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Willdenowia 42 – 2012 Filip VerlooVe1 & enrique Sánchez Gullón2 A taxonomic revision of non-native Cenchrus s.str. (Paniceae, Poaceae) in the Medi-
terranean area
Abstract
Verloove F. & Sánchez Gullón e.: A taxonomic revision of non-native Cenchrus s.str. (Paniceae, Poaceae) in the
Mediterranean area. – Willdenowia 42: 67 – 75. June 2012. – online iSSn 1868-6397; 2012 BGBM Berlin-Dahlem.
Stable url: http://dx.doi.org/10.3372/wi.42.42107 The exact identity of non-native, naturalised populations of Cenchrus s.str. in the Mediterranean area has been criti- cally assessed. A herbarium revision confirmed the presence of three species: Cenchrus echinatus, C. longispinus and C. spinifex (syn.: C. incertus). in the present paper C. echinatus is reported for the first time from Spain and confirmed for egypt and israel. C. longispinus, up to present widely confused with C. spinifex, is reported for the first time from croatia, Greece, israel and Morocco and furthermore confirmed for France (including corse) and italy. Finally, C. spinifex is confirmed for France, italy and Spain, while records from Greece and israel proved to be in error for C. longispinus. All three species are much alike and widely confused in the studied area. in some areas (especially in parts of italy and israel) two species occur sympatrically, which largely added to the confusion. Main features for their distinction are discussed and a dichotomous key for the identification of the native and non-native species of Cenchrus s.str. in the Mediterranean area is presented. Additional key words: Gramineae, Cenchrus echinatus, Cenchrus longispinus, Cenchrus spinifex, taxonomy, chorology The three non-native species of Cenchrus that are fi- nally accepted in this study (C. echinatus, C. longispinus in its traditional circumscription Cenchrus l. is a genus and C. spinifex cav.) are reputed environmental and/or of c. 20 species, predominantly distributed in the warmer agricultural weeds in as well as outside their native dis- parts of the new World and with some additional spe- tribution range. in the Mediterranean area, C. echinatus cies in Africa and Asia (Delisle 1963; Mabberley 2008). is a relatively recent newcomer and only locally estab- Several species are troublesome environmental or agri- lished or invasive (although a future naturalisation in a cultural weeds beyond their native distribution range.
wider area seems likely). Both other species have been Cenchrus is a very complex and largely misunder- repeatedly and increasingly reported as invasive species, stood genus, especially in areas where non-native species for instance in parts of Greece, the Former Yugoslavian occur, as is the case in the Mediterranean area. in the past republic, italy or Spain (Sanz elorza & al. 2004; Borsi6 decades collections of non-native Cenchrus species from & al. 2008; Arianoutsou & al. 2010; celesti-Grapow & the Mediterranean have been ascribed to either C. echi- al. 2010). Therefore, it is not only useful but even essen- natus l., C. incertus M. A. curtis, C. longispinus (hack.) tial to acquire a better understanding of the non-native Fernald, C. pauciflorus Benth. or C. tribuloides l. A her- representatives of the genus Cenchrus in the area under barium revision undertaken in the past years (see also Verloove 2006) yielded a lot of interesting new data. The in the Mediterranean area Cenchrus counts five na- results of this revision are dealt with in this paper. tive species (see for instance Maire 1952; clayton 1 national Botanic Garden of Belgium, Domein van Bouchout, B 1860 Meise, Belgium; e-mail: [email protected] 2 paraje natural Marismas del odiel (huelva), ctra. del Dique Juan carlos i, km 3, Apdo, 720 21071 huelva, españa; e-mail: Verloove & Sánchez Gullón: non-native Cenchrus in the Mediterranean area 1980; Valdés & Scholz 2009+): C. biflorus roxb. (Alge- goslavian republic), Albania, Greece (incl. islands) and ria, egypt, libya and Morocco), C. ciliaris l. (Algeria, Turkey in europe; Turkey, Syria, lebanon and israel in egypt, israel, lebanon, libya, Morocco, Sicily, Syria and Asia and finally egypt (incl. Sinai), libya, Tunisia, Alge- Tunesia; furthermore introduced in Greece and Spain), ria and Morocco in Africa. C. pennisetiformis hochst. & Steud. (egypt), C. prieurii (Kunth) Maire (Algeria, Morocco) and C. setigerus Vahl (egypt). The non-native representatives have been as- Results and discussion
cribed to several species and confusion lingers on since many decades, especially in italy. it surely is no coin- The non-native species of Cenchrus s.str. in the Medi-
cidence that italy is the initial area of naturalisation (in the 1930s) of two non-native species of Cenchrus in the Cenchrus echinatus l., Sp. pl. 2: 1050. 1753.
Mediterranean. populations were initially ascribed to C. tribuloides (plicker 1943; pellegrini 1947; D'errico Distribution. — native of southern united States, central 1949; Tosco & Ariello 1951) and later corrected to and South America and the West indies (Delisle 1963). C. pauciflorus by Grilli (1962). eventually, cecconelli More or less widely naturalised elsewhere in warm-tem- (1975) concluded that all italian collections pertain to perate and (sub-)tropical regions of the world (for in- C. longi spinus (subsequently confirmed by Guzik & pa- stance pacific islands, philippines, Australia, Arabian pe- cyna 1999). however, recent italian floristic checklists ninsula, e Africa, china, etc.). in the Mediterranean area (conti & al. 2005; celesti-Grapow & al. 2010) correctly known so far from egypt (omitted by Valdès & Scholz accept two species: C. spinifex (as C. incertus) and C. 2009+) and israel (Delisle 1963; cope 2005). here re- longi spinus but both are obviously still widely confused ported for the first time from Spain. in israel Cenchrus (see below). in France all records from the past decades echinatus now has become a noxious weed in irrigated were assigned to "C. incertus". however, a previous gardens and lawns (Danin 2004).
revision already proved that all collections from corse in fact represent C. longispinus (Verloove 2006). like- Illustrations. — Fig. 1A–B; caro & Sánchez (1967b); wise, plants from the French Mediterranean area (dep. häfliger & Scholz (1980); Stieber & Wipff (2003); cope Vaucluse) are C. longispinus while genuine C. spinifex & Gray (2009).
is only confirmed from southwestern France. in Spain all populations of non-native Cenchrus have been ascribed Specimens examined. — Egypt: Sinai, nuweiba, so far to "C. incertus" which seems to be confirmed by Straßenrand, 17.3.1995, Borkowsky s.n. (B); Sinai, nu- the present revision (although a second non-native spe- weiba, parkanlage, 13.3.1996, Borkowsky s.n. (B); el- cies, C. echinatus, recently turned up in Andalusia). re- hammam, 60 km west of Alexandria, weeds of cultiva- cent records of "C. biflorus" from Morocco (Birouk & tion, 30.9.2001, L. Boulos 19528 (K); Assouan, berge al. 1991) are here identified as C. longi spinus. elsewhere empierrée du nil, en pente, près de l'embarcadère, in the Mediterranean area non-native species of Cenchrus 26.3.2004, J. Lambinon 04/Eg/50 (lG); Baltim, north- have been reported from several different countries but ern nile delta, waste ground, 14.9.1994, I. Mashaly & L. many records now turned out to be erroneous (for in- Boulos 20247 (K).
stance from croatia, Greece and israel).
France: Dep. Var, Îles d'hyères, s.d. [<1900], De-caisne s.n. (Br).
Israel: near Tel zur, Sharon plain, sandy clay soil, 29.11. Material and methods
1968, J. Mattatia s.n. (lG); lotan, 50 km n of elat, weed in an irrigated flower plot, 14.3.2011, A. Danin s.n. (Br); The results of this paper are entirely based on the revi- Kiriat Mozkin, Acco plain, 27.8.2011, M. Iehuda s.n. sion of material from selected herbaria: the herbarium of (Br); Kfar chabad, 20 km e of Tel Aviv, 29.8.2011, S. the Botanischer Garten und Botanisches Museum Berlin, Dadon s.n. (Br); Sharon, ramat-Aviv (Tel-Aviv), botani- Germany (B), the national Botanic Garden of Belgium cal garden, 30.8.2011, M. Ron s.n. (Br); Ketura, 60 km (Br), the Museo di Storia naturale in Firenze, italy (Fi), n of elat, Arava valley, 20.9.2011, A. Danin s.n. (Br); the university of Gent, Belgium (GenT), the royal Bo- Arad, har hanegev (negev highlands), 23.10.2011, N. tanic Gardens of Kew, england (K), the university of Dar Ben s.n. (Br); Ketura, 50 km n of elat, irrigated liège, Belgium (lG), the Museo civico di Storia natu- ornamental plot, 15.12.2011, A. Danin s.n. (Br).
rale in Milano, italy (MSnM) and the università degli Spain: Huelva: palos de la Frontera, nuevo puerto (uTM Studi of Torino, italy (To). in addition, material of the 29SpB8516), ruderal carretera, 9.9.2008, E. Sánchez private herbaria of the authors, as well as of J.-M. Tison Gullón 152 (herb. Sánchez, herb. Verloove); palos de and A. Soldano (Vercelli, italy) were also revised. la Frontera, nuevo puerto (uTM 29SpB8516), ruderal The studied area encompasses the entire Mediterra- aceras próxima a silos de cereal y grano, 20.9.2008, E. nean basin: Spain (incl. Balearic islands), France (incl. Sánchez Gullón 163 (Br, MGc 69266, SeV 228686); corse), italy (incl. Sardegna, Sicily), F.Y.r. (Former Yu- palos de la Frontera, nuevo puerto (uTM 29SpB8516), Willdenowia 42 – 2012 Fig. 1. Burs of the three non-native Cenchrus species in the Mediterranean area – A – B: C. echinatus, A from egypt, L. Boulos 19528 (K), B from Spain, E. Sánchez-Gullón 163 (Br); c: C. longispinus, from Morocco, J. Lewalle 11202 (Br); D–e: C. spinifex, D from Spain, F. Verloove 5523 (Br), e from italy, C. Ricceri & P. Debolini s.n. (Fi). – Scale bars = 1 mm.
cuneta carretera, 11.10.2010, E. Sánchez Gullón 311 the lowermost spines may be in part bristle-like but they (Br); huelva, ruderal cuneta carretera junto via del tren, are not as flexible and numerous as in C. echinatus and 15.9.2011, E. Sánchez Gullón 333 (Br).
they lack the typical retrorse spinules. C. echinatus is obviously most closely related to C. brownii roem. Notes. — Cenchrus echinatus is a troublesome weed and & Schult., another weedy new World species but less widely naturalised beyond its native distribution range but widespread than C. echinatus. Their separation is not al- is a relatively recent newcomer in the Mediterranean area, ways straightforward. however, the relatively larger burs where it was initially confined to its eastern part. Accord- (> 4.5 mm wide) that are rather loosely spaced (rachis ing to Delisle (1963) it was already spreading in israel in often visible) are typical of C. echinatus. in C. brownii, the early 1960s. Dafni & heller (1990) give 1970 as the in turn, the burs are much smaller (2 – 4.5 mm wide) and year of introduction for C. echinatus, while "C. incertus" very closely crowded (completely obscuring the rachis). (doubtlessly in error for C. longispinus, see below) would other distinguishing features (e.g. length of the outer have been present since 1953. This would be in contradic- bristles and colour of burs and spines) proved to be less tion with Delisle (1963) and is therefore rather unlikely. More recently C. echinatus was also recorded in egypt (cope 2005). The incipient naturalisation of C. echinatus Cenchrus longispinus (hack.) Fernald in rhodora 45:
in southwestern europe is here probably reported for the first time. in Spain (palos de la Frontera), it is more or less established in sandy, ruderal road verges in the vicin- Distribution. — native of united States and southern ity of a grain storage (recorded at least since 2008 and by canada, Mexico, central America and the West indies 2011 reported as locally spreading). its burs easily adhere (Delisle 1963). it is furthermore naturalised in Australia, to and are readily dispersed by man and mammals, which South Africa and the Mediterranean area. here reported could enhance a future wider naturalisation.
for the first time from the Former Yugoslavian republic Cenchrus echinatus is distinguished from most of its (croatia), Greece, israel and Morocco.
congenitors by the presence of a basal ring of numer-ous fine bristles that subtend the flattened spines. in C. Illustrations. — Fig. 2c; Grilli (1962: fig. 4, as Cenchrus longispinus (and to some extent in C. spinifex as well) pauciflorus); caro & Sánchez (1967b); Weston (1974: Verloove & Sánchez Gullón: non-native Cenchrus in the Mediterranean area fig. 1e); häfliger & Scholz (1980); Stieber & Wipff lignano pineta (13°04 – 05'e, 45°37 – 38'n), 30.8.1996, (2003); cope & Gray (2009). J. Stepánek & J. Stepáneková s.n. (B); Mün dung Taglia-mento, Dünen, 27.7.2006 (B). – Lazio: circeo, Baia Specimens examined. — Croatia: presso Dubrovnic, d'Argento, in località Molella, a 300 m del lago di Sabau- costa, 9.1966, Della-Beffa s.n. (To).
dia, terreno sabbioso, 1.11.1954, A. Cacciato s.n. (Fi); France: Haute-Corse: Alistro, sables maritimes à circeo (S. Felice), spiaggia, lungo la litoraneo per Ter- partir de la lisière des maquis jusqu'à la plage, très racina, copiosissimo,10.9.1968, A. Cacciato s.n. (Fi). – abondant, 22.9.1951, T. Marchioni s.n. (K); le pineto Marche: Tra pesaro e Fano, nella spiaggia, 8.1958, A. de Biguglia au sud de Bastia, langue de sable fermant Brilli-Cattarini s.n. (Fi, MSnM); Senigallia, dune verso l'étang de Biguglia, 11.8.1972, P. Sotiaux s.n. (Br); la Mazzochetta, 27.7.1987, A. Soldano 5987 (MSnM Alistro, plage [plante trouvée par T. Marchioni en août 24.117). – Pesara-Urbino: Solitarie in locis graminosis 1951 sur la plage d'Alistro, signalée par litardière sous et arenosis, ad domum deversorii Miramare in oppidu- le nom de Cenchrus tribuloides l. (candollea 14: 125, lo Marotta, 14.9.1995, F. Cernoch & J. Schubert 56.082 1953). l'année suivante le même auteur a corrigé sa dé- (lG). – Piemonte: Trino, ruderali presso il po, 8.10.1975, termination en Cenchrus incertus M. A. curtis (etudes A. Soldano 295 (herb. A. Soldano). – Sicily: prope Mes- corses nov. sér. 1: 41, 1954)], 22.7.1975, R. Descha- sanam, in pratis humidis et […] rivulas, 5.1871, M. Gan- tres s.n. (lG); linguizzetta, plage d'Alistro, 9.1986, doger s.n. (K). – Toscana: Forte dei Marmi (cin quale), J.-M. Tison s.n. (herb. J.-M. Tison); à l'eSe de Folelli, abbondante sulle dune costieri e negli incolti dell'interno, embouchure du Fium Alto, haut de plage rudéralisé entre s.d., F. Montacchini s.n. (To); Torre del lago, dune ma- l'embouchure et le petit village de vacances situé au sud, rine, 6.9.1970, P. V. Arrigoni s.n. (Fi); ronchi [di Massa], assez abondant, 24.8.1987, J. Lambinon 87/Co/670 (lG); are nile, 23.9.1974, A. Soldano 249 (herb. A. Soldano); Alistro, friche sur sable près de la mer [avec Conyza Torre del lago, sabbiosi lato strada per la superstrada, bonariensis, C. canadensis, C. sumatrensis, Portulaca 24.9.1980, A. Soldano (MSnM 24.118); poveromo (Mas- oleracea, Tribulus terrestris, Amaranthus albus, A. hy- sa), 8.1985, E. Banfi 28.054, 28.058, 28.061, 28.062, bridus, Chondrilla juncea, Chamaemelum mixtum, …], 28.064 (MSnM); Monte Argentario (Gr), Falde di pog- 9.9.1990, J. Lambinon 90/460 (B, Br, Fi, lG, MSnM, gio pertuso, nei pressi dello stabilimento balneare di also distributed by Soc. ech. pl. Vasc. eur. Bass. Médit. "Mamma licia", retroduna residua, 16.9.1994, R. Baldi- n° 16676; sub C. incertus); Dep. Vaucluse: Buisson, cul- ni (Fi). – Veneto: lido, Vene dig, 8.1952, Baschant s.n. tures sur terrain sableux, 20.9.1999, B. Girerd s.n. (lG); (B); Basso Veneto, [< 1961], P. Zanardini s.n. (Fi); Foci Buisson, quartier de la Buissonnière, 2.10.1999, B. Gir- del Sile, 14.9.1951, G. Moggi s.n. (Fi); Venezia, litorale erd s.n. (herb. J.-M. Tison).
del cavallino, beim leuchtturm der punta dei Sabbioni, Greece: Makedonien, nomos Kavalas, eparchia Sandboden, 19.7.1959, C. Simon s.n. (lG); Ve nice, lido nestou, Keramoti, Südrand des ortes gegenüber Thasos di Jasuzo (sic), 1964, I. J. Gibson 4 (K); Venezia, lido (40°51'20''n, 24°42'3''e), Sandstrand, 2 m, 27.9.2010, T. de Jésolo, extrémité orientale, dépression sableuse rudé- Raus & al. 32284 (B); südöstlich von Kera moti, nom. ralisée, 26.7.1973, P. Auquier 3197 (lG).
& ep., Xanthis, ne, fruchtend im Dünensand, Salsola Morocco: rabat (Souissi), terrain maraîcher, 1.7.1985, kali-Xanthium strumarium-Ass. oberdorfer & Tüxen, J. Lewalle 11202 (Br, lG); rabat, champ, 7.8.1988, J. 29.9.2010, I. Dinter 9831b (B).
Lewalle 12202 (Br, JAcA).
israel: Sharon, near or-Akiva, Menashe streams sedi-mentation, sand, 15.7.2011, D. Melamed s.n. (Br); Kfar Additional collections examined. — Hungary: prov. Masaric, Acco plain, 27.8.2011, M. Iehuda s.n. (Br); Kfar nógrád, Börz / prov. Bács-Kiskun, rand des Kiskunsági Masaryk, Acco plain, 29.8.2011, S. S. Cohen s.n. (Br).
nemzeti park, 2 km Se Fülöpháza (c. 25 km W Kecs- Italy: Abruzzo: Martinsicuro, 30.7.1986, A. Soldano kemét), Salzwiesen, 16.8.1994, D. Podlech 52202 (lG).
(herb. A. Soldano). – Campania: campolungo, destra orografica della Foce del Sele (Salerno), spiaggia calpes- Notes. — Cenchrus longispinus most closely resembles tata, 15.9.1992, L. Astolfi & R. Nazzaro s.n. (Fi, nAp; C. spinifex but confusion is also likely with C. echinatus. see Astolfi & nazzaro 1992 sub Cenchrus incertus). – From the latter it is best distinguished by the absence Emilia-Romagna: rimini, in arenosis maritimis, c. 3 m, of a basal ring of numerous flexible, retrorsely barbel- 10.8.1968, E. Mayer 73073 (B); Marina di ravenna, op late bristles (although some bristle-like spines may oc- het strand van de Adriatische zee, 9.7.1972, J.-E. De cur but these are never entirely and distinctly retrorsely Langhe 357/72 (Br); Marina di ravenna, op het strand, barbellate). Moreover its spines emerge at irregular in- 9.7.1972, R. D'hose 357/72 (Br). – Friuli-Venezia- tervals throughout the body of the bur (in C. echinatus Giulia: lignano Sabbiadoro, sabbiosi presso la foce the spines of equal size originate at more or less the same del Tagliamento, 16.7.1983, A. Soldano 5633 (herb. level). however, most problematic is the distinction of A. Soldano); sine loco, 1984, H. Melzer s.n. (B); urbs C. longispinus and C. spinifex. Both have been largely udine, mare hadriaticum, opp. lignano, in locis areno- intermixed so far in the studied area. C. longispinus al- sis in litore maris ad ostium flum. Tagliamento ad vicum ways has more spines, the inner being terete to slightly Willdenowia 42 – 2012 flattened and the outer (lowermost) often bristle-like and study but all proved to be ascribable to C. longispinus. relatively slender. in C. spinifex, on the contrary, spines Moreover, pictures of "C. incertus" from other croatian are always fewer, the inner distinctly flattened (up to localities surely also pertain to C. longispinus (see for 3 mm wide at base) and bristle-like outer spines are near- instance: island rab, lopar, 2007, J. nejc, record no. ly always lacking. photographs of burs (Fig. 1, compare 30847 in Flora croatica Database 2004+). The records c and D–e) clearly show the differences between both from Greece here presented were previously also as- species, much more than words can do.
cribed to C. incertus (raus & Schuler 2005).
Cenchrus longispinus is by far the most misunder- in israel, two non-native species of Cenchrus are stood non-native species of the genus in the Mediterra- known (Dafni & heller 1990): C. echinatus and "C. in- nean area and, in fact, appears to be the most widespread. certus" but both have been widely confused. The latter its oldest centre of naturalisation in europe probably is apparently confined to the northern half of the country is on the Adriatic coast in italy: it is known since at least and is, at least for the time being, a non-invasive weed 1933 from lido del cavallino in Venezia province (cor- (Danin 2004). All records seen from this area are here betta 1964) and now is widely naturalised in this area. corrected to C. longispinus as well. Moreover, additional its distribution and invasive status elsewhere in italy is photographs of "C. incertus" from israel (see for instance uncertain, largely as a result of lingering confusion with Gold & eshel 2012) also belong with C. longispinus.
Cenchrus spinifex. According to Guzik & pacyna (1999) in Morocco non-native Cenchrus must be either a plants naturalised in italy, known as "C. incertus", repre- fairly recent introduction (compare with le Floc'h & al. sent in fact C. longispinus. however, genuine C. spinifex 1990, Valdés & al. 2002) or it must have been overlooked also exists in italy (see below). part of the confusion sure- for quite some time. Dobignard & chatelain (2010) only ly is induced by the fact that in many italian regions both cite C. biflorus and C. ciliaris for Morocco but the pres- species occur sympatrically: in all(!) regions where C. ence of the former is questioned (see also ibn Tattou & spinifex is confirmed in the present study, also C. longi- Fennane 2008). indeed, the only collections seen of C. spinus occurs. in some places, for instance in poveromo biflorus (all from the surroundings of rabat and dating in Tuscany, both grow nearly side by side. celesti-Grapow back to the 1980s) pertain to C. longispinus. According to & al. (2010) correctly accepted both C. longispinus and A. Dobignard (pers. comm. 2011) reports of C. biflorus in C. spinifex (as C. incertus) for italy but their distribution Morocco are referable to J. lewalle but collections of the and degree of naturalisation are obviously wrongly as- latter all proved to belong to C. longispinus.
sessed, giving C. longi spinus only for Veneto (casual) and in neighbouring regions, outside the studied area, Friuli-Venezia-Giulia (naturalised). The present study, in identical identification problems have occurred in the contrast, confirms its presence also in Abruzzo, campa- past: plants from hungary (voucher in lG!) and ukraine nia, emilia-romagna, lazio, Marche, piemonte, Sicily (Guzik & pacyna 1999), for instance, are ascribable to and Tuscany. its status of occurrence should be critically Cenchrus longispinus, not to C. spinifex. in hungary C. assessed but C. longispinus is probably naturalised (or in- longispinus (as C. incertus) is considered to be an inva- vasive) in all regions except piemonte and Sicily.
sive xenophyte in disturbed open grassland in the Great in France, Cenchrus longispinus has been recorded hungarian plain (Szigetvári 2002). The same applies to since 1951, at first in corse. All voucher specimens seen ukraine where it is considered among the worst invasive from that area (see above), indeed, belong to C. long- species (Mosyakin 2006).
ispinus (Verloove 2006). They were initially wrongly re-ferred to as C. tribuloides and subsequently as C. incertus Cenchrus spinifex cav., icon. 5: 38, t. 461. 1799
(see for instance Deschâtres 1986, natali & Jeanmonod = Cenchrus incertus M. A. curtis in Boston J. nat. hist. 1996, Vagnet & Vadam 2005). At present, C. longispinus 1: 135. 1837.
is commonly naturalised on the eastern coast of corse, = Cenchrus carolinianus Walter, Fl. carol.: 79. 1788, at least between Bastia and Alistru (Jeanmonod & Gam- nom. rejic. (see Brummitt 1995: 608).
isans 2007). More recently, a locally naturalised popu- = Cenchrus pauciflorus Benth., Bot. Voy. Sulphur: 56. lation from continental France (Buisson; dep. Vaucluse) published as C. incertus (Girerd & roux 2000), also rep- ?= Cenchrus bambusoides caro & e. A. Sánchez in resents C. longispinus. in this locality it was still con- Kurtziana 4: 44. 1967.
firmed in 2010 by christophe Girod (pers. comm. J.-M. Tison) but environmental and climatological conditions Distribution. — native of southern united States, Mexi- seem to be less favourable than elsewhere in the Mediter- co, central and South America and the West indies. More ranean area and a future, wider naturalisation in conti- or less widely naturalised as a noxious weed, for instance nental France appears to be rather unlikely.
in S Africa, china, Australia, the Mediterranean area, etc.
in croatia and Greece "Cenchrus incertus" was re- cently reported as an invasive alien plant species (Borsi6 Illustrations. — Fig. 1D–e; Vivant (1961 as Cenchrus & al. 2008; Arianoutsou & al. 2010). relatively few col- pauciflorus); Grilli (1962: fig. 3 as C. pauciflorus); caro lections of non-native Cenchrus were seen for the present & Sánchez (1967b as C. incertus and C. pauciflorus); Verloove & Sánchez Gullón: non-native Cenchrus in the Mediterranean area Weston (1974: fig. 1B); häfliger & Scholz (1980 as C. The first naturalised populations of Cenchrus spinifex incertus and C. pauciflorus); Sanz elorza & al. (2004 as in europe were discovered in italy near Viareggio in 1939 C. incertus); cope & Gray (2009 as C. incertus).
(plicker 1943, as C. tribuloides). in this part of Tuscany it is still present and more or less widely naturalised in Specimens examined. — France: Pyrénées-Atlan- ruderalised coastal dunes, for instance in Marina di Massa tiques: Anglet, Blancpignon, adventice dans les clair- and Marina di carrara. in Veneto, C. spinifex seems to ières sablonneuses de la pinède, aux abords de l'eglise be well-established in the surroundings of Mesola, while des Sables, du cimetière et de l'Allée de l'empereur sur local occurrences are here confirmed from Abruzzo and une distance de plus d'un km, 24.8. et 1.10.1960, J. Jal- emilia-romagna. elsewhere in italy C. spinifex has lu 7149 (Br, lG; also distributed by Soc. Fr. ech. pl. been widely confused up to present with C. longispinus. Vasc. n° 3688); Anglet, Blancpignon, près de l'église celesti-Grapow & al. (2010) also cite the former (as C. St. Joseph, 19.7.1975, N. Cnops 75.129 (Br); Anglet, à incertus) from campania, lazio, Marche, Molise, puglia la chambre d'Amour, arrière-plage rudéralisée, [plante and Valle d'Aosta but these records are either erroneous pérennante, épines fortes et peu nombreuses], 7.2005, (and ascribable to C. longispinus, see above) or require J. M. Tison s.n. (herb. J.-M. Tison; photocopy herb. confirmation. in italy C. spinifex exclusively occurs in Verloove); id., Bayonne, Anglet (chambre d'Amour), regions where C. longispinus also is present. This surely arrière-plage rudéralisée, très commun, 9.9.2006, F. Ver- added to the confusion between both species.
loove 6480 (herb. Verloove).
in southwestern France Cenchrus spinifex was re- Italy: Abruzzo: presso la foce del Borsacchio, si è dif- corded in the surroundings of Anglet near Bayonne from fusa più presso l'abitato di roseto, 10.1950, G. Zodda s.n. 1960 onwards (Vivant 1961, as C. pauciflorus). it is still (Fi); roseto degli Abruzzi, sabbiosi, 30.7.1986, A. Sol- (very) locally abundant in this area (for instance near dano 5390 (herb. A. Soldano). – Emilia-Romag na: chambre d'Amour, see above) but many of its original Bei comacchio, lido Dogli Stucchi, Dünengelände, localities probably disappeared (pers. comm. J. Vivant). 21.7.1969, H. Scholz s.n. (B). – Toscana: Marina di Mas- To our knowledge, C. spinifex is absent from other parts sa, fra la colonia edison e la foce del Brugiano, nel tratto of France: all populations from corse belong to C. longi- di littorale arenoso, in numerosi esemplari, 23.6.1947, P. spinus (Verloove 2006) and the same holds true for plants Pellegrini s.n. (Fi); Marina di Massa, poveromo, dune from the Vaucluse (see above). li toranee [con Echinophora spinosa, Solidago litora- in northeastern Spain the species was discovered in lis, Stachys maritima, Pycnocomon rutifolium, etc., Torre de la Mora, near Tarragona, for the first time in 16.7.1978, E. Banfi s.n. (lG); Marina di Massa, povero- 1972, correctly referred to as Cenchrus incertus by Tor- mo, sabie marittime, 20.8.1985, E. Banfi s.n. (lG); pov- rella & al. (1974). in this area it has become a widespread eromo (Massa), 8.1985, E. Banfi 28.063 (MSnM); pov- and noxious weed in coastal dunes, at least between eromo di Massa (MS), spiaggia del Tornado, 2.9.1992, cambrils and Baix llobregat (see map in Sanz elorza E. Ferrarini s.n. (Fi); Marina di Massa, arid grassland & al. 2004 and BDBc 2011). C. spinifex is recently near the sea, very common (perennial), 22.6.2006, F. Ver- also naturalising in Garrotxa (oliver 2009). Since 1996 loove 6361 (Br, ro; herb. Verloove); Marina di carrara, it is known from few localities in país Vasco (campos partaccia, sandy grassland, dunes, locally, 22.6.2006, F. & herrera 2008), relatively close to the localities in the Verloove 6632 (Br; herb. Verloove). – Veneto: Mes- French pays Basque (see above). in 2002, C. spinifex (as ola, bosco della Mesola, sabbie del litorale, notato an- C. incertus) was recorded for the first time in Andalusia che all'interno, 22.7.1964, P. Stampi s.n. (Fi); Mesola, (San Juan del puerto, near huelva; Sánchez Gullón & al. bosco della Mesola, strada adiacenti al canale elciola, 2006) but did not establish. Soon afterwards, in 2007, 22.7.1964, P. Stampi s.n. (Fi); Mesola, bosco della C. spinifex was observed in abundance (and obviously Mesola, al Taglio della Falce, 14.8.1964, P. Stampi s.n. naturalised) in a worked-up, sandy roadside by a military (Fi); Mesola, bosco della Mesola, 7.1965, P. Stampi s.n. base in el puerto Santa Maria, close to cádiz (Verloove (Fi); Mesola, bosco della Mesola, Taglio della Falce, & Sánchez Gullón 2008). in Spain C. spinifex is now 24.7.1975, C. Ricceri & P. Debolini s.n. (Fi).
considered to be an invasive species on sandy beaches, Spain: Cadíz: el puerto Santa Maria (Fuentebravia), especially in the northeastern part of the country (Sanz sandy road verge at military base, 8.10.2007, F. Verloove elorza & al. 2004).
6993 (B, Br, lG). – Huelva: huelva, ruderal en oril- reports of Cenchrus spinifex from other countries in las carretera n-442, 4.10.2002, E. Sánchez Gullón 91 the Mediterranean (for instance Greece, israel, libya, (Br). – Tarragona: cambrils (Vilafortuny), sandy rud- Serbia and Turkey; see Valdés & Scholz 2009+) should eral road verge, one tall specimen, 27.9.2003, F. Verloove be critically reviewed. At least part of the reports is as- 5523 (herb. Verloove).
cribable to C. longispinus.
Cenchrus spinifex is usually considered to be con- Notes. — Distinguishing features between this species specific with C. pauciflorus nowadays (Delisle 1963; and Cenchrus longispinus are discussed under the latter. Stieber & Wipff 2003; zuloaga & al. 2003) but Ward confusion with C. echinatus is unlikely.
(2010) recently rejected this synonymy. According to Willdenowia 42 – 2012 him both are best distinguished on life form (respec- presented here. This key is mostly based on the results tively annual and perennial) but it is doubtful if this war- of our examination of numerous herbarium collections rants specific rank. Duration seems to be variable in C. and considerably differs from that of Delisle (1963) spinifex in the studied area and therefore often critical who probably gave too much weight on the degree of to assess: some plants are reported to be annuals (see fusion of the spines. C. setigerus, for instance, was ac- for instance Sanz elorza & al. 2004) while others are commodated in a dichotomy with "spines connate only obviously (short-lived) perennials. Ward (2010) admits at the base", while they are connate usually for at least that where C. incertus and C. pauciflorus meet, fertile half their length. Moreover, even in species having burs hybrids are produced. Twentyman (1972) experimen- with distinctly fused inner bristles, this holds true only tally showed that culm length, habit and the ability for for one side of the bur, while on the other side the spines overwintering merely depends on day-length and envi- are always nearly free to the base, largely exposing the ronmental conditions. This issue possibly requires ad- ditional research in the Mediterranean area but, at least For convenience, Cenchrus ciliaris, a species often for the time being, all of these plants are best referred to transferred to Pennisetum recently, and its look-alike C. as C. spinifex (incl. C. pauciflorus).
pennisetiformis are also included in the key.
likewise, Cenchrus bambusoides is sometimes ac- Mature burs are absolutely required for a reliable de- cepted as a good species (caro & Sánchez 1967a; Ward 2010). in general habit it looks like C. spinifex but its leaves are inrolled on drying, without an obvious keel (in- 1. Burs with numerous outer flexible bristles, these dis- stead of flat or folded). it probably merely belongs to the tinctly antrorsely barbellate; all other bristles also flex- variability of the latter (Stieber & Wipff 2003; zuloaga & ible and therefore burs never prickly at maturity . 2 al. 2003). Moreover, plants with foliar characteristics of – Burs with or without outer flexible bristles, these (if C. bambusoides apparently have not been recorded so far present) either not at all or hardly barbellate or (pre- in the studied area.
dominantly) retrorsely barbellate; all (other) bristles The application of the binomial Cenchrus spinifex is stout, stiff and/or flattened, burs therefore always not uncontested. in addition to the aforementioned taxo- prickly at maturity (often only faintly so in C. seti- nomic difficulties there is still a nomenclatural problem. Delisle (1963) already evoked that the correct name for 2. inner bristles 15 – 27 mm long, all more or less equal C. incertus might be C. spinifex, which, indeed, antedates in length; plant annual . . . . . . . C. prieurii the former (1837 versus 1799). As he had not been able – inner bristles 7 – 14 mm long, one distinctly longer and to study the holotype of C. spinifex and the correctness wider than the others; plant perennial or annual . 3 of the isotype label had not been verified, he rejected 3. caespitose perennial, ultimately with hard, knotty this binomial. in the recent treatment of Cenchrus for base; inner bristles connate only at their extreme bas- north America (Stieber & Wipff 2003) the plants here es, forming a disc c. 0.5 – 1 mm in diameter . . .
concerned are referred to as C. spinifex but it is unclear whether or not these authors had effectively resolved the – Annual to pauciennial; inner bristles connate to form nomenclatural problem. Moreover, C. incertus is not even a disc c. 1.5 – 3 mm in diameter . C. pennisetiformis mentioned as a synonym by these authors. in the recent 4. inner bristles only fused at the base, each with 1 – 3 catalogue of new World grasses (zuloaga & al. 2003), C. distinct grooves on the outer face (hence the back dis- incertus is upheld and it is stated that "… if C. spinifex tinctly veined) . . . . . . . . . . C. biflorus turns out to be conspecific, a conservation proposal will – inner bristles fused for at least 1/3 of their length, with be considered". According to Ward (2010), C. spinifex is or without grooves on the outer surface . . . . 5 an unidentifiable name that should be rejected. Symon 5. Burs cup-shaped, with inner bristles short and broad (2010), who examined a digital image of the holotype, on (2 – 4 mm long); outer bristles always fewer in number the contrary confirms that both are conspecific and hence and mostly lacking; inner bristles fused for c. 1/3 – 1/2 C. spinifex should finally be accepted as the correct name of their length . . . . . . . . . . C. setigerus for this species. Joseph Wipff recently also investigated – Burs ovoid to globose, with inner bristles usually the type of the latter name and states that it definitely longer; outer bristles sparse or numerous, rarely ab- refers to the taxon traditionally called C. incertus (pers. sent; inner bristles mostly fused for at least 1/2 of their comm. March 2012).
6. Burs with numerous flexible, distinctly retrorsely bar- bellate outer bristles; inner bristles originating almost Key to the species of Cenchrus s.str. in the Mediter-
in a single whorl and forming flattened spines, more ranean area
or less erect at maturity . . . . . . C. echinatus
in an attempt to avoid future misidentifications within – Burs without flexible, retrorsely barbellate outer bris- Cenchrus s.str. (excl. Pennisetum; see Verloove 2012) tles [in some burs a few bristle-like spines may be in the Mediterranean area, a revised identification key is present that are not thin and flexible as in C. echinatus Verloove & Sánchez Gullón: non-native Cenchrus in the Mediterranean area and that under high magnification appear slightly bar- caro J. A. & Sánchez e. A. 1967a: notas críticas sobre es- bellate with a mixture of retrorse and antrorse barbs pecies de Cenchrus (Gramineae). – Kurtziana 4: 39 – 50.
but not with exclusively retrorse barbs as in C. echi- caro J. A. & Sánchez e. A. 1967b: las especies de natus]; inner flattened spines originating at irregular Cenchrus (Gramineae) de la república Argentina. – intervals throughout the body of the bur, distinctly Kurtziana 4: 95 – 129.
cecconelli e. 1975: Cenchrus longispinus Fernald sull' 7. Spines relatively long and numerous (usually alto litorale adriatico occidentale. – Delect. Semin. c. 30 – 50), slenderly pointed; most of the outer spines Sporar. orto Bot. ist. Tecn. Geometri G. G. Marinoni very slender (bristle-like) and ranging from patent to udine 26: 3 – 13.
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