DE GRUYTER open HACQUETIA 13/2 • 2014, 285-296 DOI: 10.2478/hacq-2014-0004 TAXONOMICAL, MORPHOLOGICAL, ECOLOGICAL AND CHOROLOGICAL NOTES ON OXYBASIS CHENOPODIOIDES AND O. RUBRA (CHENOPODIACEAE) IN ITALY Duilio lAMGNICQi Abstract Gn the basis of floristic surveys, examination of herbarium specimens and review of literature, notes on the taxonomy, morphology, and ecological features are given of Oxybasis chenopodioides and morphologically fairly similar Oxybasis rubra. Their distributions in Italy according to Provinces are discussed and mapped. Key words: Amaranthaceae, Chenopodium, distribution, Italy, new records. Izvleček Na podlagi florističnih raziskav, proučevanj herbarijskih primerkov in pregleda literature smo podali opombe k taksonomiji, morfologiji in ekologiji vrste Oxybasis chenopodioides in morfološko zelo podobne Oxybasis rubra. Prikazali in obravnavali smo njihovo razširjenost v provincah Italije. Ključne besede: Amaranthaceae, Chenopodium, razširjenost, Italija, novi podatki. 1. INTRODUCTION The genus Chenopodium L. s. lat. has a worldwide distribution, and it includes about 150-200 species (Western Australian Herbarium 1998-2009, Mosyakin & Clemants 2002, 2008, Zhu et al. 2003, Verloove & Lambinon 2006, APG III 2009, Fuentes-Bazan et al. 2012a). It is considered taxo-nomically critical because of the morphological variability and lack of good characters and, to some extent, hybridization. Recent molecular studies clearly show that the genus in its widest sense is polyphyletic resulting in six independent lineages (Kadereit et al. 2003, 2005, Fuentes-Bazan et al. 2012a, 2012b). Consequently, a new taxonomic classification of the Chenopodioideae Burnett was proposed (Fuentes-Bazan et al. 2012a), including four tribes (Atriplicae Duby, Anserineae Dumort., Dysphaniae Pax, Axyridae G. Kadereit & Sukhor.) and ten genera (Axyris L., Blitum L., Ceratocaarpus L., Chenopodiastrum S. Fuentes, Uotila & Borsch, Chenopodium L., Dys- phania R. Br., Krascheninnikovia Gueldenst., Li-pandra Moq., Oxybasis Kar. & Kir., Teloxys Moq.). The genus Oxybasis was resurrected to accomodate the Chenopodium species with branched stem, leaves not in basal rosette, inflorescence in not fleshy glomerules and usually dimorphic flowers with 3-5 tepals and horizontal and vertical seeds (Fuentes-Bazan et al. 2012a, Sukhorukov 2013). Four European and Italian taxa (Iamonico 2012b) are included in the genus: O. glauca (L.) S. Fuentes, Uotila & Borsch (= Chenopodium glau-cum L.), O. urbica (L.) S. Fuentes, Uotila & Borsch (= Chenopodium urbicum L.), O. chenopodioides (L.) S. Fuentes, Uotila & Borsch (= Chenopodium botryodes Sm.), O. rubra (L.) S. Fuentes, Uotila & Borsch (= Chenopodium rubrum L.). Oxybasis chenopodioides and O. rubra represent a critical pair of species having fairly similar gross morphology, even though they do not seem to be close relatives (see e.g. Fuentes-Bazan 2012a). Variability of vegetative morphology and difficulties to see essential floral and fruit charac- 1 Laboratory of Phytogeography and Applied Geobotany, Department DPTA, University of Rome Sapienza, via Fla-minia 72, 00196 Rome, Italy, d.iamonico@yahoo.it ters have resulted in misinterpretations of names and misidentifications. The aim of this work is to provide clarity to the taxonomy, morphology and ecology and revise the chorology of the species in Italy as a part of the treatment of Chenopodicaceae for the new Flora of Italy, Ed. S. Pignatti (see e.g. Iamonico 2009, 2010, 2011a, 2012a, 2013, Iamonico & Ka-dereit 2012, Iamonico & Sukhorukov 2013). 2. MATERIALS AND METHODS The present study is based on an extensive analysis of literature (protologues included), personal field investigations (specimens collected by the author are preserved in the Herbarium Flaminio, HFLA), examination of herbarium material from AO, FI, RO and TR and checking of images of Linnean specimens, available on the web at Linnean Society Collections Online 2009). The nomenclature follows Conti et al. (2005) and Iamonico (2012b). The descriptions are based on Italian material and personal observations. 3. RESULTS AND DISCUSSION Notes on Taxonomy and morphology Oxybasis chenopodioides was first described as Bli-tum chenopodioides by Linnaeus (1771) in Ta-taria; nunc in Suecia". Hiitonen (1933) proposed subgeneric rank for Blitum L. under Chenopodi-um. Aellen (1933) proposed the new combination Chenopodium chenopodioides (L.) Aellen (= Blitum chenopodioides L.), and discussed the affinity of the species with the related C. crassifolium Hornem. and C. rubrum, the first one considered a synonym of C. chenopodioides. As regard the relationship with C. rubrum, Aellen (1933) examined the photograph of two Linnaean specimens (Nos. 14.3 and 14.4, both received by Linnaeus from Arduino and labelled as "tataricum"), concluding that the original description was not written on the basis of these specimens because of the disharmony between their characters and the original description. He also supposed that the sheets could represent the taxon called C. rubrum. Problems and doubts of the correct application and typification of the name B. cheno-podioides led some authors (e.g. Brenan 1964) to use the next safe name: C. botryodes Sm. The Italian floras (e.g. Fiori 1923, Zangheri 1976, Pignatti 1982, Conti et al. 2005) always accept C. botryo-des (sometimes C. chenopodioides doubtfully indicated in synonymy). On the basis of an extensive discussion, Uotila (2001a) removes the possibility that the name C. chenopodioides might be a synonym of C. rubrum, and a neotype was designated. Currently, the name C. chenopodioides was accepted in several important works (e.g. Akeroyd 1993, Uotila 1990, 2001b, Clemants & Mosyakin 2003, Zhu et al. 2003). On the basis of recent molecular studies, Fuentes-Bazan et al. (2012a) recently resurrected the genus Oxybasis and proposed the new combinations O. chenopodioides and O. rubra. On the basis of the protologues (Linnaeus 1753, 1771), O. chenopodioides and O. rubra differ in the tepals features: connate nearly to tip [Linnaeus (1771) reported "Calix concavus, con-nivens ."] and free or connate only at base, respectively. Other characters [shape of the leaves (subtriangular vs. subrhomboidal) and margins (entire or slightly toothed with obtuse tooths vs. toothed with acute tooths)] were often considered as diagnostic, but they partially overlap (Uotila 2001). In the Italian specimens the only constant diagnostic character is the difference in the te-pals. As to the leaves, they are usually subtrin-gular with entire margins (sometimes coarsely toothed with obtuse tooths) and with truncate base in O. chenopodioides, while subrhomboidal with toothed margins (tooths acute or acuminate with tip to distal part of the blade) and cuneate base in O. rubra. A further diagnostic character can be seen in the field: O. chenopodioides is usually prostrate and O. rubra erect (Table 1). Oxybasis chenopodioides (L.) S. Fuentes, Uotila & Borsch, Willdenowia 42: 15 (2012) Bas.: Blitum chenopodiodes L., Mant. Pl. Altera: 170 (1771) - Type (designated by Uotila 2001: 95-96): Russia, Republic of Dagestan, XIX century, C. Steven s.n. (neotype H-1037202). ^ Chenopodium chenopodioides (L.) Aellen, Os- tenia, Festschr. Osten.: 98 (1903). ^ Chenopodium botryoides Sm., Engl. Bot. 32, tab. 2247 (1811). Description: Annual herb (therophyte), 550 cm tall. Stem usually prostrate or prostrate-ascending, reddish, glabrous (rarely slightly farinose), branched at the basal part. Leaf blade ovate, subtriangular (2-4 x 4-7 cm), entire (sometimes with 1 basal tooth on both sides) or sometimes (distal leaves) irregularly toothed with ob- Table 1: Diagnostic characters between Oxybasis chenopodioides and O. rubra. Tabela 1: Razlikovalni znaki med vrstama Oxybasis chenopodioides in O. rubra. Oxybasis chenopodioides Oxybasis rubra Tepals of lateral flowers connate almost to tip Leaf blade subtriangular, entire (somentimes with 1 basal tooth on both sides) or somentimes (distal leaves) irregularly toothed with obtuse teeth and truncate base Plants usually prostrate or prostrate-ascending Tepals of lateral flowers connate halfway or less Leaf blade subrhomboidal, toothed with acute or acuminate teeth (rarely entire) and base cuneate Plants erect tuse tooths, stalked (petiole 1-4 cm), slightly succulent, often reddish; base truncate, apex obtuse. Floral glomerules arranged in terminal and/or lateral spike-like inflorescences; flowers dimorphic: terminal flowers bisexual, with 5 tepals; lateral flowers bisexual and/or female, usually with 3 tepals (tepals always connate nearly to tip); fruit covered by the tepals, with one seed, usually vertical, brown, shiny (diameter 0.6-0.8 mm). 2n = 18 (Uotila 1990). Vegetation and ecology: O. chenopodioides prefers to colonize habitats characterized by a seasonal cycle of submersion (mainly during the winter, with rain) and emersion (during the summer and autumn). The life cycle is short, 2-3(-4) months between August and November, and is completed during the submersion phase. Soil is humid, scarcely evoluted, scarcely structured and with a fine texture (pers. obs.). O. chenopodioides frequently grows along sandy coastal habitats near the sea level, even though (for example in Sardinia; see lamonico & Calvia 2010) it may be also found far from the coast line, up to 160-180 m a.s.l. (Fig. 1). During the later summer, O. chenopodioides can be often found from halophil-ous vegetation communities ascribed to the class Mannu river Oxybasis chenopodioides population * i* - » i, JWt V J- t « Figure 1: Oxybasis chenopodioides population along the Manu river (near Lake Coghinas), at locality Muros, in Olbia-Tempio Province of Sardinia. Photo by G. Calvia. Slika 1: Populacija Oxybasis chenopodioides ob reki Manu (v bližini jezera Coghinas), lokacija Muros v provinci Olbia-Tempio, Sardinija. Foto: G. Calvia. Figure 2: Oxybasis chenopodioides and Salicornia patula associated on shore of the Manu river in Delta del Po Park, Emilia-Romagna Region of Ravenna Province, north-eastern Italy. Photo by S. Montanari. Slika 2: Oxybasis chenopodioides in Salicornia patula na obali reke Manu v delti reke Pad, regija Emilia-Romagna, provinca Ravenna, severovzhodna Italija. Foto: S. Montanari. Thero-Salicornietea, order Thero-Salicornietalia (for example, in Emilia-Romagna region; see lamoni-co & Montanari 2010); the phytocenoses are characterized by the dominance of O. chenopodioides, Suaeda maritima. and Salicornia patula (Fig. 2). O. chenopodioides can be considered a halo-nitro-philous species. General Distribution: Oxybasis chenopodioides is distributed in Europe, central and south-western Asia, northern Africa, and North America (Clemants & Mosyakin 2003, Zhu et al. 2003, Uo-tila 2011). Occurrence in Italy: Earlier Oxybasis cheno-podioides has been recorded in the Regions of Friuli-Venezia Giulia, Veneto, Lazio and Campania (Conti et al. 2005), but it was found recently from four further Regions: Emilia-Romagna (Iamonico & Montanari 2010), Sardina (Iamon-ico & Calvia 2010), Puglia (Iamonico 2011b) and Toscana (Iamonico et al. 2013). Concerning the Lombardia region (northern Italy) Pignatti (1982) reported "Chenopodium botryoides lit. raram. all'interno (Bergam.)" (= '"Chenopodium botryoides . coasts . rarely in the inland (Bergamo)") and probably referred to Fiori and Paoletti (1898, who reported "Qua e la nel Bergamasco ."), Fiori (1923) (who reported "Bergam.") and Rodegh-er & Venanzi (1894). The same record was also published by Martini (2012) in the Flora Vasco-lare della Lombardia centro-orientale (= Vascular Floras of the central-eastern Lombardia). These authors also indicated another station in Bergamo province (along the Brembo river) on the basis of D'auria & Zavagno (1995). Conti et al. (2005) recently excluded O. chenopodioides (sub Cheno-podium botryoides) from Lombardia ["segnalata in precedenza per errore" ("recorded in the past by mistake'")] considering the species as casual (G. Galasso, personal communication). In fact, there are not clear reasons to consider O. chenopodioides as alien species for Bergamo province. This species naturally occurs in humid areas (both coastal and inland) of Europe, Asia and North Africa and it can be considered autochtonous for Italy. The discover by D'Auria & Zavagni (1995) along the riverbanks of the Brembo river allows to hypotize that O. chenopodioides is authoctonous for the Bergamo territory. We found at FI one specimen collected in locality "Valle di Sermide" (Mantova Province, Lombardia Region) in the XIX century (ante 1842) sub Ch. rubrum and previously revised (1977) by P. Uotila. We confirmed (D. Iamonico, January 2010) the determination by Uotila, so highlighting for the first time the occurrence fo this species in the Province. Finally, the Herbarium examination allowed also to found one specimen in FI (sub Ch. mela-nospermum) collected in the XIX century (ante Dec 1866) in Messina Province (Sicily) in locality "Murtiddi". We confirmed (D. Iamonico, January 2008) the previous determination by P. Uotila (1977) (see the final specimen list). O. chenopodioides was not indicated in the recent Catalogue of plants growing in Sicily (Giardina et al. 2007) and in the subsequent update (Raimondo & Spada-ro 2009). G. Domina (personal communication) confirms the absence of findings and literature data of the species for Sicily. Therefore, the specimen found reprensents a new floristic data for the region. Since the date of collection, it has to be considered as disappeared occerrence. At the current state of knowledge, we cannot know the real reasons for this apparent extinction. We know that the genus Chenopodium s. lat. is very poorly known and studied in Italy and that the urbanized areas (where several Chenopodium taxa grow) are less investigated then the natural ones (Iamonico 2009), so it is possible that further field investigations could allow to rediscover the taxon. Oxybasis chenopodioides has a fragmented distribution in Italy and it can be considered rare. The main reason for this is the rarity of the habitat in which the specie occurs. These habitat are threatened by human activities (mainly tourism) that could cause the extition of O. chenodopioides and other associated taxa. The extinction can be confirmed for the area named "Valli di Sermide" (Mantova Province, Lombardia region, northern Italy) where the habitat cannot be see nowday because of human recoveries (G. Galasso, personal communication). List of Italian Regions and Provinces in which O. chenopodioides occurs (Fig. 4a): - Friuli-Venezia Giulia: Udine (Pospichal 1897, Gortani & Gortani 1905, sub C. botryoides); - Veneto: Rovigo (Benetti & Marchiori 1995, Masin et al. 2009, sub C. botryoides); - Lombardia: Bergamo (Rodegher & Venanzi 1894, Fiori 1923, sub C. rubrum ß-botryoides, Pig-natti 1982, sub C. botryoides, D'Auria & Zavag-no 1995, Martini 2012), Mantova (first record for the Province - see the final specimina list); - Emilia-Romagna: Ravenna (Iamonico & Mon-tanari 2010); - Toscana: Lucca (not recorded since 1861), Pisa (not recorded since 1862), Siena (Iamonico et al. 2013). - Lazio: Latina, Roma (not recorded since 1920) (Anzalone et al. 2010, sub C. botryoides); - Campania: Napoli (Terracciano 1910, 1917, sub C. boryoides, Agostini 1956, sub C. boryo-ides, Motti & Ricciardi 2005); - Puglia: Foggia (Iamonico 2011b); - Sardegna: Olbia-Tempio (Iamonico & Calvia 2010); - Sicily: Messina (first record for the region -see final specimina list). Oxybasis rubra (L.) S. Fuentes, Uotila & Borsch, Willdenowia 42: 15 (2012) = Chenopodium rubrum L., Sp. Pl. 1: 218 (1753) -Type (designated by Uotila 1993): Herb. Linnaeus 313.5 (lectotype LINN!). ^ Blitum rubrum (L.) Rchb., Fl. Germ. Excurs. 2(1): 582 (1832). Description: Annual herb (therophyte), 10100 cm tall. Stem erect, green or ± reddish, glabrous or nearly so, branched. Leaf blade ovate or lanceolate, subrhomboidal (2.5-6 x 4-8 cm), toothed with teeths acute or acuminate (rarely entire), stalked (petiole 2-4 cm), sometimes slightly succulent, often red; base cuneate, apex acute. Floral glomerules arranged in terminal and/or lateral spike-like inflorescences; flowers dimorphic: terminal flowers bisexual, with 5 tepals; lateral flowers bisexual and/or female, usually with 3 tepals (tepals free or connate up to the middle at most); fruit with one seed, usually vertical, brown, shiny (diameter 0.6-1.1 mm). 2n = 18, 36 (Uotila 1990). Vegetation and ecology: Oxybasis rubra mostly occurs in ruderal habitats and wastelands, at altitudes between 0-600 m a.s.l. It can be found in vegetation communities of the class Stellarietea mediae Tüxen, Lohmeyer & Preising ex von Rochow 1951, order Chenopodietalia muralis Br.-Bl. 1936 (personal observation). Recently, the new association Polygono orientalis-Chenopodietum rubri (class Bidentetea tripartitae, order Bidentetalia tripartitae - Sciandrello 2009) was described from Sicily for phytocenoses that occur in habitat characterized by seasonel cycles of submersion (during the winter-spring period) and emersion (during the summer-autumn period) on clayey-muddy, nitrophilous and humid soils. O. rubra is a dominant species in this association. Floristic investigations (carried out in 2008) in Finale Ligure locality (Province of Savona in Liguria, northwestern Italy) also revealed that O. rubra grows in sandy coastal zone (Figure 3) potentially occupied by the association Salsolo kali-Cakiletum mar-itimae (Class Cakiletea maritimae, order Euphorbi-etalia peplis). O. rubra seems to have wider ecological demands than O. chenopodioides: it can colonize both synanthrophic habitats (mostly open wastelands with low soil humidity) and riv-erbanks (with humid soils also during the emersion phase). The species can be considered nitro-philous or hygro-nitrophilous. General Distribution: Oxybasis rubra is distributed in Europe (mainly in Central territories, while it is rare in the Mediterranean area and it does not occurs in the the extreme east and north territories), central and south-western Asia, and in North America (Clemants & Mosyakin 2003, Zhu et al. 2003, Uotila 1990, 2011). The occurrence in Italy has a general interest because of the rarity of this species in the Mediterranean area (see e.g. Jalas & Suominen 1980, Greuter et al. 1984). Occurrence in Italy: Oxybasis rubra was reported from nine Regions, Friuli-Venezia Giulia, Veneto, Trentino-Alto Adige, Lombardia, Ligu-ria, Emilia-Romagna, Toscana, Abruzzo, Campania and Basilicata by Conti et al. (2005), and recently from two more, Valle d'Aosta by Pistar-ino et al. (2010) and Puglia by lamonico & Dura (2013) and it is mentioned but not confirmed from Umbria (Iamonico in press). In the past the species was recorded by mistake from Lazio (Iamonico & Iberite 2008) and Sardegna Regions (Conti et al. 2007). Recent floristic surveys (January 2008) allowed to confirm the presence of O. rubra in Savona Province (locality Finale Ligure), where no record is known since the 1980s (G. Barberis, personal communication). Oxibasis rubra has a wider distribution in Italy than O. chenopodioides since it appears to have a Figure 3: Oxybasis rubra in sandy substrate in Savona Province of Liguria Region, north-western Italy. Photo by D. Longo. Slika 3: Vrsta Oxybasis rubra na peščeni podlagi v province Savona v regiji Liguria, severozahodna Italija. Foto: D. Longo. larger ecological demand, also colonizing rud-eral habitats at altitudes uop to 600 m a.s.l. However, its distribution is fragmented, especially in central and southern Italy. The species can be considered common in northern Italy, and less frequent in central and southern Italy. List of Italian Regions and Provinces in which O. rubra occurs (Fig. 4b): - Friuli-Venezia Giulia: Gorizia (Pospichal 1897, Poldini 2002, 2009), Pordenone (Poldini 2002), Trieste (Marchesetti 1896, Mazzena & Polli 1982), Udine (Poldini 2002); - Veneto: Belluno (Argenti & Lasen 2004), Pa-dova (Masin & Tietto 2005, 2006), Vicenza (Curti & Scortegagna 1998), Verona (Bianchi-ni 1976); - Trentino-Alto Adige: Bolzano (Wilhalm et al. 2006), Trento (Prosser 1995); - Lombardia: Bergamo (Pignatti 1982); - Piemonte: Alessandria (Iamonico & Soldano 2012b); - Valle d'Aosta: Aosta (Pistarino et al. 2010); Figure 4: Distribution map of Oxybasis chenopodioides (a) and Oxybasis rubra (b) in Italy according to Provinces. Provinces with present records are marked in black, old records in gray and doubtful records in light gray with question marks. Slika 4: Karte razširjenosti vrst Oxybasis chenopodioides (a) in Oxybasis rubra (b) v provincah Italije. Province z novimi podatki so označene s črno, stari podatki so predstavljeni s sivo in vprašljivi podatki s svetlo sivo in vprašajem. Liguria: Genova (Bertoloni 1804), Imperia (L. Aita et G. Barberis, personal communication) and Savona (in this latter Province the species is confirmed in the locality Finale Ligure). Emilia-Romagna: Ferrara (Beguinot 1910, Zavatti 1999-2000), Modena (Delfini 2003, Alessandrini et al. 2010), Parma (Ghilliani et al. 2004, Petraglia et al. 2005, Ghilliani 2010), Ravenna (Zangheri 1966, Lazzari et al. 2007, Bassi 2004); Toscana: Lucca [Montelucci (1964) reported ""^.'specialmente in base alla verticalita dei semi' e cheforse si avvicinano alla var. botryoides (Sm.) Fiori" ("^ 'especially based on the vertical seeds' that are maybe similar to the var. botryoides (Sm.) Fiori"), so the species can be considered doubt for this province), Massa-Carrara (Ferrarini & Marchetti (1994) indicated a question mark, so the species is to be considered doubt), Pisa (Caruel 1860, Baroni 1897-1908); Umbria: Perugia (Bencivenga et al. 1977-1978, Iamonico in press); Abruzzo: Chieti (Leporatti et al. 2001a, 2001b); Campania: Napoli (Tenore 1823, 1831, Motti & Ricciardi 2005); Puglia: Taranto (Iamonico in press, Iamonico & Dura 2013); Basilicata: Matera (Gavioli 1947); Sicilia: Caltanissetta (Giardina et al. 2007, Sciandrello 2009). 4. ACKNOWLEDGEMENTS Thanks are due to Directors and Curators of all quoted Herbaria for their support during my visits, loans of specimens/photographs or requests for information. I am also greatful to A. Alessan-drini (Bologna), C. Argenti (Belluno), P. V. Ar-rigoni (Firenze), G. Barberis (Genova), F. Conti and F. Bartolucci (Barisciano), E. Del Guacchio (Salerno), G. Galasso and E. Banfi (Milano), M. Iberite (Latina), F. Prosser (Rovereto), A. Sol-dano (Vercelli), M. Vidali (Trieste) and T. Wilhalm (Bolzano) for information on the distribution. I thank my friends G. Calvia (Berchidda), D. Longo (Genova) and S. Montanari (Pangipane) for the permission to reproduce their photographs. Finally, I am greatful to P. Uotila and the other anonymous reviewer for the constructive comments. 5. REFERENCES Aellen, P. L. 1933: Nomenklatorische Bemerkungen zu einigen Chenopodien. Ostenia, Fest-schr. Osten: 89-101. Agostini, R. 1956: Aster squamatus Hieron. nuova avventizia della flora campana. Delpinoa 9: 164. Akeroyd, J. R. 1993: Chenopodium L. In: Tutin, T. G., Heywood, V. H., Burges, N. A., Moore, D. M., Valentine, D. H., Walters, S. M. & Webb D. A. (eds.), Flora Europaea 1. Ed. 2. Cambridge University Press, Cambridge, pp. 111-114. Alessandrini, A., Delfini, L., Ferrari, P., Fiandri, F., Gualmini, M., Lodesani, U. & Santini, C. 2010: Flora del Modenese. Censimento, Ana-lisi, tutela. 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