flat or depressed, 4–7 mm 
Figure 2: Pinus spp. in Tunisia. 
A: P. coulteri, mature close-up seed cone; 
B–C: P. brutia, crown, leaves and details 
of dehiscent mature seed cones. (Aîn 
Draham, NW Tunisia). Photographs 
R. El Mokni.
Slika 2: Pinus spp. v Tuniziji. 
A: P. coulteri, zrel semenski storž od 
blizu; B–C: P. brutia, krošnja, listi in 
podrobnosti zrelih semenskih storžev. 
(Aîn Draham, SZ Tunizija). Fotografije 
R. El Mokni.
B
A
C
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New records of coniferous species for the non-native woody flora of Tunisia and 
North Africa
A medium tree of (8)15–20 m in height, with an ir-
regular habitus and a clear crown; leaves up to 25 cm long 
with 2 mm wide, persistent, rigid with acute apex and 
finely denticulate margins, united in pairs in a sheath of 
10–20 mm; male cones yellow conical of 5 to 10 mm ag-
glomerated at the tops of the branches; female cones shiny 
light brown 8–15(18) x 5.5–7 cm, subsessile, solitary or 
grouped, subconical with woody 
scales, apical rhomboidal shield 
strongly keeled and mucronate; 
seed cones up to 22 cm long; seeds 
7–8 mm with a 20–30 mm wing.
Distribution: The subspecies 
has a W Mediterranean native 
range from Spain, Baleares to 
Italy, Sicily and Malta in North-
ern Mediterranean basin and only 
Algeria and Morocco in Northern 
Africa (Raab-Straube, 2014+ in 
Euro+Med PlantBase; POWO, 
2022b; APD, 2022b). 
Habitat in Tunisia (Figure 3): 
Pinus pinaster  subsp.  escarena  is 
found within mixed oak forests 
(Quercus canariensis Willd. and 
Q. suber L.) in the region of Aîn 
Draham (Babouch) and the re-
gion of Fernana (Aîn Debba) 
within the governorate of Jen-
douba (NW Tunisia) where it was 
introduced and cultivated since 
many decades in clayey slopes. 
Both populations, mainly that 
of Aîn Debba shows many indi-
viduals (ca. 70) of different ages 
and diameter at breast height thus 
many are from seedlings. 
Pinus pinaster Aiton subsp. escarena (Risso) K. Richt., 
Pl. Europ. 1: 1. 1890. 
Syn. P. escarena Risso, Hist. Nat. Europ. Mérid. 2: 340, 
459. 1826. 
First official report as an established taxon for the flora 
of Tunisia.
Figure 3: Pinus pinaster subsp. escarena 
in Tunisia. A: crown in bottom view; 
B: detailed part of the bark trunk; 
C: pollen cones above and seed cones 
below; D: solitary light brown shiny seed 
cone. (Tabarka & Fernana, NW Tunisia), 
Photographs R. El Mokni.
Slika 3: Pinus pinaster subsp. escarena 
v Tuniziji. A: krošnja od spodaj; 
B: podroben del lubja debla; C: pelodni 
storži zgoraj in semenski storži spodaj; 
D: posamezen svetlo rjav svetleč semenski 
storž. (Tabarka & Fernana, SZ Tunizija), 
fotografije R. El Mokni.
C D
A
B
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New records of coniferous species for the non-native woody flora of Tunisia and 
North Africa
Pinus pinaster Aiton subsp. pinaster
Syn. P. pinaster Aiton subsp. atlantica Villar, Bol. Soc. 
Esp. Hist. Nat. 33: 427. 1934. 
First official report as an established taxon for the flora 
of Tunisia.
Trees up to 30 m tall; leaves 10–20 cm long; resin ducts 
at base of leaves (within sheath) limited to the two lateral, 
larger ducts, at mid-leaf several additional small ducts; 
seed cones 10–18 cm long.
Distribution: The subspecies is known as native in Por-
tugal, Spain, France (Atlantic coast), Corse and Italy. It is 
reported as introduced only in Sardegna (Raab-Straube, 
2014+ in Euro+Med PlantBase). In North Africa, the tax-
on was cited as introduced without any precision about 
its actual status or doubtfully present in Morocco and Tu-
nisia (POWO, 2022c; APD, 2022c).
Habitat in Tunisia (Figure 4): Pinus pinaster subsp. 
pinaster is found within coastal kermes oak (Quercus coc-
cifera L.) in the region of Nadhour (Les Grottes), within 
Bizerta governorate (NE Tunisia). It was introduced since 
many decades and is extending with many individuals 
(ca. 50) of different ages and diameter from seedlings. 
Notes: The taxonomy of Pinus pinaster s.lat. has not 
been thoroughly studied and is still debated; however, 
three sub-species are generally recognized (Farjon, 1998), 
which show small differences related mainly to the anat-
omy of the leaves and to the length of leaves and seed 
cones.
Updated key to species of the genus Pinus 
Subsection Pinaster3 in Tunisia 
1a. Fascicle sheaths of new leaves 20–30 mm long; leaves 
thick, 1.5–2 mm wide; seed cones 10–20(–22) cm 
long, persistent, opening slowly; umbos on scales 
forming a central, rigid spine  .................... P. pinaster
1b. Fascicle sheaths of new leaves shorter than 20 mm; 
leaves 0.7–1.5(–1.8) mm wide; seed cones shorter 
than 13(–15) cm, deciduous or persistent; umbos un-
armed or with a minute prickle  .............................. 2
2a. Bark on trunk breaking into large plates; fascicle 
sheaths of new leaves short, to ca. 10 mm long; seed 
cones broadly ovoid to subglobose when closed; seeds 
15–20(–22) mm long, thick, with a rudimentary 
wing much shorter than the seed ...................P. pinea
2b. Bark on trunk breaking into small or elongated plates; 
fascicle sheaths of new leaves (9)10–20 mm long; seed 
cones ovoid-conical or oblong-conical when closed; 
seeds less than 10 mm long, with a wing at least 2 × 
the length of the seed .............................................. 3
3a. Leaves in fascicles of only 2, (6–)7–10(–12) cm long, 
0.7–0.8 mm wide ................................... P. halepensis
3b. Leaves in fascicles of 2 or sometimes 3, (5–)10–18(–
29) cm long, 1–1.5 mm wide ..................... P. brutia
Subfam. 3. Laricoideae (Rendle) Pilg. & Melch. (1954).
Genus 3. Pseudotsuga Carr., Traité Conif., ed. 2: 256 
(1867).
Pseudotsuga menziesii (Mirb.) Franco var. glauca 
(Beissn.) Franco, Conif. Duarum Nom.: 6. 1950 [& 
Bol. Soc. Brot., ser. 2, 24: 76. 1950]. 
Syn. Tsuga douglasii (Sabine ex D. Don) Lindl. var. glauca 
Beissn., in Jäger & Beissner, Ziergeh. Gärt. Park., ed. 2: 
446. 1884; Pseudotsuga menziesii (Mirb.) Franco subsp. 
glauca (Beissn.) E. Murray, Kalmia 12: 24. 1982.
First official reports of the genus and the species as es-
tablished taxa for the flora of North Africa.
A small tree; trunk monopodial, straight, columnar; 
bark on large trunks rough and scaly; branches spreading 
horizontally; branchlets slender, flexible; leaves spreading 
3 In this subsect. many plantations of Pinus canariensis C. Sm. are intro-
duced and cultivated with hundreds of individuals within Northern Tuni-
sia mainly and could more probably naturalizing from seedlings!
Figure 4: Pinus pinaster subsp. pinaster in Tunisia. A: summital crown 
part with leaves, juvenile and mature seed cones; B: mature close-up 
seed cones. (Bizerta, NE Tunisia), Photographs R. El Mokni.
Slika 4: Pinus pinaster subsp. pinaster v Tuniziji. A: vršni del krošnje z 
listi, juvenilnimi in zrelimi semenskimi storži; B: zreli semenski storži, 
bližnji pogled. (Bizerta, SV Tunizija), fotografije R. El Mokni.
A
B
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more or less radially, (1.5–)2.5–3.5(–4) cm long, 1.2–
1.5(–1.7) mm wide, more or less petiolate and slightly 
twisted at base, linear, straight, flattened, obtuse or acute 
at apex; pollen cones 1.5–2 cm long, yellow, with red 
brown perular scales at base; seed cones pendulous on 0.5–
1 cm long peduncles, deciduous, ovoid-conical, obtuse or 
pointed at apex, 4–9(–10) cm long, 2–4 cm wide with 
opened scales, green, with yellowish green bracts, ripen-
ing to brown or dull grey brown, with light brown bracts. 
Seed scales obovate-cuneate, convex, 2–2.5 × 2–2.5 cm 
at mid-cone; base more or less cuneate; bracts ligulate, 
trilobate at apex, with cusp longer than lateral lobes, 2.5–
3.5 cm long, exserted, not reflexed, often curved towards 
cone apex; seeds ovoid-cuneate, 6–8 × 4–5 mm, light 
brown, often with dark spots; wings ovate oblong, 9–14 × 
5–8 mm, yellowish brown.
Distribution: the species is native to North America 
with two subspecies: the coastal Douglas fir (Pseudotsuga 
menziesii var. menziesii) occurs from British Columbia 
southward along the Pacific Coast to central California; 
and the Rocky Mountain Douglas fir (Pseudotsuga men-
ziesii var. glauca) occurs from central British Columbia 
along the Rocky Mountains into the mountains of central 
Mexico (Farjon, 2010; Hermann & Lavender, 1990). The 
first seeds were introduced in Europe by David Doug-
las in 1827 and then planted at Drop more Park (Buck-
inghamshire, UK), where there is a tree which is usually 
considered the oldest Douglas fir of Europe (Elwes & 
Henry, 1909). Initially planted as ornamental, Douglas 
fir started to be used as a forest species by the end of the 
nineteenth century. This fir became a major reforestation 
species in Western Europe after the Second World War, 
mainly with the support of national or regional forest 
grants. In Europe, 80% of the total Douglas fir area is to 
be found in three countries: France (half of the European 
area), Germany and United Kingdom. Outside Europe, 
Douglas fir has also been introduced in several countries 
of the southern hemisphere (South Africa, South Amer-
ica, New Zealand and Australia) (Bastien & al., 2013) 
and also in Turkey as an important fast-growing tree spe-
cies for industrial plantations (with a limited distribution 
of approximately 140 ha). In North Africa, no report for 
the genus (Raab-Straube, 2014+ in Euro+Med PlantBase; 
POWO, 2022d). 
Habitat in Tunisia (Figure 5): Pseudotsuga menziesii 
subsp. glauca is found (ca. 30 individuals) within mixed 
oak forests (Quercus canariensis Willd. and Q. suber L.) in 
the region of Tabarka (Majen Roumi) within Jendouba 
governorate (NW Tunisia) where it seems introduced 
since colonization and show many juvenile individuals of 
different heights and diameter at breast height growing 
from seedlings.
Notes: the variety glauca remains a smaller tree than the 
var. menziesii (up to 100 m tall) and has usually shorter, 
wider (1.5–2.5(–3.5) cm × 1.4–1.7 mm) and more glau-
cous green leaves with usually smaller seed cones [vs. dark 
green leaves (2–)2.5–3.5(–4) × 1.2–1.5 mm and seed 
cones 4–9(–10) cm long].
A
B C
Figure 5: Pseudotsuga menziesii var. glauca in Tunisia. A: habit of some 
trees in their habitat within Kroumirian forests; B: detailed part of the 
bark trunk; C: detailed of mature seed cone with typical ligulate, trilo-
bate bracts at apex, longer than lateral lobes. (Tabarka, NW Tunisia), 
Photographs R. El Mokni. 
Slika 5: Pseudotsuga menziesii var. glauca v Tuniziji. A: habitus dreves 
na njihovem rastišču v Kroumirskih gozdovih; B: podrobno prikazan 
del lubja debla; C: detajl zrelega semenskega storža z značilnimi 
jezičastimi, trilobatimi ovršnimi lističi na vrhu, daljši od stranskih 
režnjev. (Tabarka, SZ Tunizija), fotografije R. El Mokni.
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Updated key to the genera of the Pinaceae 
family in Tunisia
1a. Adult green leaves bundled with 2–5(-8) together on 
dwarf shoots (1 species with a single leaf on a dwarf 
shoot); seed cones biennial (rarely triennial), with dis-
tinction between each year’s growth apparent as an 
umbo and apophysis on each scale; seed held to the 
wing in a pair of claws ...................................... Pinus
1b. Adult green leaves either solitary or in pseudowhorls 
of more than 10 together on short shoots; seed cones 
mostly annual (if biennial without umbo); seed held 
to the wing in a cup ................................................ 2
2a. Seeds without resin vesicles; seed scales with a 
broad basis, persistent, mature seed cones spreading 
or pendulous from mostly plagiotropic branches; 
leaves equally distant on all (segments of ) shoots 
bract scales of cones large; exserted and trilobate at 
apex .......................................................Pseudotsuga
2b. Seeds with resin vesicles; seed scales with a narrow, 
petiolate basis, persistent or deciduous; leaves spirally 
and remotely arranged on long shoots, in dense pseu-
do-whorls on short shoots; mature seed cones erect 
maturing in two years, disintegrating on tree with de-
ciduous seed scales ........................................  Cedrus 
ORDER 2. CUPRESSALES Link, Handbuch 2: 470 
(1829).
Family. Cupressaceae Gray, Nat. Arr. Brit. Pl. 2: 222. 
(1822), nom. Co
Subfam. 1. Taxodioideae Endl. ex K.Koch
Genus 1. Cryptomeria D.Don, Ann. Nat. Hist. 1: 233 
(1838).
Cryptomeria japonica (Thunb. ex L. f.) D. Don, Trans. 
Linn. Soc. London 18: 167. 1839. 
Syn. Cupressus japonica Thunb. ex L. f., Suppl. Pl.: 421. 
1781. 
First official reports of the genus and the species as 
 established taxa for the flora of North Africa.
Figure 6: Cryptomeria japonica in Tunisia. A: habit of some tree in their habitat within Kroumirian forests; B: part of the bark trunk fully covered 
by lichens; C: outspread densely arranged branches with decurrent, linear-subulate leaves; D: aggregated terminal close-up seed cones; E: dehiscent 
globose, squarrose seed cones with typical cuneate scales providing small teeth above the recurved bract tip. (Tabarka, NW Tunisia), Photographs 
R. El Mokni.
Slika 6: Cryptomeria japonica v Tuniziji. A: habitus dreves na njihovem rastišču v Kroumirskih gozdovih; B: del lubja v celoti prekrit z lišaji; 
C: razprte, gosto razporejene veje s padajočimi, črtasto šilastimi listi; D: končni semenski storžki; E: razpršeni kroglasti, kvadratasti semenski storžki 
s tipičnimi klinastimi luskami, ki tvorijo majhne zobce nad ukrivljeno konico ovršnega lista. (Tabarka, SZ Tunizija), fotografije R. El Mokni.
A B
C
D E
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A
B C
D
Tree, 30–40(–60) m high, evergreen, monoecious; 
trunk straight, slender, usually not over 1–2 m in diam-
eter; branches densely arranged, outspread or ascending, 
young branches green, glabrous, forming a conical crown 
in young trees and a rounded crown in mature trees; leaves 
helically arranged in ranks of 5, decurrent, spreading but 
incurved in various degrees, directed forward, linear-sub-
ulate, slightly flattened laterally, distinctly keeled abaxi-
ally, stiff, green, 3–20(–25) × 1–2 mm, margins entire, 
apex acute; pollen cones numerous, axillary and crowded 
towards ends of 2nd-year branchlets, 3–6 × 2–3 mm, 
elongating to 10 mm at anthesis; seed cones terminal on 
down-curved branchlets with normal leaves, often aggre-
gated or solitary, occasionally with proliferating vegetative 
short shoot at apex, globose to subglobose, squarrose with 
spreading bract-scale complexes, soft woody, 12–20(–25) 
mm in diameter; seeds red-brown, (1–)2–5 per bractscale 
complex (some ovules may abort), 4–5 × 3 mm, flattened, 
irregularly ovate with 2 wings; wings unequal, 1–1.5 mm 
wide, forming a strip around the seed.
Distribution: Cryptomeria japonica is an endemic and 
widely distributed species in Japan. Locally it is the most 
important commercial forestry species. It has been grown 
extensively also in forestry plantations in China and 
has naturalized in some areas there. It is considered as 
a popular ornamental tree in many temperate countries 
and in different parts of the world including India 
(Maity  &  Moktan,  2019). Sanbu-sugi, mainly planted 
in Sanbu district and Chiba Prefecture, is one of the best 
logging cultivars of C. japonica (cf Miyajima, 1973).
Habitat in Tunisia (Figure 6): Cryptomeria japonica is 
found within mixed oak forests (Quercus canariensis Willd. 
and Q. suber L.) in the surroundings of Tabarka within 
Jendouba governorate (NW Tunisia) where it seems intro-
duced since colonization and show more than 25 individu-
als of different heights and diameter at breast height.
Genre 2. Cupressus L., Sp. Pl. 2: 1002. 1753. 
Cupressus arizonica Greene var. glabra (Sudw.) Little, 
Madroño 18: 162. 1966. 
Syn. C. glabra Sudw., Amer. Forestry 16: 88. 1910; Cal-
litropsis glabra (Sudw.) D. P. Little, Syst. Bot. 31 (3): 
473. 2006 (nom. ut. rej., Art. 56); Hesperocyparis glabra 
(Sudw.) Bartel, Phytologia 91 (1): 181. 2009. 
First official report as a casual taxon for the flora of 
 Tunisia.
Large shrubs or trees 5–20(–30) m tall; trunk monopo-
dial or multistemmed from near the ground; bark smooth 
on branches and younger trees, soon turning grey, then 
exfoliating with thin scales and strips, exposing reddish 
bark remaining smooth well into maturity of trees, even-
Figure 7: Cupressus arizonica var. glabra in Tunisia. A: habit of a tree 
in its habitat within Kroumirian forests; B: detailed part of the bark 
trunk; C–D: mature close-up seed cones (Fernana, NW Tunisia), 
Photographs R. El Mokni. 
Slika 7: Cupressus arizonica var. glabra v Tuniziji. A: habitus drevesa 
na njegovem rastišču v Kroumirskih gozdovih; B: podroben del lubja 
na deblu; C–D: zreli semenski storži od blizu (Fernana, SZ Tunizija), 
fotografije R. El Mokni.
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tually near base of large trunks becoming fissured, up to 
1 cm thick, exfoliating with small flakes; branches long 
and spreading, more often ascending or erect, forming 
a conical, broad conical, pyramidal, or oval and round-
topped, dense crown; branchlets numerous, stout; leaves 
usually conspicuously and actively glandular; pollen cones 
numerous, terminal, solitary, subglobose, elliptic-oblong 
or cylindrical, 3–5 × 1.7–2.2 mm; seed cones terminal, of-
ten grouped or clustered, subglobose to irregularly broad 
ovoid, 15–27(–30) mm, while growing with prominent 
bosses large or small on the scales; seeds red-brown, not 
glaucous or slightly glaucous. 
Distribution: the taxon is native to USA: Arizona (Co-
conino, Gila, Maricopa and Yavapai Co.), Mexico North-
east, Mexico Northwest, New Mexico, Texas. It is re-
ported as introduced in Argentina Northeast, Free State, 
Italy, Turkey, Turkey-in-Europe (POWO, 2022e) whereas 
Raab-Straube (2014+) reported the taxon as cultivated in 
Algeria, Azerbaijan, with Nakhichevan, Italy, Morocco, 
Portugal, Romania, Sardinia, Spain, with Gibraltar and 
Andorra.
Habitat in Tunisia (Figure 7): Cupressus arizonica 
Greene var. glabra is found within an opened cork oak 
forest (Quercus suber L.) where many coniferous taxa 
were planted in the region of Fernana (Aîn Debba) with-
in Jendouba governorate (NW Tunisia) where it seems 
introduced since colonization and show more than 7 
individuals of different heights and diameter at breast 
height.
Notes: the variety glabra is distinguished from the other 
varieties [var. arizonica, var.  nevadensis  (Abrams) Lit-
tle, var. stephensonii (C.B. Wolf ) Little and var. montana 
(Wiggins) Little] of the Cupressus arizonica complex by 
having mainly seed cones 2–2.5 (–3.2) cm in diameter (vs. 
1.5–2.5 cm in var. montana, (1.5–)2–3(–3.5) cm in var. 
nevadensis, ( 1–)2–2.5 cm in var. stephensonii and (1.2–
)2–3 cm in var. arizonica) remaining closed at maturity 
(vs. opening at maturity for the var. montana) and seeds 
(3–)4–5(–8) mm long, with minute wings, usually waxy 
(vs. (3–)4–5(–6) mm long, with wings 1 mm wide, usually 
not waxy in var. arizonica, 3–4(–5) mm long, with wings 
1–5 mm wide, usually not waxy in var. montana, 3–6 mm 
long, with wings to 2 mm wide, occasionally slightly 
waxy in var. nevadensis and (4–)5–8 mm long, with wings 
1–3 mm wide, usually waxy in var. stephensonii).
Updated key to species of the genus 
Cupressus in Tunisia 
1a. Seed cones usually in dense, serotinous clusters; pol-
len sacs often 5 or more per microsporophyll; glands 
present on scale leaves; seed cones spherical or a little 
elongated, (1.5–)2–3 cm long, gray or brown at ma-
turity, often waxy before this, with 3 or 4(–6) pairs of 
seed scales; seeds (5–)8–15(–20) per scale, (3–)4–6(–
8) mm long, light to dark brown .......... C. arizonica
1b. Seed cones solitary or in groups, not densely clustered; 
pollen sacs up to 4 per microsporophyll; glands on all 
scale leaves; Seed cones usually oblong, (1.5–)2–3(–4) 
cm long, brown at maturity, not waxy; with 3–5 or 
6 or 7 pairs of seed scales; seeds (1–)8–20(–25) per 
scale, 4–7 mm long, reddish brown ... C. sempervirens
 
Genus 3. Taxodium Rich., Ann. Mus. Natl. Hist. Nat. 
(Paris) 16: 298. 1810.
Taxodium distichum4 (L.) Rich., Ann. Mus. Natl. Hist. 
Nat. (Paris) 16: 298. 1810.
T. distichum (L.) Rich. var. distichum
Syn. Cupressus disticha L., Sp. Pl. 2: 1003. 1753. 
First official reports of the genus and the species as nat-
uralized taxon for the flora of Tunisia.
A winter deciduous tall tree, to 40(–46) m tall; trunk 
monopodial, erect; base often swollen, fluted or but-
tressed; ‘pneumatophores’ or ‘knees’ common, numer-
ous on root systems of trees growing in or near water, 
conical, 1(–4) m tall; bark on trunk fissured, exfoliating 
in more or less fibrous, long strips, light brown, turning 
grey; branches spreading or ascending, forming a conical 
or pyramidal crown in young trees, old trees becoming in-
creasingly flat-topped with long main branches in upper 
half of tree; short shoots spreading in two ranks, or in 5–8 
ranks, mostly persistent but with occasionally assurgent 
or nearly erect branchlets; leaves linear, flattened, pecti-
nately arranged, twisted near base, 10–15(–22) mm long, 
straight or slightly curved, obtuse or mucronate; pollen 
cones solitary or in pairs, globose to ovoid-oblong, 3–5 
× 2–3 mm, yellowish green turning purplish to brown; 
microsporophylls (5–)6–10(–15), ovate-peltate, keeled, 
bearing (2–)4–9(–10) globose pollen sacs; seed cones often 
clustered, subglobose or more or less ovoid but irregu-
lar, (15–)20–35(–40) mm long; seeds ca. 20–40 per cone, 
4–7 mm long, lustrous brown, with a large, (sub)lateral, 
pale coloured hilum; wings very small or vestigial, form-
ing up to 3 narrow ridges of ca. 1 mm wide.
Distribution: the species is indigenous to South-East 
USA: Alabama, Arkansas, Delaware, Florida, Georgia, Il-
linois, Indiana, Kentucky, Louisiana, Maryland, Missis-
sippi, Missouri, North Carolina, Oklahoma, South Caro-
lina, Tennessee, South Texas and Virginia.
4 The same taxon is known in Algeria within the National Park of El Kala 
since many decades but any indication if casual or naturalized is reported!
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New records of coniferous species for the non-native woody flora of Tunisia and 
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Figure 8: Taxodium distichum var. distichum in 
Tunisia. A: habit of some trees in their habitat 
within Kroumirian forests; B: detailed part of the 
bark trunk; C: branchlets with linear, flattened, 
pectinately arranged leaves; D: clustered, subglo-
bose mature close-up seed cones; E: habit of some 
tree in the winter season. (Tabarka, NW Tunisia), 
Photographs R. El Mokni. 
Slika 8: Taxodium distichum var. distichum v Tuniz-
iji. A: habitus nekaterih dreves na njihovem rastišču 
v Kroumirskih gozdovih; B: podroben del lubja 
debla; C: veje s črtastimi, sploščenimi, glavničasto 
razporejenimi listi; D: grozdasti, subkroglasti zreli 
semenski storži od blizu; E: habitus drevesa v 
zimski sezoni. (Tabarka, SZ Tunizija), fotografije 
R. El Mokni.
A
C
D
B
E
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New records of coniferous species for the non-native woody flora of Tunisia and 
North Africa
Habitat in Tunisia (Figure 8): Taxodium distichum 
var. distichum is found within lowland river flood plains 
and swamps in the surroundings of the region of Tabar-
ka within Jendouba governorate (NW Tunisia) where it 
seems that it has been introduced since colonization and 
show many individuals (ca. 600) of different heights and 
diameter at breast height.
Notes: the var. distichum can be easily distinguished 
from the var. imbricarium (Nutt.) Sarg. by the presence 
of numerous ‘Pneumatophores’ or ‘knees’ [vs. uncom-
mon and if present more rounded (not conical)], its lin-
ear, flattened leaves pectinately arranged, twisted near 
base, 10–15(–22) mm long, straight or slightly curved, 
obtuse or mucronate (vs. largely appressed but with free 
apices leaves, not twisted at base, short acicular (subulate) 
to narrowly lanceolate, 3–10 mm long, keeled, acute or 
pungent in the var. imbricarium).
Subfam. 2. Cupressoideae Rich. ex Sweet
Genus 1. Platycladus Spach, Hist. Nat. Vég. Phan. 11: 
333. 1841. 
Platycladus orientalis (L.) Franco, Portugaliae Acta 
Biol., sér. B, Sist. Vol. “Julio Henriques”: 33.1949. 
Syn. Thuja orientalis L., Sp. Pl. 2: 1002. 1753, Biota ori-
entalis (L.) Endl., Syn. Conif.: 47. 1847.
First official reports of the genus and the species as cas-
uals to naturalizing taxa for the flora of Tunisia.
Trees to 20–25 m tall, evergreen, monoecious; trunk 
monopodial or multistemmed from low above ground, 
with stem(s) erect or ascending; bark on large stems be-
coming fibrous, exfoliating in thin, long sheets; branches 
spreading, ascending or erect, contorted in old trees, 
higher order branches mostly spreading, forming a py-
ramidal, rounded or irregular crown; leaves decussate, de-
current, scale-like, on lateral branchlets dimorphic, facials 
more or less rhombic to obtrullate, with appressed, obtuse 
apex, green or greyish green; pollen cones terminal on ul-
timate branchlets, solitary, subglobose, 2–3 × 2 mm; mi-
crosporophylls (6–)8–10(–12), decussate, peltate, bearing 
3–6 abaxial pollen sacs; seed cones terminal, ampulliform, 
glaucous or purplish cones 15–20(–25) × 10–18 mm 
H G
F
Figure 8 (continued): Taxodium distichum var. distichum in Tunisia. F: dehiscent seed cone; G: pneumatophores or knees; H: pollen
cones yellowish green turning purplish to brown (Tabarka, NW Tunisia), Photographs R. El Mokni.
Slika 8 (nadaljevanje): Taxodium distichum var. distichum v Tuniziji. F: razpadli semenski stožec; G: dihalne korenine ali kolena; H: pelodni storži 
rumenkasto zeleni, ki se spreminjajo v vijolično do rjavo (Tabarka, SZ Tunizija), fotografije R. El Mokni.
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New records of coniferous species for the non-native woody flora of Tunisia and 
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Figure 9: Platycladus orientalis 
in Tunisia. A: habit of a tree in 
its habitat; B: detailed part of 
the bark trunk; C: leaves with 
terminal, solitary pollen cones; 
D: terminal, ampulliform, 
glaucous seed cones; E–F: mature 
purplish seed cones. (Monastir, 
NE Tunisia), Photographs  
R. El Mokni. 
Slika 9: Platycladus orientalis v 
Tuniziji. A: habitus drevesa na 
njegovem rastišču; B: podroben 
del lubja debla; C: listi s 
končnimi, posamičnimi 
pelodnimi storži; D: terminalni, 
stekleničastimi, sivkastimi 
semenski storži; E–F: zreli 
škrlatni semenski storži. 
(Monastir, SV Tunizija),  
fotografije R. El Mokni.
A B
C D E F
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New records of coniferous species for the non-native woody flora of Tunisia and 
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which open widely and turn brown when the seeds have 
ripened; seeds 6–10(–12), ovoid to narrowly ovoid, 5–7 × 
3–4 mm, greyish brown with a large, light brown hilum; 
wings absent or rudimentary.
Distribution: the species is indigenous to China 
(S Gansu, Hebei, Henan, Shaanxi, Shanxi, Nei Mongol 
[Inner Mongolia]), Korea, and Russian Far East. It seems 
to be the most widely introduced cupressaceous conifer 
in Asia. In many areas inside and outside China it has 
‘escaped’ from cultivation and established spontaneous 
populations. In Europe, the plant was reported as natu-
ralized only in Austria and Italy whereas it is casual in 
Czech Republic, Germany, Spain and Portugal. In North 
Africa, it is cited as cultivated only in Morocco and Alge-
ria (Raab-Straube, 2014+; APD, 2022d). 
Habitat in Tunisia (Figure 9): Platycladus orientalis is 
found within borders of ornamental gardens and road-
sides in the regions of Monastir and Mahdia (NE Tunisia) 
where many juvenile and shrubby individuals (ca. 50) of 
different heights were observed not far from adult culti-
vated specimens.
Notes: Despite that both genera show seed cones with 
3–6 decussate pairs of fertile and sterile scales. The genus 
Thuja shows thin seed cone scales, the distal pair reduced 
and connate; seeds winged whereas the genus Platycladus 
shows thickened seed cone scales with well-developed and 
spreading distal pair and unwinged seeds.
Updated key to the genera of the 
Cupressaceae family in Tunisia
1a. Leaves on lateral branchlets and cone scales helically 
arranged; leaves (almost) all linear, flat or short ac-
icular to subulate in one variety; lateral branchlets 
(semi)deciduous; pollen cones numerous on leafless 
branchlets; seed cones on thicker branches, often ag-
gregate ..................................................... Taxodium
1b. Leaves on lateral foliage branchlets scale-like or subu-
late or lanceolate, or a mixture of scale-like and lin-
ear leaves present on the same mature plant, lateral 
branchlets evergreen................................................ 2 
2a. Leaves on lateral branchlets scale-like, appressed, or 
subulate or lanceolate; seed cones squarrose; cone 
scales cuneate, with small teeth above the recurved 
bract tip ................................................Cryptomeria
2b. Leaves on lateral, short branchlets at least in part scale-
like or subulate, or a mixture of scale-like and linear 
leaves present on the same mature plant; seed cones not 
squarrose; cone scales decussate, opposite or whorled ...3 
3a. Leaves scale-like or acicular-linear; seed cones berry-
like, ± soft, indehiscent; cone scales (almost) entirely 
fused ..........................................................  Juniperus
3b. Leaves on lateral branchlets decussate, scale-like, 
rarely acicular or linear; seed cones not berry-like, 
(eventually) dehiscent; cone scales at least partly 
integer ...................................................................  4
4a. Leaves on lateral branchlets decussate, monomorphic 
or at least not clearly divided in facials and laterals; 
seed cones globose to ovoid, with > 6 peltoid scales; 
seeds numerous, small, winged ..................  Cupressus
4b. Leaves decurrent, dimorphic; seed cones (sub)globose 
or ampulliform, with ≤ 6 scales; seeds ≤ 12, wings 
sometimes absent or rudimentary ........................... 5
5a. Leaves weakly dimorphic, appressed, long decur-
rent, adnate to branchlets except for the small apex; 
seed cones (sub)globose when closed, woody with 4(-
5–6) subpeltate scales in decussate pairs; seeds 4–6 
winged ......................................................Tetraclinis
5b. Leaves decurrent, dimorphic, facials more or less 
rhombic to obtrullate, with appressed, obtuse apex, 
laterals bilaterally flattened, distal part spreading or 
reflexed, appressed or free, with incurved, obtuse 
apex; seed cones ampulliform with 3–6 decussate 
pairs of fertile and sterile thickened scales, the 
distal pair well developed and spreading; seeds up 
to 12 unwinged or more rarely with rudimentary 
wings .....................................................Platycladus
Key to the families of the Pinopsida Class in 
Tunisia 
1. Seed cones with seed scales in the axils of bracts (bracts 
can be much reduced in mature, woody cones but are 
conspicuous in immature cones at early stages of devel-
opment); cones clearly compound and never reduced; 
seeds two on the adaxial (upper) side of each fertile 
scale; adult green leaves acicular-linear ..........Pinaceae
2. Seed cones with seed scales fused with bracts (bracts 
make up the bulk of the cone), or with bracts only 
(which may be much enlarged, swollen and/or woody 
at maturity), or with scales obscure, much reduced or 
absent; seeds either single or more than two per fertile 
scale; adult green leaves scale-like, or acicular, or with 
a distinct lamina, or replaced by phylloclades (phyl-
lodes); seed cones with seed scales fused with bracts, 
or with bracts only which are much enlarged and of-
ten swollen at maturity and may then form a compact, 
globose cone; seed cones consisting of bracts forming 
the cone scales only (sometimes with rudimentary seed 
scales only visible at very early stages of development); 
seeds 1-many, axillary or on the base of each bract 
(rarely only a single seed per cone); true leaves scale-
like or acicular (needle-like), cataphylls absent, phyl-
lodes absent .......................................... Cupressaceae
23/1 • 2024, 119–136
134
Ridha El Mokni
New records of coniferous species for the non-native woody flora of Tunisia and 
North Africa
Discussion 
The list of coniferous species here presented is only 
a starting point for Tunisia and in part for North Af-
rica. Further surveying and assessment of the status of 
woody alien plants is required for many regions within 
the Tunisian country and/or within the extended area 
of the northern African countries. Subspecies (escarena 
and pinaster) of Pinus pinaster were discovered with very 
extended populations as well-established aliens in many 
localities (in the northeastern and the northwestern) of 
the country. These taxa seem to have escaped by different 
means of dissemination since long time ago but not re-
ported due perhaps to lack of investigations, or could be 
overlooked (vegetative resemblance with the autochtho-
nous subspecies, renoui) either misidentified. Moreover, 
among the recorded taxa, Pinus brutia var. pityusa can 
be considered especially dangerous since invasiveness has 
been previously observed and assessed more recently in 
other regions mainly within the northwestern part of Tu-
nisia. A prompt invasion status could be assigned in the 
nearer future to Taxodium distichum in the marshy areas 
of the northwest since actual surveys show active dis-
persal in new localities ca. 15 km far away the historical 
know locality. The remaining taxa (mainly Cryptomeria 
japonica, Cupressus arizonica var. glabra, Pinus coulteri 
and Pseudotsuga menziesii var. glauca) can be considered 
less-threatening up to now, according to our field obser-
vations. However, invasive status may vary with time, 
and taxa could modify their ecological behavior espe-
cially with the global changing of environmental con-
ditions (Richardson & Bond, 1991; Willis et al. 2010). 
Consequently, many alien species already present in an 
area and currently deemed ‘safe’ (non-invasive) may well 
become invasive.
Conclusion
Novelties presented in this study within coniferous 
group (Gymnospermae sensu lato) enhance the evidence 
of the ongoing process of introductions and naturaliza-
tion of alien (introduced trees and shrubs) plants and 
ornamentals in Tunisia and North Africa, as in many 
parts of the World. Data here firstly presented is mainly 
related to woody species of the Pinidae group escaping 
specially from botanical gardens (arboreta) and nurseries’ 
surroundings. The increase of field surveys with a proper 
identification allow a better understanding of pathways of 
introductions and vectors of invasiveness for both recent 
and old escapes and help in reducing the impacts of these 
introduction on native plant communities and invaded 
habitats.
Acknowledgments
Thanks are due to all colleagues and friends of the Re-
gional Agricultural Development Commission (RADC) 
of Jendouba and Aîn Draham, forest services of Ghar 
Dimaou (El Feidja, Aîn Sultan and El Ghorra), Fernana 
and Aîn Draham (Aîn Draham, Aîn Sellam, Aîn Zana, 
Tabarka and Babouch) for their technical support and 
their help during my botanical surveys within Kroumiria 
region since 2001. I’m so grateful to my friend and col-
league Dr. Mohsen Kalboussi (Herpetologist, Sousse Uni-
versity) with whom I share the discovery of many taxa 
during our field prospections. I’m so grateful to anony-
mous reviewers for their useful comments and sugges-
tions given to a first version of the manuscript. 
Ridha El Mokni  https://orcid.org/0000-0003-3849-1039
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