Amaranthus powellii subsp. cacciatoi comb. et stat. nov. (Amaranthaceae)

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1 Nordic Journal of Botany 30: 12 16, 2012 doi: /j x, 2012 The Author. Nordic Journal of Botany 2012 Nordic Society Oikos Subject Editor: Arne Strid. Accepted 10 October 2011 Amaranthus powellii subsp. cacciatoi comb. et stat. nov. (Amaranthaceae) Duilio Iamonico D. Iamonico Via dei Colli Albani, IT Roma, Italy. Amaranthus powellii S. Watson subsp. cacciatoi (Aellen ex Cacciato) Iamonico comb. et stat. nov. (Amaranthaceae) is proposed based on morphometric and taxonomic studies on the type material and other specimens. A detailed description is provided and the new taxon is compared with A. powellii S. Watson subsp. powellii and subsp. bouchonii (Thell.) Costea & Carretero. Amaranthus L. is a genus of about 70 species, of which about 40 are native to the Americas, with the remaining native to Australia, Africa, Asia and Europe (Costea et al. 2001). This genus is taxonomically problematic due to nomenclatural confusion resulting from frequent misapplication of names. Recent work on the European alien flora (Lambdon et al. 2008) has also found Amaranthus to be a difficult genus with largely subjective species delimitation. Most of these problems are concentrated in the A. hybridus group (according to Costea et al. 2001). During my studies on Amaranthus in Italy, I found a taxon described by P. Aellen in 1965 ( A. bouchonii Th ell. var. cacciatoi ), that was published from Italy by Cacciato (1966), but has subsequently been ignored in more recent floras (Fig. 1). Among Italian floras, var. cacciatoi was mentioned only by Cacciato (1982), who recorded it from Rome (Latium, central Italy) and by Anzalone (1996), again from the Italian capital based on research by Cacciato (1966). The recent checklist of the Italian vascular flora (Conti et al. 2005) recorded A. bouchonii from several Italian regions, though without infraspecific taxa. In Conti et al. (2007) and Celesti-Grapow et al. (2009) A. bouchonii was synonymized within A. powellii S. Watson, while, recently, Iamonico (2008) accepted the name A. powellii S. Watson subsp. bouchonii (Thell.) Costea & Carretero for the Italian flora following Greizerstein et al. (1997) and Costea et al. (2001). In floras where A. bouchonii has been accepted, the presence of var. cacciatoi has never been indicated (e.g. Aellen 1973, Greuter et al. 1984, H ü gin 1987, Akeroyd 1993, Raus 1997, Stace 1997, Wisskirchen and Haeupler 1998, Kergu é len 1999, Wohlgemuth et al , Haeupler and Muer 2000, Costea et al. 2001, 2004, Jonsell 2001, Mosyakin and Robertson 2003). Nor is var. cacciatoi listed in the main online meta-sources for plant taxonomy (International Plant Names Index 2006, International Organization for Plant Information 2006, Tropicos 2009). The purpose of this study is to define the taxon cacciatoi and to determine if it is worthy of recognition within A. bouchonii or as it has been redefined as part of A. powellii. Material and methods Specimens of A. powellii subsp. bouchonii, A. powellii subsp. powellii and taxon cacciatoi from BOZ, G, FI, MRSN, MSNM, RO, ROV, TSB and the personal herbaria of Dr C. Argenti, Dr A. Soldano and Dr A. Tisi (not listed in Index Herbariorum) were examined. The following characteristics were measured from 200 specimens (using the average measurements of 5 flowers, fruits and seeds from each exsiccatum ), for a total of about measurements: 1) plant height (m), 2) stem vestiture (glabrous or pubescent in inflorescence region) *, 3) terminal spike length (mm), 4) inflorescence structure (lax and not erect, with many lateral branches or stiff and erect, unbranched or with very few widely spaced branches) *, 5) bracteole length (mm), 6) bracteole base maximum width (mm), 7) tepals length (mm), 8) tepal width (mm), 9) flower symmetry (actinomorphic or zygomorphic) *, 10) fruit length (mm), 11) fruit width (mm), 12) fruit dehiscence/or indehiscence *, 13) seed length (mm), 14) seed width (mm), 15) seed surface color (black or brown) *, 16) seed surface sculpturing (smooth or minutely punctiform sculputured) *, 17) fruit length/width ratio, 18) bract length/tepal length ratio, 19) seed length/ width ratio. Most of the characters are quantitative but those marked with an asterisk ( * ) were coded as binary. The variables were processed with the software of statistical and multivariate analysis NCSS Cluster analysis 12

2 Th e highest contribution to the first principal component (axis 1) is provided by the ratio of bract length/tepal length, the inflorescence structure and the fruit dehiscence vs indehiscence. The contribution of these characters to the other three main components (axis 2, 3, 4) are low. The second component (Fig. 3a, 3d, 3e) does not clearly separate any group (the largest contribution made by the tepal length and tepal width). The third (Fig. 3b, 3d, 3f) and the fourth (Fig. 3c, 3e, 3f), on the other hand, show a wellseparated group where the specimens from Rome (taxon cacciatoi ) can be recognized, while specimens refered to taxa powellii and bouchonii are not separated. The highest contributions to these latter components (axis 3, 4) were made by the flower (symmetry). With regards to distribution, the taxa show no geographic separation and, in particular, taxon cacciatoi is sympatric with taxon powellii in central Italy (see Fig. 4 for the distribution of A. powellii s.l. in Italy, according to Iamonico 2008, 2009b, Iamonico and Del Guacchio 2010). However, the dispersal of the fruits and seeds of Amaranthus species is often performed by humans, mainly by sewage works (Costea et al. 2004, Iamonico 2009a, 2010), making the natural distributions of these taxa obscure. Although the ordination studies (Fig. 3d f) show that the taxon cacciatoi is more distinct than the other two taxa (and may consequently be considered a distinct species), there is an overlap of some measurements (Table 1). On the basis of this and the discussion about the distribution (above), I think that the most appropriate taxonomic rank for taxon cacciatoi is that of subspecies. Figure 1. Holotype of Amaranthus bouchonii var. cacciatoi Aellen ex Cacciato (G). was performed by the UPGMA method. The data matrix was also subjected to ordination (principal component analysis, PCA). Results and discussion The analysis of the morphological features and qualitative characters (Table 1) allows the recognition of distinct sub-units recognizable as taxa. The dendrogram (Fig. 2) shows two main clusters (1 and 2). The first one (1) is divided into two sub-clusters. Sub-cluster 1.1 is entirely composed of specimens of var. cacciatoi from the city of Rome. Subcluster 1.2 is comprised of A. powellii subsp. bouchonii specimens (from northern Italy) and it is further divided into two sub-clusters (1.2a and 1.2b); each including a large proportion of specimens from several localities. Cluster 2 is composed of A. powellii subsp. powellii specimens. The results of the cluster analysis are confirmed by the PCA results (Fig. 3); the cumulative percentage of eigenvalues for the first four axes is about 66.5%. The PCA (Fig. 3a c) shows three clearly separated groups (corresponding to sub-clusters 1.1, 1.2 and cluster 2 in Fig. 2). The sub-clusters 1.2a and 1.2b (as defined in Fig. 2) are not discernable in the PCA ordination. Amaranthus powellii S. Watson subsp. cacciatoi (Aellen ex Cacciato) Iamonico comb. et stat. nov. (Fig. 1) Basionym : Amaranthus bouchonii var. cacciatoi Aellen ex Cacciato, Ann. Bot. Roma 28 (1966, p. 618). Type : Italia, Lazio, Roma, in ruderatis, communis in ipse Urbe. 12 Aug 1965 A. Cacciato 98 (holotype: G!). Annual therophyte. Stems erect, m tall, sparsely pubescent (in some specimens more densely pubescent on the inflorescence axis). Leaves subovate or rhombic, cm. Inflorescence erect, with many lateral branches and a terminal spike 2 9 cm long, subequal or longer than the lateral spikes. Bracteoles rigid, narrowing towards the apex, mm long, about times longer than the tepals. Tepals 5, equal or slightly unequal, mm long, lanceolate to linear, with lateral margins parallel. Fruit dehiscent, ovoid, mm long, about times longer than wide, the pericarp smooth. Seeds usually broadly ovate, mm, the surface smooth, shiny, black or brown. Etymology The name was chosen by P. Aellen (Basel) in honour of the Italian botanist Dr Alfredo Cacciato who was an expert on the genus Amaranthus in Italy in the 1960 s and 1970 s. 13

3 Table 1. Morphological features of Amaranthus cacciatoi, A. bouchoni i and A. powellii. A. cacciatoi A. bouchonii A. powellii Plant height (m) Stem indumentum Slightly pubescent in the inflorescence region Glabrous or slightly pubescent in the inflorescence region Glabrous or slightly pubescent in the inflorescence region Inflorescence structure Lax and not erect, with many lateral branches Lax and not erect, with many lateral branches Stiff and erect, unbranched or with very few widely spaced branched Terminal spike length (cm) (12.0) (2.0) Bracteole length (mm) Bracteole base width (mm) Tepal length (mm) Tepal width (mm) Flower symmetry Tepals usually equal, with parallel margins Tepals usually very unequal, with or without parallel margins Tepals usually very unequal, with or without parallel margins Fruit length (mm) Fruit width (mm) Fruit Dehiscent Indehiscent Dehiscent Seed length diameter (mm) Seed width diameter (mm) Seed colour Black or brown Black or brown Black or brown Seed surface Smooth Smooth Smooth Ratio fruit length to fruit width Ratio bract length to tepal length (3.5) Ratio longer diameter to shorter diameter of seed Habitat and distribution Amaranthus powellii subsp. cacciatoi has been found in the south southeast metropolitan area of Rome (Lazio, central Italy), in ruderal habitats, at altitudes above m a.s.l. Flowering has been observed between Jun Nov. It is widely accepted that Amaranthus powellii s.s. has been introduced to Europe from North and South America (Akeroyd 1993, Costea et al. 2001, Mosyakin and Robertson 2003), presumably by human activities, as for several other Amaranthus species (Costea et al. 2004, Iamonico 2009a, 2010). However, Amaranthus powellii subsp. bouchonii has been described from Europe (southwest France, Thellung 1926) and it is considered to be of European origin, although some authors are uncertain of its origin (Costea et al. 2001). With regards to A. powellii subsp. cacciatoi, it is probable that it has been introduced from the New World and has evolved independently in Europe, or that it is of hybrid origin, or the product of introgression with other species. Up to now subsp. cacciatoi has only been recorded in the Lazio region and more studies will have to determine if it is endemic to this area or not. Figure 2. UPGMA dendrogram based on 19 morphological characters (see Material and Methods and Table 1) showing the phenetic relationships among the examined exsiccata. Differences between the subspecies Compared to Amaranthus powellii subsp. bouchonii, the fruit of subsp. cacciatoi is always circumscissily dehiscent rather than indehiscent. In addition, subsp. cacciatoi has actinomorphic or slightly zygomorphic flowers (with the tepals equal or slightly unequal, with lateral parallel margins), while subsp. bouchonii has clearly zygomorphic flowers (with the tepals very unequal and their margins either parallel or not). The terminal spike of the inflorescence is longer than (in some cases subequal to) the lateral spikes in both subspecies, however in subsp. cacciatoi the terminal spikes are 2 9 cm long, while in subsp. bouchonii they are usually longer (up to 20 cm). The morphological differences between subsp. cacciatoi and subsp. powellii are in the flowers (actinomorphic or slightly zygomorphic in subsp. cacciatoi, clearly zygomorphic in subsp. powellii ) and in the inflorescence (lax and not erect, with many lateral branches in subsp. cacciatoi ; stiff and erect, unbranched or with very few widely spaced branches in subsp. powellii ). 14

4 Figure 3. Ordination (PCA) based on 19 morphological characters (see Material and Methods and Table 1). The cumulative percentage of eigenvalues for the first four axes is over 66.5% (cacc taxon cacciatoi, bouch taxon bouchonii, pow taxon powellii ). A key to subspecies of Amaranthus powellii s.l. follows: 1. Inflorescence stiff and erect, unbranched or with very few widely spaced branches; bract/tepal length ratio A. powellii subsp. powellii Inflorescence lax and not erect, with many lateral branches; bract/tepal length ratio Fruit circumscisly dehiscent; terminal spikes 2 9 cm long. Flowers actinomorphic or slightly zygomorphic (tepals equal or subequal, with parallel margins)... A. powellii subsp. cacciatoi Fruit indehiscent; terminal spikes usually longer (up to 20 cm). Flowers zygomorphic (tepals very unequal and their margins parallel or not).... A. powellii subsp. bouchonii Amaranthus powellii subsp. bouchonii Italy. Friuli-Venezia Giulia: Provincia di Udine, Sacile: San Giovanni di Livenza, 4 Sep 1998, L. Poldini 0041/3 (TSB); Lombardia: Milano, zona 18 Baggio, via Albona: numero dispari, tra via A. da Gandino e via Cabella, marciapiede presso il cordolo che separa l aiuola, 120 m a.s.l., Specimens examined Amaranthus powellii subsp. cacciatoi Italy. Lazio: Roma, Piazza R. Malatesta 4 Nov 1960, A. Cacciato (RO); Roma, Centocelle, 29 Aug 1961, A. Cacciato s. n. (RO); Roma, 20 Aug 1962, A. Cacciato s. n. (RO). Roma, Cinecitt à, 14 Jun 1963, A. Cacciato s. n. (RO); Roma, ruderati a Torpignattara, 14 Aug 1964, A. Cacciato (FI); Roma, in ruderatis, communis in ipse Urbe, 12 Aug 1965, A. Cacciato 98 (holotype: G); Roma, prati a Cinecitt à, 10 Aug 1965, A. Cacciato s. n. (RO); Roma, 28 Aug 1966, A. Cacciato 13 (G); Roma, 4 Oct 1968, A. Cacciato s. n. (RO); Roma, lungo via di Portonaccio, 20 Aug 1968, A. Cacciato s. n. (RO); Roma, 14 Jul 1970, A. Cacciato s. n. (RO); Roma, via Appia Nuova, all altezza di via delle Capannelle, incolti al margine della strada, 15 Sep 2008, D. Iamonico s. n. (herb. Iamonico, RO); Roma, Parco degli Acquedotti (zona Tuscolana-Cinecitt à ), incolti, 17 Aug 2009, D. Iamonico s. n. (herb. Iamonico, RO). Figure 4. Distribution map showing provinces in Italy with records of Amaranthus powellii subsp. cacciatoi ( ), A. powellii subsp. bouchonii ( ) and A. powellii subsp. powellii ( ). 15

5 8 Nov 1991, G. Galasso s. n. (MSNM); Piemonte: Vercelli, lato sponda destra del Sesia a monte del ponte ferroviario, 130 m a.s.l., 15 Nov 1979, A. Soldano 2482 (herb. Soldano); Cureggio (NO), in luogo incolto presso Fontaneto d Aragona, 30 Sep 1983, G. Abb à 89 (MRSN); Alessandria, Fugarolo, pianura, 94 m a.s.l., 25 Aug 2008, A. Tisi s. n. (herb. Tisi); Toscana: Massa, cinque vie, 5 Nov 1975, A. Soldano s. n. (herb. Soldano); Trentino-Alto Adige: Provincia di Trento, loc. Colle a circa 500 m a N di Mori, in campo di patate, 605 m a.s.l., 15 Aug 1999, F. Prosser s. n. (ROV); Bolzano, Siebeneich (Terlan), Margarethenwald 0.4 km NW Darumhof, ruderalstelle, 270 m a.s.l., 8 Oct 2004, W. Stockner PVASC5127 (BOZ). Amaranthus powellii subsp. powellii Italy. Abruzzo: L Aquila, lago di Campotosto, 1314 m a.s.l., 8 Nov 2002, Leg. D. Tinti, Det. D. Iamonico 2167 (APP); Lazio: Roma, Parco Urbano di Aguzzano, Terreno da riporto, 100 m a.s.l., 8 Nov 2007, D. Iamonico s. n. (RO, herb. Iamonico); Roma, Valmontone, incolto al margine della Strada Vicinale della Vecchia, nei pressi della stazione ferroviaria, 845 m a.s.l., 15 Oct 2008, D. Iamonico s. n. (herb. Iamonico); Friuli-Venezia Giulia: Udine, Villa Santina, 2 Nov 2001, L. Poldini s. n. (TSB); Trentino- Alo Adige: Bozen, Feldthurns, Schrambach, Villn ö sser Haltestelle, Rand der Bahnstrecke, 530 m a.s.l., 22 Aug 2002, A. Hilpold PVASC3631 (BOZ); Trento, 3 Sep 1890, Leg. E. Gelmi, Det. F. Festi s. n. (TR); Trento, Piano di Vallarsa: campo lungo la stradina per Poiani, Campo a riposo, 890 m a.s.l., 25 Oct 2005, F. Prosser s. n. (ROV); Veneto: Padova, Tra Faedo e Fontanafredda in una cascina (Colli Euganei, provincia di Padova, Italia), Su macerie, 100 m a.s.l., 7 Aug 1989, F. Prosser s. n. (ROV); Verona, Navene, lungolago a N delle Terme, Scarpata ghiaiosa, 67 m, 9 Nov 2006, Leg. A. Bertolli, F. Prosser, Det. F. Prosser s. n. (ROV). Acknowledgements I am grateful to Dr L. Peruzzi (Univ. of Pisa), Dr M. Iberite (Univ. of Rome Sapienza) and Dr M. Costea (Wilfrid Laurier Univ., Canada) for constructive comments on the first version of the manuscript. Thanks to directors and curators of BOZ, G, FI, MRSN, MSNM, RO, ROV and TSB for their support of my visit and loan of specimens/photographs and to Dr C. Argenti, Dr A. Soldano and Dr A. Tisi for information and loan of material conserved in their personal herbaria. References Aellen, P Amaranthus L. In: Guinochet, M. and Vilmorin, R. (eds), Flore de France 1. É ditions du Centre National de la Recherche Scientifique, pp Akeroyd, J Amaranthus L. In: Tutin, T. G. et al. (eds), Flora Europea, 2nd ed. Vol. 1. Cambridge Univ. Press, pp Anzalone, B Prodromo della Flora Romana (Elenco preliminare delle piante vascolari spontanee del Lazio). Aggiornamento. Parte 1a. Ann. Bot. Roma 52 (1994), Suppl. 11: 13. Cacciato, A Il genere Amaranthus a Roma e nel Lazio. Ann. Bot. Roma 28: Cacciato, A Amaranthus L. In: Pignatti, S. (ed.), Flora d Italia 1. Edagricole, pp Celesti-Grapow, L. et al Inventory of the non-native flora of Italy. Plant biosystem 143(2): Conti, F. et al. (eds) An annotated checklist of the Italian vascular flora. Palombi and Partner. Conti, F. et al Integrazioni alla checklist della flora vascolare italiana. Natura Vicentina 10 (2006): Costea, M. et al Preliminary results towards a revision of the Amaranthus hybridus complex (Amaranthaceae). Sida 19: Costea, M. et al The biology of Canadian weeds Amaranthus retrofl exus L., A. powellii S.Watson and A. hybridus L. Can. J. Plant Sci. 84: Greizerstein, E. et al Karyological studies in five wild species of amaranths. Cytologia 62: Greuter, W. et al Med-checklist 1. Pteridophyta, Gymnospermae, Dicotyledones (Acanthaceae Cneoraceae). Conserv. Jard. Bot. Ville Geneve. Haeupler, H. and Muer, T Bildatlas der Farn- und Bl ü tenpflanzen Deutschlands. Ulmer. H ü gin, G Einige Bemerkungen zu wenig bekannten Amaranthus -sippen (Amaranthaceae) Mitteleuropas. Willdenowia 16: Iamonico, D Notulae Amaranthus powellii S. Watson subsp. powellii Amaranthus powellii S. Watson subsp. bouchonii (Thell.) Costea & Carretero (Amaranthaceae). Notulae alla Checklist della flora italiana: 6. Inform. Bot. Ital. 40: 263. Iamonico, D. 2009a. Note ecologiche su Amaranthus blitoides S. Watson (Amaranthaceae) e suoi caratteri d invasivit à. Lagascalia 29: Iamonico, D. 2009b First record of Amaranthus powellii subsp. powellii (Amaranthaceae) in Lazio region (central Italy) with taxonomical, morphological, corologycal and ecological notes. Acta Bot. Malac. 34: Iamonico, D Biology, life-strategy and invasiveness of Amaranthus retrofl exus L. (Amarnathaceae) in central Italy: preliminary remarks. Bot. Serb. 34: Iamonico, D. and Del Guacchio, E Amaranthus powellii S. Watson subsp. powellii (Amaranthaceae), nuova per la flora esotica della Campania. Delpinoa 49 (2007), in press. International Organization for Plant Information International Plant Names Index index.html. Jonsell, B. (ed.) Flora Nordica. Chenopodiaceae to Fumariaceae 2. R. Swedish Acad. Sci. Kergu é len, M Index sinonimique de la flore de France. Lambdon, P. W. et al Alien flora of Europe: species diversity, temporal trends, geographical patterns and research needs. Preslia 80: Mosyakin, S. L. and Robertson, K. R Amaranthus L. In: Flora of North America Editorial Committee (ed.), Flora of North America North Mexico 4, part 1. Oxford Univ. Press, pp Raus, Th Amaranthus L. In: Strid, A. and Tan, K. (eds), Flora Hellenica 1. Koeltz Scientific Books. Stace, C. A New flora of British Isles, 2nd ed. Cambridge Univ. Press. Th ellung, A Amaranthus bouchonii Th ell. spec. (?) nov. Monde Plant. 27: 4 5. Tropicos Tropicos. Miss. Bot. Gard. tropicos.org. Wisskirchen, R. and Haeupler, H Standardliste der Farn- und Blütenpflanzen Deutschlands. Verlag Eugen Ulmer. Wohlgemuth, T. et al Swiss web flora, ver ethlm. 16

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