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Summary: Botanical surveys undertaken in Cen-tral and Northern Tunisia (North Africa), mostlybetween 2012 and 2018, have revealed first recordsand/or new distributional records for non-nativesucculents (sensu lato) belonging to the Cactaceaefamily. Some can be considered naturalised or evenpotentially invasive [e.g. Opuntia dillenii (KerGawl.) Haw.], whereas others are here reported forthe first time for North Africa, either as recentlynaturalised or casual, e.g.: Echinopsis eyriesii(Turpin) Zuccarini, Opuntia dejecta Salm-Dyck, O.lindheimeri Engelm., O. monacantha Haw and O.robusta J.C. Wendl. ex Pfeiff. Short descriptions aregiven for each of these species, comments on theiractual status were pointed out and field photo-graphs are provided.

Zusammenfassung: Botanische Untersuchungen inZentral- und Nordtunesien (Nordafrika), haupt-sächlich zwischen 2012 und 2018, haben ersteAufzeichnungen und/oder neue Beobachtungen fürnicht-einheimische Sukkulenten (sensu lato) ausder Cactaceae-Familie ergeben. Einige können alseingebürgert oder sogar potentiell invasiv angese-hen werden [als Opuntia dillenii (Ker Gawl.)Haw.], während andere hier zum ersten Mal fürNordafrika berichtet werden, entweder vor kurzemeingebürgert oder vorübergegend: Echinopsis ey-riesii (Link) Zucc. ex Pfeiff. & Otto, Opuntia dejectaSalm-Dyck, Opuntia lindheimeri Engelm., O.monacantha Haw und O. robusta J. C. Wendl. exPfeiff. Für jede dieser Arten werden kurze

Beschreibungen gegeben, Kommentare zu ihremaktueller Status werden angegeben und Fotos wer-den bereitgestellt.

Introduction Succulents are found across the globe in nearly

all types of habitat, but most often in arid or semi-arid parts of the world. They are specially adaptedto deal with dry, desert-like conditions, and able tostore water in one or more of their organs; theirleaves, stems, or roots are often filled with water-storing tissue and are thus unusually fleshy andenlarged. The Cactaceae  family is distinguishedfrom the other succulent plant families (i.e.Agavaceae, Aloaceae, Euphorbiaceae, etc.) by thepresence of small rounded woolly areas normallylocalized along the ribs of the plant (known as are-oles) from which new segments, flowers and spinesare developing (retrieved from https://www.iucn.org/ssc-groups/plants-fungi/cactus-and-succulent-plants-specialist-group/what-are-cacti-and-succu-lents). Cactus roots are generally less specialized inmorphology than the above-ground vegetativeparts. They have a shallow distribution, which en-ables them to exploit limited rainfall, and are richin root hairs. Associations with fungi and bacteriacan improve the capture of mineral nutrients(Dubrovsky & North, 2002; Land Protection, 2006).

Recent classifications of Cactaceae recognizefour subfamilies (Anderson, 2001; Wallace & Gib-son, 2002; Nyffeler, 2002) including Cactoideaewith 1,222 species (the highest diversity) where

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New records of cacti (Opuntioideae & Cactoideae, Cactaceae) fromTunisia

Ridha El Mokni1,2,3, Filip Verloove4, Alessandro Guiggi5 & Mohamed HédiEl Aouni11. University of Carthage, Laboratory of Botany and Plant Ecology (SNA-214), Department of Life Sciences, Faculty of Sciences of Bizerta, Jarzouna, 7021, Bizerte, Tunisia. (email: riridah@hot-mail.com)2. University of Jendouba, Laboratory of Silvo-Pastoral Resources, Silvo-Pastoral Institute ofTabarka, BP. 345, 8110-Tabarka, Tunisia.3. University of Monastir, Laboratory of Botany, Cryptogamy and Plant Biology, Department of Pharmaceutical Sciences “A”, Faculty of Pharmacy of Monastir, Avenue Avicenna, 5000-Monastir,Tunisia.4. Botanic Garden of Meise, Nieuwelaan 38, B-1860 Meise, Belgium.(email: filip.verloove@botanicgardenmeise.be) 5. DISTAV, Polo Botanico, Università degli Studi di Genova, International Cactaceae Research Cen-ter (ICRC), Corso Dogali, 1M - 16136 Genova, Italy. (email: alex.guiggi@libero.it)

Photographs: Ridha El Mokni.

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species have stems with ribs or tubercles and aroleswithout glochids (Gibson & Nobel, 1986) and Op-untioideae with about 186 species where speciesare articulated cacti with flat, cylindrical or globu-lar stem-segments (Gibson & Nobel ,1986; Ter-razas-Salgado & Mauseth, 2002).

The first reports of the cultivation of cacti in Eu-rope date back to the 1500’s, following their intro-duction from the Americas (Howard & Touw, 1981)and soon became common in European collectionsand gardens (Anderson, 2001). Today, Europe pro-duces millions of propagated cacti per year from itshorticultural industry. However, there remains apersistent demand for species collected from thewild (McGough et al., 2014). Thus, many introducedsucculents have been reported as alien species (seee.g., Guillot & van der Meer, 2004; Guillot & Lodé,

2009; Guiggi, 2008, 2010, 2014; Guillot et al., 2014;Verloove, 2016; Verloove et al., 2017a & b). InNorth Africa, very few papers have dealt with theinventory of Cactaceae (Véla, 2013), hence the orig-inality of the present work where 12 taxa are pre-sented and illustrated, most of them not previouslyreported for the non-native Tunisian or even NorthAfrican flora. Several species, e.g. Opuntia dejectaSalm-Dyck, Opuntia lindheimeri Engelm. O.monacantha Haw. and O. robusta H.L. Wendl. exPfeiff. are here reported for the first time fromNorth Africa.

Materials and MethodsField work carried out by the first and the

fourth authors (pro parte) within Central andNorthern Tunisia (North Africa), mostly between

Figure 1. Austrocylindropuntia subulata, A. Monastir city, coastal area, 3rd January 2017; B. Monastir city,coastal area, clayey slope, 30th May 2019; C. & D. Monastir city, coastal area, clayey slope, 12th June 2018.

A B

C D

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2012 and 2018, has revealed new national and evenNorth African records. All records here reported aredocumented by short descriptions of each species.The actual degree of naturalization for each taxonis assessed based on numerous sources such asRichardson et al. (2000), Occhipinti-Ambrogi &Galil (2004), Richardson & Pyšek (2006), Pyšek etal. (2004) or Richardson (2011). Further commentson distribution, habitats occupied, etc. and fieldphotographs of most of them are also presented.The second and third authors assisted with theidentification of most of the species.

The paper is divided in two parts in both ofwhich the taxa are arranged in alphabetical se-quence. Nomenclature of the taxa presented ismostly in accordance with recent sources (cf. APD,2019; Korotkova & Raab-Straube, 2017; onlinedatabase http://www.llifle.com/Encyclopedia/CACTI/Family/Cactaceae/).

Results 1. OpuntIOIDeAe SuBFAMIly1.1 Austrocylindropuntia subulata (Muehlenpf.)Backeb., Cactaceae Jahrb. Kakteen. PereskioideaeOpuntioideae 1941(2): 12. 1942. (Figure 1) ≡ Opuntia subulata (Muehlenpf.) Engelm., Gard.Chron. 19: 627. 1883.≡ Pereskia subulata Muehlenpf., Allgem. Garten-zeitung 13: 347. 1845.≡ Cylindropuntia subulata (Muehlenpf.) F.M.Knuth in Backeb. & F.M. Knuth, Kaktus-ABC: 121.1936.

New for the non-native flora of Tunisia and sec-ond record for the flora of North Africa (APD, 2019;Korotkova & Raab-Straube, 2017).

TUNISIA: Jendouba: Route Bulla Regia, one singlepopulation on the roadside, 36°31′20″N,08°45′05″W, 169m a.s.l., 24.03.2018, R. El Moknis.c.; idem, Jleyleya, one single extended populationon the roadside, 36°32′57″N, 08°43′06″W, 206ma.s.l., 13.10.2004–01.08.2017, R. El Mokni s.c.;Route Boussalem, after the Bridge “Five Dinars”,36°24′ 02″N, 08°53′58″W, 157m a.s.l., 01.08.2017,R. El Mokni s.c.; Tabarka, one single population onthe roadside towards touristic route, 36°57′00″N,08°46′20″W, 29m a.s.l., 24.10.2014-24.01.2018, R.El Mokni s.c.; Beja: Ouechtata, one single popula-tion on the roadside, 36°57′35″N, 08°59′11″W, 69ma.s.l., 26.03.2018, R. El Mokni s.c.; towards Mjaz ElBab, one single population on the roadside,36°39′40″N, 09°30′54″W, 154m a.s.l., 01.08.2017, R.El Mokni s.c.; Bizerta: one little population withinlandscaping, 37°17′26″N, 09°52′17″W, 07m a.s.l., R.El Mokni s.c.; Route Sejnan Nefza, one single pop-ulation on the roadside, 37°04′36″N, 09°10′27″W,142m a.s.l., 26.03.2018, R. El Mokni s.c.; Mateur,route towards Sidi Nsir, one huge population on theroadside, 37°02′03″N, 09°39′18″W, 43m a.s.l.,13.09.2004-26.03.2018, R. El Mokni s.c.; Tunis:

Bab-Bnat, one single population on the roadside,36°47′51″N, 10°09′51″W, 67m a.s.l., 05.03.2018, R.El Mokni s.c.; Sousse: towards motorway, one sin-gle extended population on the roadside,35°50′01″N, 10°33′52″W, 45m a.s.l., 02.01.2017, R.El Mokni s.c.; Monastir: Menzel Harb towardsMonastir’s airport, one huge population growing onthe roadside, 35°45′49″N, 10°43′12″W, 13m a.s.l.,11.03.2018, R. El Mokni s.c.

Probably native only to the Andes of Peru, Aus-trocylindropuntia subulata is now one of the mostwidely distributed opuntioids of South America,particularly in Argentina and Bolivia where it iswidely cultivated as natural fencing. It has widelynaturalised in parts of southern Europe and in thetropics where it was initially introduced as an or-namental. 

Description:  Stem (trunk) 6 to 10cm in diame-ter, the old bark smooth and brown, its areolesbearing clusters of 8 spines or more, branches nu-merous, elongate, unsegmented, 30–50cm long, 4–7cm in diameter, more or less clustered but notwhorled, at first almost at right angles to mainstem but soon erect, somewhat fragile, brightgreen. Areoles, shortly woolly, sitting in the retusegrooves on upper parts of the tubercles.Leaves,  persistent for more than a year, green,more succulent than on other species, awl-like,nearly at right angles to branch, straight or some-what bowed above, nearly terete, pointed, 5 to 12cmlong, grooved on the underside. Spines, usually 1 or2 or sometimes spineless (more numerous in olderstems), slender, erect, strong, straight, pale yellowto greyish white, to 8cm long. Flowers borne towardthe ends of the branches, red, not opening widely to6cm long. Pericarpels long, tuberculate, awl-like,with erect scales to 2cm long. Sepals reddish,minute, 4 to 8mm long or less. Petals broader thanthe sepals, orange-red or greenish yellow. Stylerose-red except for the whitish base, including thestigma-lobes about 3cm long, about as long as thelongest stamens. Stigma-lobes 5 or 6, slender, or-ange-yellow. Fruits, ovoid to oblong to club shaped,more or less persistent, sometimes spiny, 6 to 10cmlong, leafy, with a deep umbilicus, sometimes pro-liferous. Seeds  few, large, 10 to 12mm long (re-vealed from an online databasehttp://www.llifle.com/Encyclopedia/CACTI/Fam-ily/Cactaceae/4092/Austrocylindropuntia_subu-lata). Two subspecies are sometimes recognised,subsp. subulata and subsp. exaltata (A. Berger)D.R. Hunt.

Austrocylindropuntia subulata is mentionedhere for the first time from Tunisia, and for the sec-ond time as naturalised in North Africa (APD 2019;Korotkova & Raab-Straube, 2017). In Europe, it isrecorded as a naturalised alien in Italy, Sardiniaand Sicily, as alien (status unknown) in Portugal,and as a casual alien in Corsica (Korotkova &

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Raab-Straube, 2017). In the Canary Islands it iscommonly naturalised on all islands except El Hi-erro (Acebes Ginovés et al., 2010).

1.2 Cylindropuntia imbricata (Haw.) F.M.Knuth in Backeberg & Knuth, Kaktus-ABC: 125.1936. (Figure 2)≡ Cactus imbricatus Lem., Les Cactées: 88. 1868.≡ Cereus imbricatus Haw., Suppl. Pl. Succ.: 70.1819.≡ Grusonia imbricata (Haw.) G.D. Rowley, Tephro-cactus Study Group 12(3): 44. 2006. ≡ Opuntia imbricata (Haw.) DC., Prodr. 3: 471.1828.

Confirmed for the non-native flora of Tunisia asnaturalised (APD 2019; Korotkova & Raab-Straube2017).

TUNISIA: Sidi-Bouzid/Gafsa: some populationswithin the National park of Bouhedma, 34°28′18″N,09°28′36″W, 237m a.s.l., 24.04.2017, R. El Moknis.c.

Cylindropuntia imbricata is a cactus found inthe southwestern United States and northern Mex-ico (Durango, Zacatecas, and San Luis Potosí), in-cluding some cooler regions in comparison to manyother cacti. It occurs primarily in the arid regions ofthe Southwestern United States in the states ofTexas, New Mexico, Arizona and Nevada.

Description: Tree-like, often 3m high or higher,with a more or less definite woody trunk 2.5cm indiameter; ultimate joints 2 to 3cm in diameter,strongly tuberculate; leaves 8 to 24mm long, terete;tubercles 2 to 2.5cm long, flattened laterally; spines8 to 30, 2 to 3cm long, brown, covered with paperysheaths; flowers bowl-shaped, diurnal, purple or

magenta, rarely rose-pink, borne at ends ofbranches, 4 to 6cm long, sometimes 8 to 9cm broad,purple; ovary tuberculate, bearing a few bristlesfrom some of the upper areoles, spineless, occasion-ally bearing a few bristles from some of the upperareoles; perianth purple; anthers yellow on purplefilaments; stigma pale yellow; fruit naked, egg-shaped with a hollow at the wide end where theflower fell off, yellow, 2.5 to 4.4cm long, 2–4cm in di-ameter, strongly tuberculate or, when long persist-ent, smooth; seeds 2.5 to 3.5(–4)mm in diameter. Inaddition to the typical variety (var. imbricata), twovarieties are recognised from West Texas: var. ar-gentea (M.S. Anthony) Backeb. with silvery spines,and var. arborescens (Engelm.) A.D. Zimmermanwhich is taller (Britton & Rose, 1919).

The species is naturalised in arid regions andalong watercourses in South America, South Africa,the Mediterranean basin and Australia. In NorthAfrica, C. imbricata is reported as an introducedcultivated alien (status unknown) for Morocco,Tunisia and Libya (APD, 2019; Korotkova & Raab-Straube, 2017). In the Canary Islands it is onlyknown from a single locality in Gran Canaria (Ver-loove et al., 2017a)

1.3 Opuntia ficus-indica (L.) Mill. f. amyclaea(Ten.) Schelle, Handb. Kakteenkult: 51. 1907.Nomenclature according to Kiesling (1998). (Figure3)≡ Opuntia amyclaea Tenore, Fl. Neap. Prodr. App.5: 15. 1826.= O. megacantha Salm-Dyck, Hort. Dyck.: 363.1834. = O. maxima Salm-Dyck, Cact. Hort. Dyck.: 68.1850, non Mill.

Figure 2. Cylindropuntia imbricata, Bouhedma (Sidi-Bouzid), abandoned land, 24th April 2017.

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TUNISIA: Monastir, scattered individuals growingon a sun-exposed clayey slope close to the coast,35°46′54″N, 10°48′12″W, 40m a.s.l., 04.12.2017, R.El Mokni s.c.

Description: Compared with Opuntia ficus-in-dica it has oblong to suborbicular, glaucous-pru-inose, distinctly flattened cladodes that are alwaysspiny. Molecular data seem to suggest that it prob-ably is nothing else than the wild progenitor of do-mesticated O. ficus-indica (Labra et al., 2003).Other authors, however, accept it as a species dis-tinct from O. ficus-indica, based on morphometricanalyses (Reyes-Agüero et al., 2005). Forma amy-claea has an erect habit; joints oblong to elliptic, 30to 40cm long, about twice as long as broad, thick,dull green, a little glaucous; leaves few mm long,acute, red; areoles small, with 1 or 2 short bristlesfrom the lower parts of areoles; spines 1 to 4, nearlyporrect, usually less than 3cm. long, white or horn-coloured, the stoutest angled; glochids brown, soon

disappearing; flowers yellow; fruit yellowish red,not very juicy (cf. Britton & Rose, 1919).

This taxon is new to the flora of Tunisia and asecond record for the flora of North Africa (APD,2019). In Tunisia, O. ficus-indica f. amyclaea canobviously be classified as well naturalised as it ap-pears in several subpopulations. However, likeother species from this genus, it certainly has thepotential to spread and become an invasive alien.

1.4 Opuntia dejecta Salm-Dyck, Hort. Dyck.: 361.1834. (Figure 4)≡ Nopalea dejecta (Salm-Dyck) Salm-Dyck, Cact.Hort. Dyck. 1849: 64. 1850.

TUNISIA: Monastir, 35°46′03″N, 10°50′07″W, 38ma.s.l., abandoned land, 18.11.2017, R. El Mokni s.c.

Probably a native of Panama, this species isvery common in cultivation in tropical America(Britton & Rose, 1919).

Figure 3. Opuntia ficus-indica f. amyclaea, Monastir city within the coastal side, clayey slope, 7th April2018.

Figure 4. Opuntia dejecta, Monastir city, abandoned land, 17th November 2017.

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Figure 5.Opuntia dillenii complex, A. Monastir city within coastal side, rocky slope, 29th November 2017;B. Bouhedma (Sidi-Bouzid), abandoned land, 24th April 2017; C. Tamera, Sejnane (Bizerte) within road-sides, 12th April 2018, D. Monastir city, rocky slope within abandoned herbaceous stratum, 7th April 2018.

A B

C

D

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Description: Opuntia dejecta is a plant of 1 to2m in height, with a definite trunk, very spiny, theold areoles often bearing 6 or 8 spines; joints nar-row, 10 to 15cm long, only moderately thick, oftendrooping, bright green even in age, bearing usuallytwo somewhat spreading spines at an areole; spinesat first pale yellow or pinkish, in age grey, thelongest 4cm long; flower, including ovary and style,is 6–7cm long; sepals obtuse; petals erect, dark red;stamens long-exerted, dark red. (Britton & Rose,1919).

Opuntia dejecta is new to the flora of Tunisia,the flora of Africa (APD, 2019) and is recorded forthe second time for the Mediterranean area, whereit was so far reported only as naturalised in Sicily(Korotkova & Raab-Straube, 2017). In Tunisia, O.dejecta is obviously a recent escape and can be clas-sified only as casual. However, like other specieswithin this genus, it certainly has the potential tospread to the nearby coastal regions.

1.5 Opuntia dillenii (Ker Gawl.) Haw., Suppl. Pl.Succ.: 79. 1819. (Figure 5)≡ Cactus dillenii Ker Gawl., Bot. Reg. 3: t. 255.1818.≡ Opuntia stricta (Haw.) Haw. var. dillenii (KerGawl.) L.D. Benson, Cact. Succ. J. (Los Angeles) 41:126. 1969.

TUNISIA: Monastir, scattered populations growingon a sun-exposed clayey slope of the rocky coast inthe north east of Monastir, 35°46′54″N, 10°48′12″W,40m a.s.l., 04.12.2017, R. El Mokni s.c.; idem, scat-tered individuals growing on the sandy coast at thenorth east of Monastir, 35°44′57″N, 10°49′36″W,27m a.s.l., 12.11.2017, R. El Mokni s.c.; idem, scat-tered massive populations/subpopulations growingon both sides of the Monastir Sousse tourist route,35°45′49″N, 10°43′12″W, 13m a.s.l., 11.03.2018, R.El Mokni s.c.; idem, scattered subpopulations grow-ing on both sides of the Menzel Kamel-Ouerdenineroute, 35°38′26″N, 10°39′37″W, 75m a.s.l.,11.03.2018, R. El Mokni s.c.; Bizerta, widely culti-vated as ornamental in pots and public gardens,escaping to roadsides or coastal rocks, 37°19′52″N,09°51′52″W, 12m a.s.l., R. El Mokni s.c.; Mateur-massif of Ichckeul, several scattered massive popu-lations growing everywhere within the massif ofIchckeul, including the National Park, 37°07′27″N,09°39′36″W, 33–476m a.s.l., 24.03.2014, R. El Moknis.c.; Sejnane–Tamera, scattered subpopulationsgrowing on roadsides, 37°04′41″N, 09°08′57″W,122m a.s.l., 12.04.2018, R. El Mokni s.c.

Plants of O. dillenii sprawl or grow erect andreach a height of about 50 to 200cm. They consist offlattened, elliptic and obovate stem segments, andthese are impressive for many stout spines. Thejoints are dull green or bluish-green in colour, 10 to40cm long, 5 to 15cm broad, 1 to 2cm thick, and ir-regularly trimmed with relatively few areoles.

These are some millimeters across and bear smallscale-like leaves on young joints, later on 1 to 10spines each. The spines are very variable in colour(yellow to brown) and length (1.5 to 6cm) and cansometimes be lacking. They spread in all directionsand are accompanied by glochids, short barbed bris-tles which form striking tufts (Backeberg, 1970;Benson, 1982; Ellenberg, 1989; Anderson, 2005).

At flowering, O. dillenii plants have 6 to 8cmwide flowers that are finally lemon yellow in colourbut can be slightly reddish in early stages of devel-opment. The purple-skinned fruits are pear-shaped(Figure 5D) measure 4 to 7cm in length and about3cm in diameter. They normally show no spinesand their glochids mostly fall off from the few are-oles at maturity. The juicy flesh of fruits is purplein colour and contains many rounded seeds whichare –with an arillus-like envelop - about 4mm in di-ameter and tan coloured (Backeberg, 1970; Benson,1982; Ellenberg, 1989; Anderson, 2005; Land Pro-tection, 2006).

There is considerable dispute whether Opuntiadillenii belongs in Opuntia stricta or should be con-sidered distinct (Böhm, 2008). Opuntia dillenii dif-fers from O. stricta mainly by its unarmed stemsegments with groups of 1–7 large spines on most ofthe areoles. Both are low-growing plants (usually50–100cm tall) with relatively large flattened andelongated (i.e. elliptic or obovate) stem segments.

Opuntia dillenii originates from southeasternparts of North America, the east coast of Mexico,the Bermuda’s, the West Indies and from the northof South America (Britton & Rose, 1919). Wild pop-ulations are nowadays reported in the same areas(Backeberg, 1970; Benson, 1982; Schultze-Motel,1986; Scheinvar, 2002; Anderson, 2005) and addi-tionally at the Gulf Coast of Texas (Benson, 1982),the southeastern coastal margin of Brazil (Schein-var, 1984), and in Ecuador (Anderson, 2005). Thenative locations of O. dillenii are mostly at an alti-tude near sea level in sand dunes, at the edges ofmaritime forests, or associated with tropical plants(Benson, 1982; Pinkava, 2003). O. dillenii was in-troduced into many parts of the world. It is foundaround the Mediterranean (Schultze-Motel, 1986;Gentile, 1991), on the Canary Islands (Perfumi &Tacconi, 1996; Loro et al., 1999; Anderson, 2005;Díaz Medina et al., 2007), in the west and east ofAfrica (Schultze-Motel, 1986), on Madagascar andMauritius (Schultze-Motel, 1986; Ellenberg, 1989),in North Yemen (Ellenberg, 1989), in India (Backe-berg, 1970; Badami & Thakkar, 1984;, Gupta et al.,2002), in the southeast of Asia (Schultze-Motel,1986; Jiang et al., 2006), and in Australia (Backe-berg, 1970; Schultze-Motel, 1986; Ellenberg ,1989).Opuntia dillenii becomes naturalised under condi-tions different from those in its original areas andcan even rapidly spread out (Ellenberg, 1989). It isinvolved in dangerous infestations with several Op-untia species (Burdon & Marshall, 1981), notably

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O. stricta (pest pear) in eastern Australia (Free-man, 1992; GISD, 2010).

In Tunisia, Opuntia dillenii can be consideredan invasive species in many regions of the country.

1.6 Opuntia lindheimeri Engelm., Bost. J. Nat.Hist. 6: 207. 1850. (Figure 6)≡ Opuntia engelmannii subsp. lindheimeri (En-gelm.) U. Guzmán & Mandujano, Cactaceae Syst.Init. 16: 18. 2003.

TUNISIA: Monastir, 35°46′00″N, 10°49′37″W, 40ma.s.l., roadside, a single individual, 15.02.2018, R.El Mokni s.c.

A species native to Southwestern Louisiana,southeastern Texas (U.S.A.), and Tamaulipas(Mexico).

Description: Opuntia lindheimeri has an erecthabit, 2 to 4m high, with a more or less definitetrunk, but at times much lower and spreading;joints green or bluish green, somewhat glaucous,orbicular to obovate, up to 2.5cm long; leaves subu-late, 3 to 4mm long, somewhat flattened, pointed;areoles distant, often 6cm apart; spines usually 1to 6, often only 2, one porrect and 4cm long or more,the others somewhat shorter and only slightlyspreading, pale yellow to nearly white, sometimesbrownish or blackish at base, or some plants spine-less; glochids yellow or sometimes brownish, usu-ally prominent; petals usually yellow (sometimesred); stigma-lobes usually green; fruit purple, pyri-form to oblong, 3.5 to 5.5cm long. It is an extremelyvariable species, composed of many races, differingmainly in armament, colour of flowers, size andshape of joints and of fruit (Britton & Rose, 1919).

A very close taxon compared to Opuntia engel-mannii and still considered as a synonym by manyauthors, O. lindheimeri is mainly characterized by(i) the presence of flowers with coloured orangetepals versus yellow tepals in O. engelmannii, and

(ii) thorns completely remaining yellowish whereasthorns are brown at the base and the rest yellow inthe case of O. engelmannii (see e.g. Vázquez & Gar-cía, 2017).

New to the flora of Tunisia and the flora ofAfrica, O. lindheimeri has been reported before inEurope from Azerbaijan and Georgia in Transcau-casia (Korotkova & Raab-Straube, 2017) as well asfrom Italy and Spain (Sanz-Elorza et al., 2006;Guiggi, 2014). It is also known from a single popu-lation in Gran Canaria in the Canary Islands (Ver-loove et al., 2017a). In Tunisia, O. lindheimeri isobviously a recent escape and can be classified onlyas casual. However, like other species within thisgenus, it certainly has the potential to spread to thenearby coastal regions.

1.7 Opuntia microdasys (Lehm.) Pfeiff., Enum.Diagn. Cact.: 154. 1837. (Figure 7).≡ Cactus microdasys Lehm., Index Seminum (Ham-burg), 1827: 16. 1827.

TUNISIA: Bizerta: abandoned plantations in manysites, 37°17′26″N, 09°52′17″W, 7m a.s.l., R. ElMokni s.c.; Monastir: scattered little subpopula-tions growing on both sides of the Monastir Soussetourist route, 35°45′49″N, 10°43′12″W, 13m a.s.l.,11.03.2018, R. El Mokni s.c.; Sidi-Bouzid/Gafsa:some populations in landscaping within Nationalpark of Bouhedma, 34°28′18″N, 09°28′36″W, 237ma.s.l., 24.04.2017, R. El Mokni s.c.

An endemic of Mexico, it became an invader inAustralia and South Africa (Walters et al., 2011;Novoa et al., 2014).

Description: Opuntia microdasys (bunny ears)is a branched, shrubby succulent perennial, erectto spreading, up to 1m tall. Stem segments palegreen, brighter green when young, round to oval-shaped, 8–16 × 7–16 cm. Areoles prominent, round,close-set. Needle-like spines usually absent, some-

Figure 6. Opuntia lindheimeri, Monastir city, roadsides under plantation of Ficus benjamina, 15th Feb-ruary 2018.

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times a single short one present on an areole.Glochids dense, fine, bristle-like, yellow, white orbrown. Flowers 40mm in diameter, petals to 40mmlong, yellow with a slight reddish tint towards theslightly frilly tips of the petals. Fruits 30–40mm indiameter, fleshy, egg-shaped to globose, red to pur-plish-red, densely covered with areoles bearingdense clusters of glochids (Smith et al., 2011; Bárce-nas et al., 2017).

New to the flora of Tunisia and a second recordfor the flora of North Africa (APD, 2019), Opuntiamicrodasys is widely cultivated as a popular and at-tractive ornamental cactus. It is often recorded asan escape from cultivation, either naturalised oradventitious (casual), in Europe at least in France,Italy, Portugal, and Spain (Essl & Kobler, 2008;Novoa et al., 2014; Korotkova & Raab-Straube2017; Verloove et al., 2017a& b).

1.8 Opuntia monacantha (Willd.) Haw., Suppl.pl. succ.: 81. 1819. (Figure 8).≡ Cactus monacanthos Willd., Enum. pl. 33. 1813(1814).= Opuntia vulgaris Britton & Rose, non Mill., Cact.1: 156. 1919.New for the flora of Tunisia and the flora of NorthAfrica.

TUNISIA: Bizerta: roadside, a single individual,37°16′22″N, 09°52′25″W, 31m a.s.l., 15.02.2018, R.El Mokni s.c.; Monastir: roadside, a single individ-ual, 35°46′00″N, 10°49′37″W, 40m a.s.l., 03.12.2017,R. El Mokni s.c.

This South American species (Argentina andBrazil) is widely cultivated as an ornamental.

Description: Plant 2 to 4 or even 6m high, oftenwith a definite trunk, usually with a large muchbranched top; trunk cylindric, 15mm in diameter,either spiny or smooth; joints ovate to oblong, nar-rowed at base, 10 to 30mm long, bright shininggreen; leaves subulate, 2 to 3mm long; areoles filledwith short wool; glochids brownish; spines 1 or 2,sometimes more (on the trunk often 10 or more)from an areole, erect, 1 to 4cm long, yellowishbrown to dark reddish brown; flowers yellow to red-dish, 7.5cm broad; sepals broad, each with a broadred stripe down the middle; petals golden yellow,widely spreading; filaments greenish; style white;stigma-lobes 6, white; ovary spineless, 3.5cm long;fruit obovoid, 5 to 7.5cm long, reddish purple, long-persisting, sometimes proliferous (Britton & Rose,1919).

New to the flora of Tunisia and for the flora ofNorth Africa (APD, 2019), Opuntia monacantha is

Figure 7. Opuntia microdasys, Bouhedma (Sidi-Bouzid), abandoned land, 24th April 2017.

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cultivated as an attractive ornamental cactus. InEurope, it is recorded as naturalised at least inFrance, Corsica, Canary Islands, as alien in Sicilyand doubtfully present in Switzerland, Italy, Por-tugal, and Spain (Korotkova & Raab-Straube,2017).

1.9 Opuntia robusta J.C. Wendl. Cat. Hort. Her-renh. 1835 (Figure 9).= Opuntia camuessa F.A.C. Weber, Dict. Hort.[Bois] 2: 895. 1898.New for the flora of Tunisia and a first record forthe flora of North Africa (Hansen & Sunding, 1993;APD, 2019).

TUNISIA: Monastir: two subpopulations of few in-dividuals, growing on both sides of the MonastirSousse tourist route, 35°45′49″N, 10°43′12″W, 13m

a.s.l., 11.03.2018, R. El Mokni s.c.

Opuntia robusta is widely distributed in north-ern and central Mexico (Coahuila, Durango, NuevoLeón, San Luis Potosí, Tamaulipas, Aguascalientes,Guanajuato, Guerrero, Hidalgo, México, Querétaroprovinces). In the United States it is found in Ari-zona. This species has expanded into North andSouth America and has been introduced into NewZealand and the Mediterranean region of southernEurope, South Africa, Madagascar and Australiawhere it is a significant environmental and pastoralweed, particularly in the states of South Australia,New South Wales, Queensland and Victoria. 

Description: Opuntia robusta, popularly knownas the ‘wheel cactus’ in reference of the circularshape of the segments, is a much branched, oftenerect, succulent shrub or small tree, sometimes 4–

Figure 8. Opuntia monacantha, Monastir city, roadsides under plantation of Ficus benjamina, 29th No-vember 2017.

Figure 9. Opuntia robusta, Monastir city, coastal area, 30th May 2019.

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6m high, but commonly 1–2m, with more or less dis-tinct trunks. The branch segments are armed withtufts of numerous barbed bristles (glochids) andcommonly spineless or with 1 to 5 sharp spines to4cm long arising from each areole. Flowering inspring (April to May), both plants with male, femaleand hermaphrodite flowers can be found in Opuntiarobusta. The fruits (in autumn, September to Octo-ber), which may be red, orange, yellow or green, areglobular to ellipsoid, at first more or less tubercu-late, 7 to 9cm long, deep red, but also orange, yel-low or green, with light brown felt and yellowglochids c. 3mm long, produced in profusion, possessthick peel (with sharp glochids) and juicy red pulpcontaining numerous seeds; they are edible but bestfor animal feeding (revealed from an online data-base http://www.llifle.com/Encyclopedia/CACTI/Family/Cactaceae/32724/Opuntia_robusta).

New to the flora of Tunisia and a first report forthe flora of North Africa (APD, 2019), Opuntia ro-busta has been reported from Gran Canaria, LaPalma, and Tenerife in the Canary Islands (AcebesGinovés et al., 2010) although many claims requireconfirmation, resulting from confusion with themorphologically similar O. ficus-indica f. amyclaea(Verloove et al., 2017a & b). In Europe, it is onlyrarely reported in the wild (Guiggi, 2014). It isrecorded as naturalised in France, Corsica andSicily and as a casual alien in Italy (Korotkova &Raab-Straube, 2017). Moreover, it is reported as aninvasive weed in several countries, including Aus-tralia, Botswana, and South Africa (Walters et al.,2011; Novoa et al., 2014).

,CACtOIDeAe SuBFAMIly2.1 Cereus hildmannianus K. Schum., Fl. Bras.4(2): 202. 1890. (Figure 10).= Cereus peruvianus auct. non (L.) Mill.New for the non-native flora of Tunisia and for theflora of North Africa (APD, 2019; Korotkova &Raab-Straube, 2017).

TUNISIA: Monastir: some scatteredindividuals/subpopulations in center of the townwithin some public gardens, 35°46′13″N,10°49′39″W, 41m a.s.l., 03.10.2015-30.03.2018, R.El Mokni s.c.; Jemmel, 35°37′43″N, 10°45′32″W,41m a.s.l., 03.10.2015-30.03.2018, R. El Mokni s.c.;idem 35°37′28″N, 10°45′32″W, 57m a.s.l.,03.10.2015–04.04.2018, R. El Mokni s.c.; Sousse:Bouhsina, one huge population on an ancient ex-tended building, 35°49′15″N, 10°37′33″E, 71m a.s.l.,27.12.2017, R. El Mokni s.c.; Tunis: Mornaguia, onevigourous population on the right of the motorway,36°45′21″N, 09°59′19″W, 66m a.s.l., 24.03.2018, R.El Mokni s.c.; Bizerta: one small population withinon the right of the road towards Jarzouna fromMenzl Jemil, 37°15′29″N, 10°37′33″W, 28m a.s.l.,27.12.2017, R. El Mokni s.c.

Native to Argentina, Brazil (Mato Grosso doSul, Minas Gerais, Paraná, Rio de Janeiro, RioGrande do Sul, Santa Catarina, São Paulo), Para-guay and Uruguay (Anderson, 2001).

Description:  stems, cylindrical, segmented, bluegreen to dull green, often glaucous when young, to15cm in diameter; ribs, (4–)5–6(–8), thin to obtuse,slightly indented, 2.5–3.5cm high, rounded, greenoften with large yellow patches along the sides; are-oles, distant, brown, large 2cm apart; spines, usu-ally absent, occasionally a few golden or brownspines develop afterwards; flowers, very large, noc-turnal, elongated, funnelform, 20 to 30cm long;inner perianth-segments white, broad and obtuseovary naked, 2.5 to 3cm long, blooming at night inspring through summer. The buds grow quickly,shooting out from the branches like so many smallsnakes. Within two weeks, dozens of flowers open,always at night and all or nearly all flowers opensimultaneously at the same time. By mid-morningthe following day, all had closed and drooped. Thisis apparently a water-saving strategy by cacti. Theplant usually buds and flowers about two weeks fol-lowing any warm-season rain. The fruits usuallyripen within a month. Buds may erupt well intofall, even early winter, if sufficiently warm andrains fall. During dry times, the buds often fall offprior to opening; fruits globose, pear-shaped, redwith white pulp. The Peruvian apple cactus mayproduce fruit 3–4 years after propagation from seed(revealed from an online database http://www.llifle.com/Encyclopedia/CACTI/Family/Cac-taceae/6499/Stenocereus_peruvianus).

This species is widely cultivated worldwide asan ornamental plant. It has escaped and sometimesbecame invasive. In Europe, Cereus hildmannianushas been reported before from Italy (Guiggi, 2010);it was also recently reported from Canary Islandsas casual alien (Verloove et al., 2017b; Korotkova &Raab-Straube, 2017).

2.2 Echinopsis eyriesii (Turpin) Zuccarini inPfeiffer and Otto, Abbild. Beschr. Cact. 1: under pl.4. 1839. (Figure 11).≡ Echinocactus eyriesii Turpin, Ann. Inst. Roy.Hort. Fromont 2: 158. 1830. ≡ Cereus eyriesii Pfeiffer, Enum. Cact.: 72. 1837. ≡ Echinonyctanthus eyriesii Lemaire, Cact. Gen.Nov. Sp.: 84. 1839. ≡ Echinopsis pudantii Pfersdorff, Monatsschr. Kak-teenk. 10: 167. 1900.New for the non-native flora of Tunisia, for the floraof North Africa and for the flora of the Mediter-ranean area (APD, 2019; Korotkova & Raab-Straube, 2017)TUNISIA: Monastir: some scattered individuals inan abandoned field with a Chenopodiaceae/Ama-ranthaceae vegetation alongside the road towardsSousse, 35°45′34″N, 10°44′24″W, 29m a.s.l.,03.10.2017, R. El Mokni s.c.; Tunis: Ariana, some

46 Bradleya 38/2020

scattered individuals, 36°50′40″N, 10°11′54″W, 25ma.s.l., 10.12.2017, R. El Mokni s.c.

Originating from Argentina (Buenos Aires). Ithas a large distribution within Southern Brazil,Uruguay, and the province of Entre Rios, Ar-gentina. (cf. Britton & Rose, 1922).

Description: Echinopsis eyriesii is a simple orclustered, globular to short-columnar cactus; ribs11 to 18, not tuberculate, rather thin above; areolescircular, filled with white or tawny wool; spines sev-eral, 14 to 18, very short, 5mm long or less; flowerfrom the side of plant but above the middle, large,17 to 25cm long; inner perianth-segments white,acuminate; stamens and style shorter than the pe-rianth-segments; scales on the flower-tube small,

ovate, brownish, hairy in their axils (cf. Britton &Rose, 1922).

2.3 Hylocereus undatus (Haw.) Britton & Rose,Fl. Bermuda: 256 (fig. 278). 1918. (Figure 12).≡ Cereus undatusHaw., Philos. Mag. Ann. Chem. 7:110. 1830.New for the non-native flora of Tunisia and for theflora of North Africa. (APD, 2019; Korotkova &Raab-Straube, 2017).TUNISIA: Monastir: one epiphytic population onplanted Phoenix canariensis within an old buildingin the centre of the town, 35°46′03″N, 10°49′55″W,40m a.s.l., 27.02.2018, R. El Mokni s.c.; Tunis–Ari-ana: escaped plants from ancient planted individu-

Figure 10. Cereus hildmannianus, A. B. & C. Jemmal (Monastir), old buildings and roadsides, 13th April2018; D. & e. Ksar-Hellal (Monastir), roadsides, 12th April 2018.

A

B

C

D e

47Bradleya 38/2020

als in pots within an old building, 35°50′50″N,10°11′36″W, 34m a.s.l., 15.11.2017, R. El Mokni s.c.

This species is widely cultivated and has be-come naturalised in many countries, however, asthe native range is unknown, nothing is knownabout the size and trends of the native wild popu-lation (Durán et al., 2017).

Description: Hylocereus undatus is a sprawlingor vining, terrestrial or epiphytic cactus. It climbsby use of aerial roots and can reach a height of 10mor more growing on rocks and trees. It is very vari-able and closely related to  Selenicereus.Stems, green and 3-winged, from a few cm up to 5mlong (in mature plants), 4 to 7.5cm wide with mar-gins undulate and horny with wings that are 2.5 to

5cm wide; areoles about 4.5cm apart; spines 1 to 3(–5), conical and up to 1cm long (but usually about 2–3mm long); flowers ornate, fragrant and beautiful,25–35cm long by 30cm across, white with greenouter tepals and bracts, blooming only at night, andusually the flower lasts just one night. In tropicalclimates the plants can have up to 4–6flowering/fruiting cycles per year. In temperate cli-mates they bloom in late spring to early summer;once pollinated it produces edible red fruits withwhite pulp up to 12cm long (revealed from an on-line database http://www.llifle.com/Encyclope-dia/CACTI/Family/Cactaceae/7429/Hylocereus_undatus).

The native range of this species is unknown asit has long been in cultivation (Anderson, 2001;

Figure 11. Echinopsis eyriesii, A. Ariana city, abandoned land, 16th November 2017; B. Monastir city,coastal area, clayey slope, 24th April 2018.

A

B

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Hunt et al., 2006). Some believe it to be native tosouthern Mexico, the Pacific side of Guatemala,Costa Rica, and El Salvador, however, this has notbeen confirmed. It is the most widely cultivatedspecies in the genus (including commercial cultiva-tion) and often escapes and becomes naturalised.This species is therefore widely distributed and hasbeen recorded from tropical forests in Mexico, var-ious islands in the Caribbean, Central America,tropical South America, and southeasternAsia. The type collection is from a cultivated speci-men grown in London which was probably broughtfrom China. In Brazil, it is introduced as a gardenplant and sometimes escapes into roadside areasand maritime scrub, occurring in Pernambuco,Alagoas, Bahia, Minas Gerais and Espírito Santo(Taylor & Zappi, 2004). In addition to its wide nat-uralised range, it has also a wide altitudinal rangeoccurring from near sea level up to 2750m. (Duránet al., 2017). In Europe, the species is reported onlyfrom the Canary Islands by Korotkova & Raab-Straube (2017). It is here reported for the first timefrom North Africa.

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