Bradleya 38/2020 pages 35–50

New records of cacti (Opuntioideae & Cactoideae, Cactaceae) from Tunisia Ridha El Mokni 1,2,3 , Filip Verloove 4, Alessandro Guiggi 5 & Mohamed Hédi El Aouni 1 1. University of Carthage, Laboratory of Botany and 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 of Tabarka, 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: [email protected]) 5. DISTAV, Polo Botanico, Università degli Studi di Genova, International Cactaceae Research Cen - ter (ICRC), Corso Dogali, 1M - 16136 Genova, Italy. (email: [email protected]) Photographs: Ridha El Mokni.

Summary : Botanical surveys undertaken in Cen - Beschreibungen gegeben, Kommentare zu ihrem tral and Northern Tunisia (North Africa), mostly aktueller Status werden angegeben und Fotos wer - between 2012 and 2018, have revealed first records den bereitgestellt. and/or new distributional records for non-native succulents (sensu lato) belonging to the Cactaceae Introduction family. Some can be considered naturalised or even Succulents are found across the globe in nearly potentially invasive [e.g. Opuntia dillenii (Ker all types of habitat, but most often in arid or semi- Gawl.) Haw.], whereas others are here reported for arid parts of the world. They are specially adapted the first time for North Africa, either as recently to deal with dry, desert-like conditions, and able to naturalised or casual, e.g.: Echinopsis eyriesii store water in one or more of their organs; their (Turpin) Zuccarini, Opuntia dejecta Salm-Dyck, O. leaves, stems, or roots are often filled with water- lindheimeri Engelm., O. monacantha Haw and O. storing tissue and are thus unusually fleshy and robusta J.C. Wendl. ex Pfeiff. Short descriptions are enlarged. The Cactaceae family is distinguished given for each of these species, comments on their from the other families (i.e. actual status were pointed out and field photo - Agavaceae, Aloaceae, Euphorbiaceae, etc.) by the graphs are provided. presence of small rounded woolly areas normally localized along the ribs of the plant (known as are - Zusammenfassung : Botanische Untersuchungen in oles) from which new segments, flowers and spines Zentral- und Nordtunesien (Nordafrika), haupt - are developing (retrieved from https://www.iucn.org sächlich zwischen 2012 und 2018, haben erste /ssc-groups/-fungi/-and-succulent- Aufzeichnungen und/oder neue Beobachtungen für plants-specialist-group/what-are-cacti-and-succu - nicht-einheimische Sukkulenten (sensu lato) aus lents). Cactus roots are generally less specialized in der Cactaceae-Familie ergeben. Einige können als morphology than the above-ground vegetative eingebürgert oder sogar potentiell invasiv angese - parts. They have a shallow distribution, which en - hen werden [als Opuntia dillenii (Ker Gawl.) ables them to exploit limited rainfall, and are rich Haw.], während andere hier zum ersten Mal für in root hairs. Associations with fungi and bacteria Nordafrika berichtet werden, entweder vor kurzem can improve the capture of mineral nutrients eingebürgert oder vorübergegend: Echinopsis ey - (Dubrovsky & North, 2002; Land Protection, 2006). riesii (Link) Zucc. ex Pfeiff. & Otto, Opuntia dejecta Recent classifications of Cactaceae recognize Salm-Dyck, Opuntia lindheimeri Engelm., O. four subfamilies (Anderson, 2001; Wallace & Gib - monacantha Haw und O. robusta J. C. Wendl. ex son, 2002; Nyffeler, 2002) including Cactoideae Pfeiff. Für jede dieser Arten werden kurze with 1,222 species (the highest diversity) where

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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. species have stems with ribs or tubercles and aroles 2009; Guiggi, 2008, 2010, 2014; Guillot et al., 2014; without glochids (Gibson & Nobel, 1986) and Op - Verloove, 2016; Verloove et al., 2017a & b). In untioideae with about 186 species where species North Africa, very few papers have dealt with the are articulated cacti with flat, cylindrical or globu - inventory of Cactaceae (Véla, 2013), hence the orig - lar stem-segments (Gibson & Nobel ,1986; Ter - inality of the present work where 12 taxa are pre - razas-Salgado & Mauseth, 2002). sented and illustrated, most of them not previously The first reports of the cultivation of cacti in Eu - reported for the non-native Tunisian or even North rope date back to the 1500’s, following their intro - African flora. Several species, e.g. Opuntia dejecta duction from the Americas (Howard & Touw, 1981) Salm-Dyck, Opuntia lindheimeri Engelm. O. and soon became common in European collections monacantha Haw. and O. robusta H.L. Wendl. ex and gardens (Anderson, 2001). Today, Europe pro - Pfeiff. are here reported for the first time from duces millions of propagated cacti per year from its North Africa. horticultural industry. However, there remains a persistent demand for species collected from the Materials and Methods wild (McGough et al., 2014). Thus, many introduced Field work carried out by the first and the succulents have been reported as alien species (see fourth authors ( pro parte ) within Central and e.g., Guillot & van der Meer, 2004; Guillot & Lodé, Northern Tunisia (North Africa), mostly between

36 Bradleya 38/2020 2012 and 2018, has revealed new national and even Bab-Bnat, one single population on the roadside, North African records. All records here reported are 36°47′51″N, 10°09′51″ W, 67m a.s.l., 05.03.2018, R. documented by short descriptions of each species. El Mokni s.c. ; Sousse: towards motorway, one sin - The actual degree of naturalization for each taxon gle extended population on the roadside, is assessed based on numerous sources such as 35°50′01″N, 10°33′52″ W, 45m a.s.l., 02.01.2017, R. Richardson et al. (2000), Occhipinti-Ambrogi & El Mokni s.c. ; Monastir: Menzel Harb towards Galil (2004), Richardson & Pyšek (2006), Pyšek et Monastir’s airport, one huge population growing on al. (2004) or Richardson (2011). Further comments the roadside, 35°45′49″N, 10°43′12″ W, 13m a.s.l., on distribution, habitats occupied, etc. and field 11.03.2018, R. El Mokni s.c. photographs of most of them are also presented. The second and third authors assisted with the Probably native only to the Andes of Peru, Aus - identification of most of the species. trocylindropuntia subulata is now one of the most The paper is divided in two parts in both of widely distributed opuntioids of South America, which the taxa are arranged in alphabetical se - particularly in Argentina and Bolivia where it is quence. Nomenclature of the taxa presented is widely cultivated as natural fencing. It has widely mostly in accordance with recent sources (cf. APD, naturalised in parts of southern Europe and in the 2019; Korotkova & Raab-Straube, 2017; online tropics where it was initially introduced as an or - database http://www.llifle.com/Encyclopedia/ namental. CACTI/Family/Cactaceae/). Description: Stem (trunk) 6 to 10cm in diame - ter, the old bark smooth and brown, its areoles Results bearing clusters of 8 spines or more, branches nu - 1. OpuntIOIDeAe SuBFAMIly merous, elongate, unsegmented, 30–50cm long, 4– 1.1 Austrocylindropuntia subulata (Muehlenpf.) 7cm in diameter, more or less clustered but not Backeb., Cactaceae Jahrb. Kakteen . Pereskioideae whorled, at first almost at right angles to main Opuntioideae 1941(2 ): 12. 1942. (Figure 1) stem but soon erect, somewhat fragile, bright ≡ Opuntia subulata (Muehlenpf.) Engelm., Gard. green. Areoles, shortly woolly, sitting in the retuse Chron. 19 : 627. 1883. grooves on upper parts of the tubercles. ≡ Pereskia subulata Muehlenpf., Allgem. Garten - Leaves, persistent for more than a year, green, zeitung 13 : 347. 1845. more succulent than on other species, awl-like, ≡ subulata (Muehlenpf.) F.M. nearly at right angles to branch, straight or some - Knuth in Backeb. & F.M. Knuth, Kaktus-ABC : 121. what bowed above, nearly terete, pointed, 5 to 12cm 1936. long, grooved on the underside. Spines, usually 1 or New for the non-native flora of Tunisia and sec - 2 or sometimes spineless (more numerous in older ond record for the flora of North Africa (APD, 2019; stems), slender, erect, strong, straight, pale yellow Korotkova & Raab-Straube, 2017). to greyish white, to 8cm long. Flowers borne toward the ends of the branches, red, not opening widely to TUNISIA: Jendouba: Route Bulla Regia, one single 6cm long. Pericarpels long, tuberculate, awl-like, population on the roadside, 36°31′20″N, with erect scales to 2cm long. Sepals reddish, 08°45′05″ W, 169m a.s.l., 24.03.2018, R. El Mokni minute, 4 to 8mm long or less. Petals broader than s.c. ; idem , Jleyleya, one single extended population the sepals, orange-red or greenish yellow. Style on the roadside, 36°32′57″N, 08°43′06″ W, 206m rose-red except for the whitish base, including the a.s.l., 13.10. 2004–01.08.2017, R. El Mokni s.c. ; stigma-lobes about 3cm long, about as long as the Route Boussalem, after the Bridge “Five Dinars”, longest stamens. Stigma-lobes 5 or 6, slender, or - 36°24′ 02″N, 08°53′58″ W, 157m a.s.l., 01.08.2017, ange-yellow. Fruits, ovoid to oblong to club shaped, R. El Mokni s.c. ; Tabarka, one single population on more or less persistent, sometimes spiny, 6 to 10cm the roadside towards touristic route, 36°57′00″N, long, leafy, with a deep umbilicus, sometimes pro - 08°46′20″ W, 29m a.s.l., 24.10.2014-24.01.2018, R. liferous. Seeds few, large, 10 to 12mm long (re - El Mokni s.c. ; Beja: Ouechtata, one single popula - vealed from an online database tion on the roadside, 36°57′35″N, 08°59′11″ W, 69m http://www.llifle.com/Encyclopedia/CACTI/Fam - a.s.l., 26.03.2018, R. El Mokni s.c. ; towards Mjaz El ily/Cactaceae/4092/Austrocylindropuntia_subu - Bab, one single population on the roadside, lata). Two subspecies are sometimes recognised, 36°39′40″N, 09°30′54″ W, 154m a.s.l., 01.08.2017, R. subsp. subulata and subsp. exaltata (A. Berger) El Mokni s.c. ; Bizerta: one little population within D.R. Hunt. landscaping, 37°17′26″N, 09°52′17″ W, 07m a.s.l., R. Austrocylindropuntia subulata is mentioned El Mokni s.c. ; Route Sejnan Nefza, one single pop - here for the first time from Tunisia , and for the sec - ulation on the roadside, 37°04′36″N, 09°10′27″ W, ond time as naturalised in North Africa (APD 2019; 142m a.s.l., 26.03.2018, R. El Mokni s.c. ; Mateur, Korotkova & Raab-Straube, 2017). In Europe, it is route towards Sidi Nsir, one huge population on the recorded as a naturalised alien in Italy, Sardinia roadside, 37°02′03″N, 09°39′18″ W, 43m a.s.l., and Sicily, as alien (status unknown) in Portugal, 13.09.2004- 26.03.2018, R. El Mokni s.c. ; Tunis: and as a casual alien in Corsica (Korotkova &

Bradleya 38/2020 37 Figure 2. Cylindropuntia imbricata , Bouhedma (Sidi-Bouzid), abandoned land, 24th April 2017.

Raab-Straube, 2017). In the Canary Islands it is magenta, rarely rose-pink, borne at ends of commonly naturalised on all islands except El Hi - branches, 4 to 6cm long, sometimes 8 to 9cm broad, erro (Acebes Ginovés et al., 2010). purple; ovary tuberculate, bearing a few bristles from some of the upper areoles, spineless, occasion - 1.2 Cylindropuntia imbricata (Haw.) F.M. ally bearing a few bristles from some of the upper Knuth in Backeberg & Knuth, Kaktus-ABC : 125. areoles; perianth purple; anthers yellow on purple 1936. (Figure 2) filaments; stigma pale yellow; fruit naked, egg- ≡ Cactus imbricatus Lem., Les Cactées : 88. 1868. shaped with a hollow at the wide end where the ≡ Cereus imbricatus Haw., Suppl. Pl. Succ .: 70. flower fell off, yellow, 2.5 to 4.4cm long, 2–4cm in di - 1819. ameter, strongly tuberculate or, when long persist - ≡ Grusonia imbricata (Haw.) G.D. Rowley, Tephro - ent, smooth; seeds 2.5 to 3.5(–4)mm in diameter. In cactus Study Group 12 (3): 44. 2006. addition to the typical variety (var. imbricata ), two ≡ Opuntia imbricata (Haw.) DC., Prodr. 3: 471. varieties are recognised from West : var. ar - 1828. gentea (M.S. Anthony) Backeb. with silvery spines, Confirmed for the non-native flora of Tunisia as and var. arborescens (Engelm.) A.D. Zimmerman naturalised (APD 2019; Korotkova & Raab-Straube which is taller (Britton & Rose, 1919). 2017). The species is naturalised in arid regions and along watercourses in South America, South Africa, TUNISIA: Sidi-Bouzid/Gafsa: some populations the Mediterranean basin and Australia. In North within the National park of Bouhedma, 34°28′18″N, Africa, C. imbricata is reported as an introduced 09°28′36″ W, 237m a.s.l., 24.04.2017, R. El Mokni cultivated alien (status unknown) for Morocco, s.c. Tunisia and Libya (APD, 2019; Korotkova & Raab- Cylindropuntia imbricata is a cactus found in Straube, 2017). In the Canary Islands it is only the southwestern United States and northern Mex - known from a single locality in Gran Canaria (Ver - ico (Durango, Zacatecas, and San Luis Potosí), in - loove et al., 2017a) cluding some cooler regions in comparison to many other cacti. It occurs primarily in the arid regions of 1.3 Opuntia ficus-indica (L.) Mill. f . amyclaea the Southwestern United States in the states of (Ten.) Schelle, Handb. Kakteenkult : 51. 1907. Texas, New , and . Nomenclature according to Kiesling (1998). (Figure Description: Tree-like, often 3m high or higher, 3) with a more or less definite woody trunk 2.5cm in ≡ Opuntia amyclaea Tenore, Fl. Neap. Prodr. App . diameter; ultimate joints 2 to 3cm in diameter, 5: 15. 1826. strongly tuberculate; leaves 8 to 24mm long, terete; = O. megacantha Salm-Dyck, Hort. Dyck.: 363. tubercles 2 to 2.5cm long, flattened laterally; spines 1834. 8 to 30, 2 to 3cm long, brown, covered with papery = O. maxima Salm-Dyck, Cact. Hort. Dyck.: 68. sheaths; flowers bowl-shaped, diurnal, purple or 1850, non Mill.

38 Bradleya 38/2020 Figure 3. Opuntia ficus-indica f. amyclaea , Monastir city within the coastal side, clayey slope, 7th April 2018.

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

TUNISIA: Monastir, scattered individuals growing disappearing; flowers yellow; fruit yellowish red, on a sun-exposed clayey slope close to the coast, not very juicy (cf. Britton & Rose, 1919). 35°46′54″N, 10°48′12″ W, 40m a.s.l., 04.12.2017, R. This taxon is new to the flora of Tunisia and a El Mokni s.c. second record for the flora of North Africa (APD, 2019). In Tunisia, O. ficus-indica f. amyclaea can Description: Compared with Opuntia ficus-in - obviously be classified as well naturalised as it ap - dica it has oblong to suborbicular, glaucous-pru - pears in several subpopulations. However, like inose, distinctly flattened cladodes that are always other species from this genus, it certainly has the spiny. Molecular data seem to suggest that it prob - potential to spread and become an invasive alien. ably is nothing else than the wild progenitor of do - mesticated O. ficus-indica (Labra et al ., 2003). 1.4 Opuntia dejecta Salm-Dyck, Hort. Dyck .: 361. Other authors, however, accept it as a species dis - 1834. (Figure 4) tinct from O. ficus-indica , based on morphometric ≡ Nopalea dejecta (Salm-Dyck) Salm-Dyck, Cact . analyses (Reyes-Agüero et al ., 2005). Forma amy - Hort. Dyck. 1849 : 64. 1850. claea has an erect habit; joints oblong to elliptic, 30 to 40cm long, about twice as long as broad, thick, TUNISIA: Monastir, 35°46′03″N, 10°50′07″ W, 38m dull green, a little glaucous; leaves few mm long, a.s.l., abandoned land, 18.11.2017, R. El Mokni s.c. acute, red; areoles small, with 1 or 2 short bristles from the lower parts of areoles; spines 1 to 4, nearly Probably a native of Panama, this species is porrect, usually less than 3cm. long, white or horn- very common in cultivation in tropical America coloured, the stoutest angled; glochids brown, soon (Britton & Rose, 1919).

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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.

40 Bradleya 38/2020 Description: Opuntia dejecta is a plant of 1 to These are some millimeters across and bear small 2m in height, with a definite trunk, very spiny, the scale-like leaves on young joints, later on 1 to 10 old areoles often bearing 6 or 8 spines; joints nar - spines each. The spines are very variable in colour row, 10 to 15cm long, only moderately thick, often (yellow to brown) and length (1.5 to 6cm) and can drooping, bright green even in age, bearing usually sometimes be lacking. They spread in all directions two somewhat spreading spines at an areole; spines and are accompanied by glochids, short barbed bris - at first pale yellow or pinkish, in age grey, the tles which form striking tufts (Backeberg, 1970; longest 4cm long; flower, including ovary and style, Benson, 1982; Ellenberg, 1989; Anderson, 2005). is 6–7cm long; sepals obtuse; petals erect, dark red; At flowering, O. dillenii plants have 6 to 8cm stamens long-exerted, dark red. (Britton & Rose, wide flowers that are finally lemon yellow in colour 1919). but can be slightly reddish in early stages of devel - Opuntia dejecta is new to the flora of Tunisia, opment. The purple-skinned fruits are pear-shaped the flora of Africa (APD, 2019) and is recorded for (Figure 5D) measure 4 to 7cm in length and about the second time for the Mediterranean area, where 3cm in diameter. They normally show no spines it was so far reported only as naturalised in Sicily and their glochids mostly fall off from the few are - (Korotkova & Raab-Straube, 2017). In Tunisia, O. oles at maturity. The juicy flesh of fruits is purple dejecta is obviously a recent escape and can be clas - in colour and contains many rounded seeds which sified only as casual. However, like other species are –with an arillus-like envelop - about 4mm in di - within this genus, it certainly has the potential to ameter and tan coloured (Backeberg, 1970; Benson, spread to the nearby coastal regions. 1982; Ellenberg, 1989; Anderson, 2005; Land Pro - tection, 2006). 1.5 Opuntia dillenii (Ker Gawl.) Haw., Suppl. Pl. There is considerable dispute whether Opuntia Succ .: 79. 1819. (Figure 5) dillenii belongs in Opuntia stricta or should be con - ≡ Cactus dillenii Ker Gawl., Bot. Reg . 3: t. 255. sidered distinct (Böhm, 2008). Opuntia dillenii dif - 1818. fers from O. stricta mainly by its unarmed stem ≡ Opuntia stricta (Haw.) Haw. var. dillenii (Ker segments with groups of 1–7 large spines on most of Gawl.) L.D. Benson, Cact. Succ. J. (Los Angeles) 41 : the areoles. Both are low-growing plants (usually 126. 1969. 50–100cm tall) with relatively large flattened and elongated (i.e. elliptic or obovate) stem segments. TUNISIA: Monastir, scattered populations growing Opuntia dillenii originates from southeastern on a sun-exposed clayey slope of the rocky coast in parts of North America, the east coast of Mexico, the north east of Monastir, 35°46′54″N, 10°48′12″ W, the Bermuda’s, the West Indies and from the north 40m a.s.l., 04.12.2017, R. El Mokni s.c. ; idem , scat - of South America (Britton & Rose, 1919). Wild pop - tered individuals growing on the sandy coast at the ulations are nowadays reported in the same areas north east of Monastir, 35°44′57″N, 10°49′36″ W, (Backeberg, 1970; Benson, 1982; Schultze-Motel 27m a.s.l., 12.11.2017, R. El Mokni s.c. ; idem , scat - ,1986; Scheinvar, 2002; Anderson, 2005) and addi - tered massive populations/subpopulations growing tionally at the Gulf Coast of Texas (Benson, 1982), on both sides of the Monastir Sousse tourist route, the southeastern coastal margin of Brazil (Schein - 35°45′49″N, 10°43′12″ W, 13m a.s.l., 11.03.2018, R. var, 1984), and in Ecuador (Anderson, 2005). The El Mokni s.c. ; idem , scattered subpopulations grow - native locations of O. dillenii are mostly at an alti - ing on both sides of the Menzel Kamel-Ouerdenine tude near sea level in sand dunes, at the edges of route, 35°38′26″N, 10°39′37″ W, 75m a.s.l., maritime forests, or associated with tropical plants 11.03.2018, R. El Mokni s.c. ; Bizerta, widely culti - (Benson, 1982; Pinkava, 2003). O. dillenii was in - vated as ornamental in pots and public gardens, troduced into many parts of the world. It is found escaping to roadsides or coastal rocks, 37°19′52″N, around the Mediterranean (Schultze-Motel, 1986; 09°51′52″ W, 12m a.s.l., R. El Mokni s.c. ; Mateur- Gentile, 1991), on the Canary Islands (Perfumi & massif of Ichckeul, several scattered massive popu - Tacconi, 1996; Loro et al., 1999; Anderson, 2005; lations growing everywhere within the massif of Díaz Medina et al., 2007), in the west and east of Ichckeul, including the National Park, 37°07′27″N, Africa (Schultze-Motel, 1986), on Madagascar and 09°39′36″ W, 33–476m a.s.l., 24.03.2014, R. El Mokni Mauritius (Schultze-Motel, 1986; Ellenberg, 1989), s.c. ; Sejnane–Tamera, scattered subpopulations in North Yemen (Ellenberg, 1989), in India (Backe - growing on roadsides, 37°04′41″N, 09°08′57″ W, berg, 1970; Badami & Thakkar, 1984;, Gupta et al., 122m a.s.l., 12.04.2018, R. El Mokni s.c. 2002), in the southeast of Asia (Schultze-Motel Plants of O. dillenii sprawl or grow erect and ,1986; Jiang et al., 2006), and in Australia (Backe - reach a height of about 50 to 200cm. They consist of berg, 1970; Schultze-Motel, 1986; Ellenberg ,1989). flattened, elliptic and obovate stem segments, and Opuntia dillenii becomes naturalised under condi - these are impressive for many stout spines. The tions different from those in its original areas and joints are dull green or bluish-green in colour, 10 to can even rapidly spread out (Ellenberg, 1989). It is 40cm long, 5 to 15cm broad, 1 to 2cm thick, and ir - involved in dangerous infestations with several Op - regularly trimmed with relatively few areoles. untia species (Burdon & Marshall, 1981), notably

Bradleya 38/2020 41 Figure 6. Opuntia lindheimeri , Monastir city, roadsides under plantation of Ficus benjamina , 15th Feb - ruary 2018. O. stricta (pest pear) in eastern Australia (Free - (ii) thorns completely remaining yellowish whereas man, 1992; GISD, 2010). thorns are brown at the base and the rest yellow in In Tunisia, Opuntia dillenii can be considered the case of O. engelmannii (see e.g. Vázquez & Gar - an invasive species in many regions of the country. cía, 2017). New to the flora of Tunisia and the flora of 1.6 Opuntia lindheimeri Engelm., Bost. J. Nat. Africa, O. lindheimeri has been reported before in Hist. 6: 207. 1850. (Figure 6) Europe from Azerbaijan and Georgia in Transcau - ≡ Opuntia engelmannii subsp. lindheimeri (En - casia (Korotkova & Raab-Straube, 2017) as well as gelm.) U. Guzmán & Mandujano, Cactaceae Syst. from Italy and Spain (Sanz-Elorza et al., 2006; Init . 16 : 18. 2003. Guiggi, 2014). It is also known from a single popu - lation in Gran Canaria in the Canary Islands (Ver - TUNISIA: Monastir, 35°46′00″N, 10°49′37″ W, 40m loove et al., 2017a). In Tunisia, O. lindheimeri is a.s.l., roadside, a single individual, 15.02.2018, R. obviously a recent escape and can be classified only El Mokni s.c. as casual. However, like other species within this genus, it certainly has the potential to spread to the A species native to Southwestern Louisiana, nearby coastal regions. southeastern Texas (U.S.A.), and Tamaulipas (Mexico). 1.7 Opuntia microdasys (Lehm.) Pfeiff., Enum. Description: Opuntia lindheimeri has an erect Diagn. Cact .: 154. 1837. (Figure 7). habit, 2 to 4m high, with a more or less definite ≡ Cactus microdasys Lehm., Index Seminum (Ham - trunk, but at times much lower and spreading; burg) , 1827 : 16. 1827. joints green or bluish green, somewhat glaucous, orbicular to obovate, up to 2.5cm long; leaves subu - TUNISIA: Bizerta: abandoned plantations in many late, 3 to 4mm long, somewhat flattened, pointed; sites, 37°17′26″N, 09°52′17″ W, 7m a.s.l., R. El areoles distant, often 6cm apart; spines usually 1 Mokni s.c. ; Monastir: scattered little subpopula - to 6, often only 2, one porrect and 4cm long or more, tions growing on both sides of the Monastir Sousse the others somewhat shorter and only slightly tourist route, 35°45′49″N, 10°43′12″ W, 13m a.s.l., spreading, pale yellow to nearly white, sometimes 11.03.2018, R. El Mokni s.c. ; Sidi-Bouzid/Gafsa: brownish or blackish at base, or some plants spine - some populations in landscaping within National less; glochids yellow or sometimes brownish, usu - park of Bouhedma, 34°28′18″N, 09°28′36″ W, 237m ally prominent; petals usually yellow (sometimes a.s.l., 24.04.2017, R. El Mokni s.c. red); stigma-lobes usually green; fruit purple, pyri - form to oblong, 3.5 to 5.5cm long. It is an extremely An endemic of Mexico, it became an invader in variable species, composed of many races, differing Australia and South Africa (Walters et al ., 2011; mainly in armament, colour of flowers, size and Novoa et al ., 2014). shape of joints and of fruit (Britton & Rose, 1919). Description: Opuntia microdasys (bunny ears) A very close taxon compared to Opuntia engel - is a branched, shrubby succulent perennial, erect mannii and still considered as a synonym by many to spreading, up to 1m tall. Stem segments pale authors, O. lindheimeri is mainly characterized by green, brighter green when young, round to oval- (i) the presence of flowers with coloured orange shaped, 8–16 × 7–16 cm. Areoles prominent, round, tepals versus yellow tepals in O. engelmannii , and close-set. Needle-like spines usually absent, some -

42 Bradleya 38/2020 Figure 7. Opuntia microdasys , Bouhedma (Sidi-Bouzid), abandoned land, 24th April 2017. times a single short one present on an areole. TUNISIA: Bizerta: roadside, a single individual, Glochids dense, fine, bristle-like, yellow, white or 37°16′22″N, 09°52′25″ W, 31m a.s.l., 15.02.2018, R. brown. Flowers 40mm in diameter, petals to 40mm El Mokni s.c. ; Monastir: roadside, a single individ - long, yellow with a slight reddish tint towards the ual, 35°46′00″N, 10°49′37″ W, 40m a.s.l., 03.12.2017, slightly frilly tips of the petals. Fruits 30–40mm in R. El Mokni s.c. diameter, fleshy, egg-shaped to globose, red to pur - plish-red, densely covered with areoles bearing This South American species (Argentina and dense clusters of glochids (Smith et al., 2011; Bárce - Brazil) is widely cultivated as an ornamental. nas et al., 2017). Description: Plant 2 to 4 or even 6m high, often New to the flora of Tunisia and a second record with a definite trunk, usually with a large much for the flora of North Africa (APD, 2019), Opuntia branched top; trunk cylindric, 15mm in diameter, microdasys is widely cultivated as a popular and at - either spiny or smooth; joints ovate to oblong, nar - tractive ornamental cactus. It is often recorded as rowed at base, 10 to 30mm long, bright shining an escape from cultivation, either naturalised or green; leaves subulate, 2 to 3mm long; areoles filled adventitious (casual), in Europe at least in France, with short wool; glochids brownish; spines 1 or 2, Italy, Portugal, and Spain (Essl & Kobler, 2008; sometimes more (on the trunk often 10 or more) Novoa et al ., 2014; Korotkova & Raab-Straube from an areole, erect, 1 to 4cm long, yellowish 2017; Verloove et al., 2017a& b). brown to dark reddish brown; flowers yellow to red - dish, 7.5cm broad; sepals broad, each with a broad 1.8 Opuntia monacantha (Willd.) Haw., Suppl. red stripe down the middle; petals golden yellow, pl. succ .: 81. 1819. (Figure 8). widely spreading; filaments greenish; style white; ≡ Cactus monacanthos Willd., Enum. pl . 33. 1813 stigma-lobes 6, white; ovary spineless, 3.5cm long; (1814). fruit obovoid, 5 to 7.5cm long, reddish purple, long- = Opuntia vulgaris Britton & Rose, non Mill., Cact. persisting, sometimes proliferous (Britton & Rose, 1: 156. 1919. 1919). New for the flora of Tunisia and the flora of North New to the flora of Tunisia and for the flora of Africa. North Africa (APD, 2019), Opuntia monacantha is

Bradleya 38/2020 43 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. cultivated as an attractive ornamental cactus. In a.s.l., 11.03.2018, R. El Mokni s.c. Europe, it is recorded as naturalised at least in France, Corsica, Canary Islands, as alien in Sicily Opuntia robusta is widely distributed in north - and doubtfully present in Switzerland, Italy, Por - ern and central Mexico (, Durango, Nuevo tugal, and Spain (Korotkova & Raab-Straube, León, San Luis Potosí, Tamaulipas, Aguascalientes, 2017). Guanajuato, Guerrero, Hidalgo, México, Querétaro provinces). In the United States it is found in Ari - 1.9 Opuntia robusta J.C. Wendl. Cat. Hort. Her - zona. This species has expanded into North and renh . 1835 (Figure 9). South America and has been introduced into New = Opuntia camuessa F.A.C. Weber, Dict. Hort. Zealand and the Mediterranean region of southern [Bois] 2: 895. 1898. Europe, South Africa, Madagascar and Australia New for the flora of Tunisia and a first record for where it is a significant environmental and pastoral the flora of North Africa (Hansen & Sunding, 1993; weed, particularly in the states of , APD, 2019). , Queensland and Victoria. Description: Opuntia robusta , popularly known TUNISIA: Monastir: two subpopulations of few in - as the ‘wheel cactus’ in reference of the circular dividuals, growing on both sides of the Monastir shape of the segments, is a much branched, often Sousse tourist route, 35°45′49″N, 10°43′12″ W, 13m erect, succulent shrub or small tree, sometimes 4–

44 Bradleya 38/2020 6m high, but commonly 1–2m, with more or less dis - Native to Argentina, Brazil (Mato Grosso do tinct trunks. The branch segments are armed with Sul, Minas Gerais, Paraná, Rio de Janeiro, Rio tufts of numerous barbed bristles (glochids) and Grande do Sul, Santa Catarina, São Paulo), Para - commonly spineless or with 1 to 5 sharp spines to guay and Uruguay (Anderson, 2001) . 4cm long arising from each areole. Flowering in Description: stems, cylindrical, segmented, blue spring (April to May), both plants with male, female green to dull green, often glaucous when young, to and hermaphrodite flowers can be found in Opuntia 15cm in diameter; ribs, (4–)5–6(–8), thin to obtuse, robusta . The fruits (in autumn, September to Octo - slightly indented, 2.5–3.5cm high, rounded, green ber), which may be red, orange, yellow or green, are often with large yellow patches along the sides; are - globular to ellipsoid, at first more or less tubercu - oles, distant, brown, large 2cm apart; spines, usu - late, 7 to 9cm long, deep red, but also orange, yel - ally absent, occasionally a few golden or brown low or green, with light brown felt and yellow spines develop afterwards; flowers, very large, noc - glochids c. 3mm long, produced in profusion, possess turnal, elongated, funnelform, 20 to 30cm long; thick peel (with sharp glochids) and juicy red pulp inner perianth-segments white, broad and obtuse containing numerous seeds; they are edible but best ovary naked, 2.5 to 3cm long, blooming at night in for animal feeding (revealed from an online data - spring through summer. The buds grow quickly, base http://www.llifle.com/Encyclopedia/CACTI/ shooting out from the branches like so many small Family/Cactaceae/32724/Opuntia_robusta). snakes. Within two weeks, dozens of flowers open, New to the flora of Tunisia and a first report for always at night and all or nearly all flowers open the flora of North Africa (APD, 2019), Opuntia ro - simultaneously at the same time. By mid-morning busta has been reported from Gran Canaria, La the following day, all had closed and drooped. This Palma, and Tenerife in the Canary Islands (Acebes is apparently a water-saving strategy by cacti. The Ginovés et al. , 2010) although many claims require plant usually buds and flowers about two weeks fol - confirmation, resulting from confusion with the lowing any warm-season rain. The fruits usually morphologically similar O. ficus-indica f. amyclaea ripen within a month. Buds may erupt well into (Verloove et al., 2017a & b). In Europe, it is only fall, even early winter, if sufficiently warm and rarely reported in the wild (Guiggi, 2014). It is rains fall. During dry times, the buds often fall off recorded as naturalised in France, Corsica and prior to opening; fruits globose, pear-shaped, red Sicily and as a casual alien in Italy (Korotkova & with white pulp. The Peruvian apple cactus may Raab-Straube, 2017). Moreover, it is reported as an produce fruit 3–4 years after propagation from seed invasive weed in several countries, including Aus - (revealed from an online database http://www. tralia, Botswana, and South Africa (Walters et al ., llifle.com/Encyclopedia/CACTI/Family/Cac - 2011; Novoa et al ., 2014). taceae/6499/Stenocereus_peruvianus). , This species is widely cultivated worldwide as CACtOIDeAe SuBFAMIly an ornamental plant. It has escaped and sometimes 2.1 Cereus hildmannianus K. Schum., Fl. Bras. became invasive. In Europe, Cereus hildmannianus 4(2): 202. 1890. (Figure 10). has been reported before from Italy (Guiggi, 2010); = Cereus peruvianus auct. non (L.) Mill. it was also recently reported from Canary Islands New for the non-native flora of Tunisia and for the as casual alien (Verloove et al., 2017b; Korotkova & flora of North Africa (APD, 2019; Korotkova & Raab-Straube, 2017). Raab-Straube, 2017). 2.2 Echinopsis eyriesii (Turpin) Zuccarini in TUNISIA: Monastir: some scattered Pfeiffer and Otto, Abbild. Beschr. Cact . 1: under pl. individuals/subpopulations in center of the town 4. 1839. (Figure 11). within some public gardens, 35°46′13″N, ≡ Echinocactus eyriesii Turpin, Ann. Inst. Roy. 10°49′39″ W, 41m a.s.l., 03.10.2015-30.03.2018, R. Hort . Fromont 2: 158. 1830. El Mokni s.c. ; Jemmel, 35°37′43″N, 10°45′32″ W, ≡ Cereus eyriesii Pfeiffer, Enum. Cact .: 72. 1837. 41m a.s.l., 03.10.2015-30.03.2018, R. El Mokni s.c.; ≡ Echinonyctanthus eyriesii Lemaire, Cact. Gen. idem 35°37′28″N, 10°45′32″ W, 57m a.s.l., Nov. Sp. : 84. 1839. 03.10.2015–04.04.2018, R. El Mokni s.c. ; Sousse: ≡ Echinopsis pudantii Pfersdorff, Monatsschr. Kak - Bouhsina, one huge population on an ancient ex - teenk . 10 : 167. 1900. tended building, 35°49′15″N, 10°37′33″E, 71m a.s.l., New for the non-native flora of Tunisia, for the flora 27.12.2017, R. El Mokni s.c. ; Tunis: Mornaguia, one of North Africa and for the flora of the Mediter - vigourous population on the right of the motorway, ranean area (APD, 2019; Korotkova & Raab- 36°45′21″N, 09°59′19″ W, 66m a.s.l., 24.03.2018, R. Straube, 2017) El Mokni s.c. ; Bizerta: one small population within TUNISIA: Monastir: some scattered individuals in on the right of the road towards Jarzouna from an abandoned field with a Chenopodiaceae/Ama - Menzl Jemil, 37°15′29″N, 10°37′33″ W, 28m a.s.l., ranthaceae vegetation alongside the road towards 27.12.2017, R. El Mokni s.c. Sousse, 35°45′34″N, 10°44′24″ W, 29m a.s.l., 03.10.2017, R. El Mokni s.c. ; Tunis: Ariana, some

Bradleya 38/2020 45 A C

B

D e

Figure 10. Cereus hildmannianus , A. B. & C. Jemmal (Monastir), old buildings and roadsides, 13th April 2018; D. & e. Ksar-Hellal (Monastir), roadsides, 12th April 2018. scattered individuals, 36°50′40″N, 10°11′54″ W, 25m ovate, brownish, hairy in their axils (cf. Britton & a.s.l., 10.12.2017, R. El Mokni s.c. Rose, 1922). Originating from Argentina (Buenos Aires). It has a large distribution within Southern Brazil, 2.3 Hylocereus undatus (Haw.) Britton & Rose, Uruguay, and the province of Entre Rios, Ar - Fl. Bermuda : 256 (fig. 278). 1918. (Figure 12). gentina. (cf. Britton & Rose, 1922 ). ≡ Cereus undatus Haw., Philos. Mag. Ann. Chem. 7: Description: Echinopsis eyriesii is a simple or 110. 1830. clustered, globular to short-columnar cactus; ribs New for the non-native flora of Tunisia and for the 11 to 18, not tuberculate, rather thin above; areoles flora of North Africa. (APD, 2019; Korotkova & circular, filled with white or tawny wool; spines sev - Raab-Straube, 2017). eral, 14 to 18, very short, 5mm long or less; flower TUNISIA: Monastir: one epiphytic population on from the side of plant but above the middle, large, planted Phoenix canariensis within an old building 17 to 25cm long; inner perianth-segments white, in the centre of the town, 35°46′03″N, 10°49′55″ W, acuminate; stamens and style shorter than the pe - 40m a.s.l., 27.02.2018, R. El Mokni s.c. ; Tunis–Ari - rianth-segments; scales on the flower-tube small, ana: escaped plants from ancient planted individu -

46 Bradleya 38/2020 A

B

Figure 11. Echinopsis eyriesii , A. Ariana city, abandoned land, 16th November 2017; B. Monastir city, coastal area, clayey slope, 24th April 2018. als in pots within an old building, 35°50′50″N, 5cm wide; areoles about 4.5cm apart; spines 1 to 3(– 10°11′36″ W, 34m a.s.l., 15.11.2017, R. El Mokni s.c. 5), conical and up to 1cm long (but usually about 2– 3mm long); flowers ornate, fragrant and beautiful, This species is widely cultivated and has be - 25–35cm long by 30cm across, white with green come naturalised in many countries, however, as outer tepals and bracts, blooming only at night, and the native range is unknown, nothing is known usually the flower lasts just one night. In tropical about the size and trends of the native wild popu - climates the plants can have up to 4–6 lation (Durán et al., 2017). flowering/fruiting cycles per year. In temperate cli - Description: Hylocereus undatus is a sprawling mates they bloom in late spring to early summer; or vining, terrestrial or epiphytic cactus. It climbs once pollinated it produces edible red fruits with by use of aerial roots and can reach a height of 10m white pulp up to 12cm long (revealed from an on - or more growing on rocks and trees. It is very vari - line database http://www.llifle.com/Encyclope - able and closely related to Selenicereus . dia/CACTI/Family/Cactaceae/7429/Hylocereus_und Stems, green and 3-winged, from a few cm up to 5m atus). long (in mature plants), 4 to 7.5cm wide with mar - The native range of this species is unknown as gins undulate and horny with wings that are 2.5 to it has long been in cultivation (Anderson, 2001;

Bradleya 38/2020 47 A B C

Figure 12. Hylocereus undatus , A. & C. Tunis (Ariana), old buildings, 30th September 2017; B. Monastir city, epiphytic on Phoenix canariensis , 1st March 2018.

Hunt et al ., 2006). Some believe it to be native to APD (AFRICAN PLANT DATABASE ) (2019). Opuntia southern Mexico, the Pacific side of Guatemala, Mill. In African Plant Database. – Published at: Costa Rica, and El Salvador, however, this has not http://www.ville-ge.ch/musinfo/bd/cjb/africa/de - been confirmed. It is the most widely cultivated tails.php?langue=an&id=193208. Accessed on species in the genus (including commercial cultiva - 24 May 2019. tion) and often escapes and becomes naturalised. ANDERSON , E.F. (2001). The Cactus Family , Port - This species is therefore widely distributed and has land, Oregon: Timber Press, pp. 144–145. been recorded from tropical forests in Mexico, var - ANDERSON , E.F. (2005). Das große Kakteen- ious islands in the Caribbean, Central America, Lexikon. Ulmer-Verlag, Stuttgart. tropical South America, and southeastern BACKEBERG , C. (1970). Das Kakteenlexikon . Gustav Asia. The type collection is from a cultivated speci - Fischer Verlag, Jena. men grown in London which was probably brought BADAMI , R.C. & THAKKAR , J.K. (1984) Minor Seed from China. In Brazil, it is introduced as a garden Oils XVIII: Examination of Twelve Seed Oils. plant and sometimes escapes into roadside areas Fette Seifen Anstrichmittel 86 : 165–167. and maritime scrub, occurring in Pernambuco, BáRCENAS LUNA , R., G OETTSCH , B.K., G óMEZ -H INOS - Alagoas, Bahia, Minas Gerais and Espírito Santo TROSA , C., G UADALUPE MARTíNEZ , J. & S áNCHEZ , (Taylor & Zappi, 2004). In addition to its wide nat - E. (2017). Opuntia microdasys (amended ver - uralised range, it has also a wide altitudinal range sion of 2013 assessment). The IUCN Red List of occurring from near sea level up to 2750m. (Durán Threatened Species 2017: e.T152311A et al., 2017). In Europe, the species is reported only 121587299. http://dx.doi.org/10.2305/IUCN.UK. from the Canary Islands by Korotkova & Raab- 2017-3.RLTS.T152311A121587299.en. Down - Straube (2017). It is here reported for the first time loaded on 03 April 2018. from North Africa. BENSON , L. (1982). The Cacti of the United States and Canada . Stanford University Press, Stan - References ford. ACEBES GINOVéS , J.R., L EóN ARENCIBIA , M.C., R O- BöHM , H. (2008). “ Opuntia dilleni –An Interesting DRíGUEZ NAVARRO , M.L. et al . (2010). Pterido - and Promising Cactaceae Taxon”. Journal of the phyta, Spermatophyta. In : ARECHAVALETA , M., Professional Association for Cactus Development RODRíGUEZ , S., ZURITA , N. & GARCíA , A. ( EDS ), 10 :148-170. Lista de especies silvestres de Canarias (hongos, BRAUN , P., L AROCCA , J., M ACHADO , M. & T AYLOR , plantas y animales terrestres) . (2nd ed.). Pp. N.P. (2017). Cereus hildmannianus (amended 119–172. Gobierno de Canarias, La Laguna. version of 2013 assessment). The IUCN Red

48 Bradleya 38/2020 List of Threatened Species 2017: GUILLOT , D. & LODé , J. (2009). Opuntia pilifera e.T152594A121480022. http://dx.doi.org/10.230 Weber, primera cita como alóctona en Europa. 5/IUCN.UK.2017-3.RLTS.T152594 Bouteloua 6: 141. A121480022.en. Downloaded on 02 April 2018. GUILLOT , D. & VAN DER MEER , P. (2004). Opuntia BRITTON N.L. & R OSE J.N. (1919). The Cactaceae, ×elisae Guillot & Van der Meer, un híbrido na - Descriptions and Illustrations of plants of the tural de Opuntia ficus-indica Haw. y Opuntia cactus family. Vol. 1. The Carnegie Institution tomentosa Salm. -Dyck var. hernandezii (DC.) of Washington. Washington, 320 p. Bravo. Toll Negre 3: 7–10. BRITTON N.L. & R OSE J.N. (1922). The genus GUILLOT , D., SáNCHEZ -G ULLóN , E. & L ODé , J. (2014). Echinopsis Zuccarini in The Cactaceae, Descrip - Nuevas citas de cuatro cactáceas en la provincia tions and Illustrations of plants of the cactus de Valencia. Bouteloua 19 : 11–17. family. Vol. III , pp. 60–77. 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