This art icle was downloaded by: [ Universit a di Palerm o] On: 06 February 2014, At : 02: 59 Publisher: Taylor & Francis I nform a Lt d Regist ered in England and Wales Regist ered Num ber: 1072954 Regist ered office: Mort im er House, 37- 41 Mort im er St reet , London W1T 3JH, UK Plant Biosystems - An International Journal Dealing with all Aspects of Plant Biology: Official Journal of the Societa Botanica Italiana Publicat ion det ails, including inst ruct ions f or aut hors and subscript ion inf ormat ion: ht t p: / / www. t andf online. com/ loi/ t plb20 A critical conspectus of Italian Isoetes (Isoetaceae) a A. Troia & W. Greut er b a Dipart iment o di Scienze e Tecnologie Biologiche Chimiche e Farmaceut iche, Sezione di Bot anica ed Ecologia Veget ale, Universit à degli St udi di Palermo, Via Archiraf i 38, 90123 Palermo, It aly b Herbarium Medit erraneum Panormit anum, Universit à degli St udi di Palermo, c/ o Ort o Bot anico, Via Lincoln 2/ A, 90133 Palermo, It aly Accept ed aut hor version post ed online: 08 Jan 2014. Published online: 05 Feb 2014. To cite this article: A. Troia & W. Greut er , Plant Biosyst ems - An Int ernat ional Journal Dealing wit h all Aspect s of Plant Biology (2014): A crit ical conspect us of It alian Isoet es (Isoet aceae), Plant Biosyst ems - An Int ernat ional Journal Dealing wit h all Aspect s of Plant Biology: Of f icial Journal of t he Societ a Bot anica It aliana, DOI: 10. 1080/ 11263504. 2013. 878409 To link to this article: ht t p: / / dx. doi. org/ 10. 1080/ 11263504. 2013. 878409 PLEASE SCROLL DOWN FOR ARTI CLE Taylor & Francis m akes every effort t o ensure t he accuracy of all t he inform at ion ( t he “ Cont ent ” ) cont ained in t he publicat ions on our plat form . However, Taylor & Francis, our agent s, and our licensors m ake no represent at ions or warrant ies what soever as t o t he accuracy, com plet eness, or suit abilit y for any purpose of t he Cont ent . Any opinions and views expressed in t his publicat ion are t he opinions and views of t he aut hors, and are not t he views of or endorsed by Taylor & Francis. 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GREUTER2 1 Downloaded by [Universita di Palermo] at 02:59 06 February 2014 Dipartimento di Scienze e Tecnologie Biologiche Chimiche e Farmaceutiche, Sezione di Botanica ed Ecologia Vegetale, Università degli Studi di Palermo, Via Archirafi 38, 90123 Palermo, Italy and 2Herbarium Mediterraneum Panormitanum, Università degli Studi di Palermo, c/o Orto Botanico, Via Lincoln 2/A, 90133 Palermo, Italy Abstract The work on the Isoetaceae account for the Critical Flora of Italy has resulted in a new, synonymic inventory of Italian Isoetes taxa, based on herbarium studies, observation by scanning electron microscopy and critical re-examination of the previously published works. Ten species are listed. The names I. longissima and I. gymnocarpa are restored as correct, to replace I. velata and I. subinermis, respectively. Eighteen lectotypes, one epitype and one neotype are designated. A dichotomous key to Italian taxa is provided. Keywords: Italian flora, Lycopodiophyta, morphology, pteridophytes, taxonomy, typification Introduction The definition, description and identification of Isoetes species remain difficult because, due to the extreme morphological simplicity of the plants, combined with phenotypic plasticity, the number of stable and useful diagnostic characters is very limited (Hickey et al. 1989). As stated by Kott and Britton (1983), “to accept any concept of species in the genus Isoetes L. one must first accept that perhaps the only constant in this group is its variation”. For practical purposes, it is desirable to define species in such a way that one can discriminate them by traits easily observed on herbarium specimens and live plants, without resorting to caryology, DNA analysis or scanning electron microscopy (SEM). Data obtained by such advanced techniques are essential to underpin a classification that has phylogenetic value, and we have used and integrated them if and when available. In the process, we have found that it is risky to rely on a single if important feature, e.g. macrospore ornamentation, to define taxa. General habit, ecological preferences, and morphology of leaves, scales/phyllopodia, sporangial area, and macro- and microspores have to be considered in conjunction. The present synopsis of Italian Isoetaceae is the first in a series of contributions toward a Critical Flora of Italy (Pignotti 2006). It consists of a new, critical synonymic inventory of Italian Isoetes taxa, with a practical key for their identification. It includes some nomenclatural novelties and new type designations. For some critical taxa here considered, particularly in the Isoetes longissima and I. histrix groups, megaspore and microspore features were observed by SEM. The SEM results are not reported here, but will be published separately. Materials and methods We have consulted the herbaria FI and PAL (including PAL-Gr) for their contents of Italian Isoetes and selected specimens from other countries. We have obtained on loan potential type specimens from B and TO, and seen high-resolution digital images available online, or provided on request, from several herbaria, including BORD, MO, MPU, P, PAD and RO (acronyms according to Thiers 2013). Aware of the assets of modern Web-based technology enabling direct online access to relevant data (Martellos & Attorre 2012), we have listed in Correspondence: A. Troia, Dipartimento di Scienze e Tecnologie Biologiche Chimiche e Farmaceutiche, Sezione di Botanica ed Ecologia Vegetale, Università degli Studi di Palermo, Via Archirafi 38, I-90123 Palermo, Italy. Tel: þ 39 091 23891260. Fax: þ 39 091 6165997. Email: angelo.troia@libero.it q 2014 Società Botanica Italiana 2 A. Troia and W. Greuter Downloaded by [Universita di Palermo] at 02:59 06 February 2014 Table I. URIs of type specimens cited in the text and available on the Internet. Specimens are listed in the same order they are cited. Taxon Specimen description (herbarium code) Isoetes echinospora Durieu Isoetes longissima Bory MO 1852527 P 466542 Lectotype Lectotype Isoetes longissima Bory Isoetes setacea var. delilei Isoetes setacea var. delilei Isoetes setacea var. delilei Isoetes setacea var. delilei Isoetes velata A. Braun P 466541 p.p. P 202851 p.p. P 202852 P 202851 p.p. MPU 014139 B 20 0096552 Isolectotype Lectotype Paralectotype Paralectotype Paralectotype Epitype TO Lectotype PAL 6049 Isolectotype B 20 0096543 FI 002102 TO Lectotype (or holotype?) Lectotype Lectotype TO Lectotype MO 1852617 P 507734 Lectotype Isolectotype (same as for I. histrix [unranked] grandis) B 20 0107668 Lectotype B 20 0107687 Lectotype P 507729 Isolectotype (same as for I. histrix [unranked] solitaria) B 20 0107589 p.p. Lectotype B 20 0107675 p.p. Lectotype PAL 6036 TO Lectotype Lectotype B 20 0107700 Lectotype B 20 0083454 Lectotype PAL 6044 P 00202838 Neotype Isolectotype Bory Bory Bory Bory Isoetes velata [unranked] sicula Ces. & De Not. ex Gennari Isoetes velata [unranked] sicula Ces. & De Not. ex Gennari Isoetes velata var. brevifolia A. Braun Isoetes tiguliana Gennari Isoetes dubia Gennari Isoetes dubia var. maculosa Gennari Isoetes histrix Bory Isoetes histrix Bory Isoetes histrix f. loricata A. Braun Isoetes histrix [unranked] grandis A. Braun Isoetes histrix [unranked] nana A. Braun Isoetes histrix [unranked] nana A. Braun Isoetes histrix f. desquamata A. Braun Isoetes histrix [unranked] solitaria A. Braun Isoetes histrix [unranked] longispina A. Braun Isoetes histrix var. lacunosa Tod. Isoetes gymnocarpa (Gennari) A. Braun Cephaloceraton histrix var. subinerme Gennari Isoetes histrix [unranked] brevispina A. Braun Isoetes sicula Tod. Isoetes durieui Bory Type Table I the uniform resource identifiers (URIs) of type images that are available on the Internet. In the Journal’s online version, the tabulated URIs function as direct links, and they also underly the citations of the specimens in the main text. A synonymic synopsis of Isoetes in Italy In the absence of a conclusive phylogenetic analysis, we have grouped the species conventionally, by habitat type: submerged (species nos 1–3), amphibious URI http://www.tropicos.org/Image/51712 http://dsiphoto.mnhn.fr/sonnerat/apiINFO/IE/IE20050719/ P00466542.jpg http://sonneratphoto.mnhn.fr/2010/08/12/4/P00466541.jpg http://sonneratphoto.mnhn.fr/2010/08/05/2//P00202851.jpg http://sonneratphoto.mnhn.fr/2010/08/05/2//P00202852.jpg http://sonneratphoto.mnhn.fr/2010/08/05/2//P00202851.jpg http://plants.jstor.org/specimen/mpu014139 http://ww2.bgbm.org/herbarium/images/B/20/00/96/55/b_20_ 0096552.jpg http://www.unito.it/unitoWAR/page/dipartimenti7/D071/D071_ campioni1 http://147.163.105.223/zoomify/view_img.asp?ic¼6049 http://ww2.bgbm.org/herbarium/images/B/20/00/96/54/b_20_ 0096543.jpg http://parlatore.msn.unifi.it/img72/FI002102.jpg http://www.unito.it/unitoWAR/page/dipartimenti7/D071/D071_ campioni1 http://www.unito.it/unitoWAR/page/dipartimenti7/D071/D071_ campioni1 http://www.tropicos.org/Image/3947 http://dsiphoto.mnhn.fr/sonnerat/apiJOIN/JF/JF20060103/ P00507734.jpg http://ww2.bgbm.org/herbarium/images/B/20/01/07/66/b_20_ 0107668.jpg http://ww2.bgbm.org/herbarium/images/B/20/01/07/68/b_20_ 0107687.jpg http://dsiphoto.mnhn.fr/sonnerat/apiJOIN/JF/JF20060103/ P00507729.jpg http://ww2.bgbm.org/herbarium/images/B/20/01/07/58/b_20_ 0107589.jpg http://ww2.bgbm.org/herbarium/images/B/20/01/07/67/b_20_ 0107675.jpg http://147.163.105.223/zoomify/view_img.asp?ic¼6036 http://www.unito.it/unitoWAR/page/dipartimenti7/D071/D071_ campioni1 http://ww2.bgbm.org/herbarium/images/B/20/01/07/70/b_20_ 0107700.jpg http://ww2.bgbm.org/herbarium/images/B/20/00/83/45/b_20_ 0083454.jpg http://147.163.105.223/zoomify/view_img.asp?ic¼6044 http://sonneratphoto.mnhn.fr/2010/08/05/2//P00202838.jpg (nos 4 and 5), and terrestrial (nos 6 – 10), and arranged them by morphological similarity within groups. (1) Isoetes malinverniana Ces. & De Not., Index Seminum (Genoa) 1858: 36. 1858 ; Calamaria malinverniana (Ces. & De Not.) Kuntze, Revis. Gen. Pl. 1-2: 828. 1891. Protologue locality: [Italy]“circa pagos Greggio et Oldenico in Pedemontii ditione Vercellensi”. Downloaded by [Universita di Palermo] at 02:59 06 February 2014 Critical conspectus of Italian Isoetes Lectotype (Soldano & Badino 1990: 66): “li 20 luglio [1858] a Greggio in un fosso raccoglitore della casa Gattinara verso Sesia”, Malinverni (RO, Herb. Cesati [image!]) (Figure 1). (2) Isoetes echinospora Durieu in Bull. Soc. Bot. France 8: 164. 1861. Protologue localities: “dans quelques lacs du centre de la France”. Lectotype (Pfeiffer 1922: 155; Prada 1983: 90): “Lac de Guéry, Puy de Dome (France), à 30 à 40 cm au dessous du niveau de l’eau”, 24 August 1861, Durieu (MO 1852527 [image!]). (3) Isoetes sabatina Troia & Azzella in Pl. Biosyst. 147 (online): 1052. 2013 [8 Apr 2013]. Holotype (Troia & Azzella 2013): “Lago di Bracciano (Roma), località Polline (UTM 33T 275.4667) at a depth of 1.2 m, on sandy substrate, 163 m a.s.l.”, 18 August 2010, Azzella (FI!; isotype: RO!). Paratypes at FI, RO, PAL!. (4) Isoetes longissima Bory in Compt. Rend. Hebd. Séances Acad. Sci. 18: 1165. 1844 ; Isoetes velata var. longissima (Bory) A. Braun in Bory & Durieu, Expl. Sci. Algérie, Atlas: tab. 37, fig. 2. 1849 ; Isoetes velata subsp. longissima (Bory) Greuter & Burdet, Med-Checklist 1: 5. 1984. Protologue locality: [Algeria] “dans une des mares des forets qui environnent le lac Houbéira, au canton de la Calle”. Lectotype (designated here): “La Calle, fond d’une Mare dans les Forets pres du Lac Houbéira”, 31 May 1841, Durieu ex herb. Bory St Vincent (P 466542 [image!]; isolectotype: P 466541 p.p., detached leaves to the right [image!]). Note – Sheet 466541 (P) bears two labels. One, referring to a bunch of detached leaves mounted on the right that is obviously part of the original gathering, reads “Fructifications 3 détachées du collet tuberculeux de l’Isoëtes longissima, pour l’étude. Eaux profondes de La Calle par Durieu 1841”. The other belongs with three whole plants mounted to the left, collected [at the locus classicus] by Durieu on 24 June 1844 and received by Bory subsequently: “Portée à la maison par Durieu le 30 décembre 1844”. In view of these dates, we do not consider the latter specimen, nor two other ones collected by Durieu on 24 May 1844 (P 466539 and 466540), to be part of the original material. ¼ Isoetes setacea var. delilei Bory in Compt. Rend. Hebd. Séances Acad. Sci. 18: 1165. 1844 ; Isoetes decipiens [‘descipiens’ ] Bory in Compt. Rend. Hebd. Séances Acad. Sci. 20: 619. 1846. Protologue locality: [Algeria] “sur plusiers points du canton de la Calle, ( . . . ) entre le lacs el Mehla et el-Houbéira; dans les sables d’Aı̈n-Treard; ( . . . ) près du douair de Ben-Atia ( . . . ), etc.” Lectotype (designated here): “Isoetes Gouani B., (La Calle, par Durieu, 7 juin 1841), individu venu dans le sable marécageux, non inondé, dans les bois d’Aı̈n-Treard”, 7 June 1841, Durieu ex herb. Bory St Vincent (P 202851 p.p., left-hand specimen [image!]). Paralectotypes [images!]: P 202852, P 202851 p.p., middle specimen (as “Isoetes Gouani B.” [ined.]), MPU 014139 (as “Isoetes descipiens Bory”). – A specimen of the same taxon from the La Calle area, MPU 014138 (as “Isoetes Bovei Bory” [ined.] [image!]), was collected by Durieu in 1844 and is probably not original material. ¼ Isoetes velata A. Braun in Bory & Durieu, Expl. Sci. Algérie, Atlas: tab. 37, fig. 1. 1849, nom. illeg. – Illegitimate due to inclusion, as a variety, of the earlier, legitimate I. longissima, but not based Figure 1. Herbarium label of the single historical type not yet available on the Internet: Isoetes malinverniana Ces. & De Not. (RO, Herb. Cesati: lectotype). Downloaded by [Universita di Palermo] at 02:59 06 February 2014 4 A. Troia and W. Greuter automatically on the same type (McNeill et al. 2012: Art. 7.5). Protologue locality: none [by implication, described from Algeria]. Lectotype (designated here): tab. 37, fig. 1, in Bory & Durieu, Expl. Sci. Algérie, Atlas, 1849. Epitype (designated here): “Lieux marécageux dans les basses forêts de liège, près La Calle”, 24 Junee 1844, [Durieu ] (B 20 0096552!). ¼ Isoetes velata [unranked] sicula Ces. & De Not. ex Gennari in Comment. Soc. Crittog. Ital. 1: 103. 1861 (September). Protologue locality: [Sicily] “Piano Stoppa, Misilmeri”. Lectotype (designated here): “Isoetes velata sicula Cesati et DNtrs mss. et herbar. Al Piano della Stoppa presso Palermo, ne’ prati inondati durante l’inverno. Primavera ed Estate”, 1858, Todaro in Erb. Crittog. Ital. No. 105 (TO ex Herb. Gennari!; isolectotypes: FI!, PAL 6049!, etc.). – Paralectotype, from the Gennari herbarium (TO), collected by the Huet du Pavillon brothers in the same locality (“In paludosis piana della Stuppa, supra Misilmeri [Sicilia]”!), probably in 1856 (as in the isoparalectotype in FI, with printed label dated 24 May 1856). ? ¼ Isoetes velata var. brevifolia A. Braun in Monatsber. Königl. Preuss. Akad. Wiss. Berlin, 1863: 605. 1864. Protologue locality: [Sardinia] “zwischen Cala d’Ostia und Domus de Maria bei Pula, 22 Mai”. Lectotype (designated here; or holotype?): “Cala d’Ostia bei Pula”, 22 May 1863, Ascherson & Reinhardt (B 20 0096543!). – In B there is a single specimen from the Braun herbarium that agrees with the protologue citation (likely the holotype; but as duplicates may exist elsewhere we prefer designating it as lectotype). The plants show some macroand micromorphological traits (e.g. megaspore ornamentation) that might justify their maintenance as a distinct taxon, but more preferably live material is needed to confirm that assumption. Note 1 – Recent work by Romero et al. (2004) and our own observations show that Isoetes longissima and I. velata, already treated as mere varieties by Braun (in Bory & Durieu, Expl. Sci. Algérie, Atlas: tab. 37, figs 1 –2. 1849), belong to one and the same taxon. Note 2 – The plant known from Spain as “I. longissima” has recently been recognised as a (5) ?¼ ?¼ (6) different species, Isoetes fluitans M. I. Romero (Romero et al. 2004). Isoetes tiguliana Gennari in Comment. Soc. Crittog. Ital. 1: 42. 1861 (February) ; Isoetes tegulensis Gennari in Comment. Soc. Crittog. Ital. 1: 106. 1861 (September), nom. illeg. ; Isoetes velata var. tegulensis (Gennari) Fiori in Fiori & Paol., Fl. Italia 1: 25. 1896. Protologue locality: “In aquaeductibus prope Teulada in Sardinia australiori”. Lectotype (designated here): “Isoetes tegulensis (Genn!), Teulada”, May 1859, Bellisai (FI 002102!; isolectotype?: TO ex herb. Gennari!). – The collector’s name is not mentioned in the protologue but appears in a subsequent paper (Gennari 1861). We chose the FI specimen as type since it bears the collecting date, whereas its presumed duplicate in TO, from the Gennari herbarium, is undated. Isoetes dubia Gennari in Comment. Soc. Crittog. Ital. 1: 104. 1861 (September). Protologue locality: [Sardinia] “rigagnolo a ponente dell’isola Maddalena”. Lectotype (designated here): “Isoetes dubia Genn! Maddalena”, May 1861, [Gennari ] (TO ex herb. Gennari!). Isolectotype in FI (!). Isoetes dubia var. maculosa Gennari in Comment. Soc. Crittog. Ital. 1: 105. 1861 (September). Protologue locality: [Sardinia] “Vive in società con la specie” [Isoetes dubia, see above]. Lectotype (designated here): “Isoetes dubia b maculosa Genn!, Maddalena”, May 1861, [Gennari ] (TO ex herb. Gennari!). Isolectotype in FI (!). Note – The plants ascribed by Gennari to Isoetes dubia resemble those of I. tiguliana except by having echinate – aculeate rather than smooth microspores. Pending further studies to assess the variability of this character, we prefer to consider both microspore types as pertaining to a single species. Isoetes histrix Bory in Compt. Rend. Hebd. Séances Acad. Sci. 18: 1167. 1844 ; Cephaloceraton histrix (Bory) Gennari in Comment. Soc. Crittog. Ital. 1: 111. 1862. Protologue localities: [Algeria] “dans l’etendue du canton de la Calle, ( . . . ) dans tous les environs de Bonne, ( . . . ) et jusque sur les pentes de l’Eydoug, . . . abondamment dans le pays d’Oran . . . ”. Downloaded by [Universita di Palermo] at 02:59 06 February 2014 Critical conspectus of Italian Isoetes ¼ ¼ ¼ ¼ “Découverte en mai 1841, par M. Durieu”. Lectotype (Motelay & Vendryès 1883: 397; Pfeiffer 1922: 128): “La Calle, ( . . . )”, 30 Mar 1841, [Durieu ] (MO 1852617 ex herb. Motelay [image!]; isolectotypes: BORD [image!], P 507734 [image!]). – The plants used by Bory to describe his new species are in Paris (P 507729 – 507734). Motelay and Vendryès (1883: 397), however, qualified as “échantillons types” material in the “Herb. Motelay”, from one of Durieu’s original gatherings. As part of that material was distributed to others, theirs is a first-step lectotypification. One such duplicate was sent to Eaton and, with his Isoetes herbarium, ended up in St Louis (MO). Pfeiffer (1922: 128) designated the latter specimen as “type”, which must be accepted as a second-step lectotypification. Isoetes histrix f. loricata A. Braun in Monatsber. Königl. Preuss. Akad. Wiss. Berlin 1863: 617. 1864 ; I. histrix var. loricata (A. Braun) Batt. & Trab., Fl. Algérie Tunisie: 407. 1905. Protologue locality: see under the two following names, designating subordinate taxa. Lectotype (designated here): same as for I. histrix [unranked] grandis, below. Isoetes histrix [unranked] grandis A. Braun in Monatsber. Königl. Preuss. Akad. Wiss. Berlin 1863: 617. 1864. Protologue locality: [Algeria] “bei Mascara, Bona, etc.” Lectotype (designated here): “Isoetes Hystrix Durieu, plateaux sablonneux des collines aux environs de Mascara”, 22 mai 1844, Durieu (B 20 0107668!). Isoetes histrix [unranked] nana A. Braun in Monatsber. Königl. Preuss. Akad. Wiss. Berlin 1863: 617. 1864. Protologue locality: [Algeria] “Oran”. Lectotype (designated here): “Isoetes Hystrix [sic!] DR. [sic!] b. nana Al. Br., Algérie. Oran. Lieux arides et nus du plateau du Djebbel Santo”, 22 mars 1842, [Durieu ] (B 20 0107687!; isolectotype: P 507729 [image!]). Isoetes histrix f. desquamata A. Braun in Monatsber. Königl. Preuss. Akad. Wiss. Berlin 1863: 617. 1864 ; I. histrix var. desquamata (A. Braun) Batt. & Trab., Fl. Algérie Tunisie: 407. 1905. Protologue locality: see under the following names, designating subordinate taxa. 5 Lectotype (designated here): same as for I. histrix [unranked] solitaria, below. ¼ Isoetes histrix [unranked] solitaria A. Braun in Monatsber. Königl. Preuss. Akad. Wiss. Berlin 1863: 618. 1864. Protologue locality: [France] “Houat und Belle-Ile”. Lectotype (designated here): “Isoetes Delalandii, Ile de Houat, Avr 1852” (B 20 0107589 p.p., the three specimens at the top of the sheet!). ¼ Isoetes histrix [unranked] longispina A. Braun in Monatsber. Königl. Preuss. Akad. Wiss. Berlin 1863: 618. 1864. Protologue locality: [Algeria] “z. B. bei La Calle”. Lectotype (designated here): the first plant, top right, on the sheet labelled: “Isoetes Hystrix Durieu b scutellata, macrospora, La Calle, pentes et plateaux des collines dont il forme presque à lui seul le gazon”, 28 March–1 April 1841, [Durieu ] (B 107675 p.p., first plant, top right, on the sheet!). ¼ Isoetes histrix var. lacunosa Tod. in Giorn. Sci. Nat. Econ. Palermo 1: 251. 1866. Protologue locality: [Sicily] “Alcamo”. Lectotype (designated here): “Isoetes histrix Br. var. lacunosa, in arenosis collium regionis inferioris – Alcamo alla Fico”, [Todaro’s writing ] (PAL 6036!). – The undated specimen here selected as type is almost certainly part of the original material. Note – Several different taxa may perhaps be hidden under Isoetes histrix, even when the similar I. gymnocarpa is segregated (see below, also Bagella et al. 2011: 456). We tried to differentiate them using morphological traits, for example the presence of a ventral or dorsal central tooth in the phyllopodia, but found this character to be unreliable. A broad study, encompassing the whole distribution area and including molecular in addition to morphological criteria, is needed to disentangle this hotchpotch. At present, we refrain from an attempt at further subdivision. (7) Isoetes gymnocarpa (Gennari) A. Braun in Monatsber. Koenigl. Preuss. Akad. Wiss. Berlin 1863: 555. 1864 ; Cephaloceraton gymnocarpum Gennari in Comm. Soc. Crittog. Ital. 1: 113. 1862 ; C. pseudo-hystrix Gennari in Ann. Agr. Sicil., 1865: 175. 1865, nom. illeg. (illegitimate due to the inclusion, as a synonym, of the earlier, legitimate C. gymnocarpum). Protologue locality: [Sardinia] “Isola di Caprera”. Lectotype (designated here): First plant, top left, of the sample labelled “Cephaloceraton Downloaded by [Universita di Palermo] at 02:59 06 February 2014 6 A. Troia and W. Greuter gymnocarpum (Genn.!), Caprera”, Maggio 1861, Gennari (TO ex herb. Gennari p.p., first plant, top left, on the sheet!). – The sheet with the original material of Isoetes gymnocarpa is a mixture. Of its two plants, one has phyllopodia with long lateral spine-like teeth as is typical of I. histrix, the other (our lectotype) is congruent with the protologue in this respect. Indeed, Gennari himself reports that his new species grows together with the similar I. histrix. However, contrary to the original description, in the lectotype specimen the velum is complete, as already found by Braun (1864) when studying the I. gymnocarpa sample he had received from Gennari. In the I. histrix complex, presence or absence of the velum does not seem to be a reliable diagnostic feature: Rolleri and Prada (2007: 218) indeed report that “en I. histrix el velo suele ser completo pero occasionalmente falta en macrosporofilos”. ¼ Cephaloceraton histrix var. subinerme Gennari in Comm. Soc. Crittog. Ital. 1: 112. 1862 ; Isoetes histrix [unranked] subinermis A. Braun in Monatsber. Königl. Preuss. Akad. Wiss. Berlin 1863: 618. 1864 ; I. histrix var. subinermis Durieu ex Motelay & Vendryès in Actes Soc. Linn. Bordeaux 36: 400. 1883 ; I. subinermis (Gennari) Cesca & Peruzzi in Fl. Medit. 11: 308. 2001. – Under the Code (McNeill et al. 2012: Art. 41.8(c)), the latter combination must be deemed as validly published in spite of the fact that the alleged basionym referred to (Isoetes histrix Bory forma subinermis Durieu in Bull. Soc. Bot. France 8: 164. 1861) is not validly published (being a nomen nudum). Protologue localities: monte Pisano [Pisa]. Lectotype (designated here): “Isoetes Hystrix var. subinermis, in pratis prope Pisas”, 30 April 1862, Beccari (B 20 0107700 ex herb. Braun!). – No original material has been traced in Gennari’s herbarium (TO), nor in other consulted Italian herbaria (FI, PI, PAL); however, the type here selected clearly belongs to the original gathering. ¼ Isoetes histrix [unranked] brevispina A. Braun in Monatsber. Königl. Preuss. Akad. Wiss. Berlin 1863: 618. 1864. Protologue localities: [Spain; Italy; Corsica; Sardinia] Guadarrama; Pisa; Bonifacio; Ingurtosu, Maddalena, Caprera. Lectotype (designated here): “Isoetes Hystrix Dur. f. desquamata b. caespitosa brevispina Al. Br. . . . Ingurtosu”, 11 Junee 1863, Reinhardt (B 20 0083454!). – The larger of the two plants on the sheet has megaspores of variable size, as is typical of hybrids. ¼ Isoetes sicula Tod. in Giorn. Sci. Nat. Econ. Palermo 1: 251. 1866 ; I. histrix subsp. sicula (Tod.) P. Fourn., Quatre Fl. France: 6. 1934. Protologue locality: [Sicily] “Ficuzza”. Neotype (designated here): “Isoetes sicula Tod., in humidis, Ficuzza al Lupo, martii”, Lojacono-Pojero (PAL 6044!). – No plants collected by Todaro have been traced, nor any other original material. The neotype specimen, collected in the ditio classica by one of Todaro’s pupils, has megaspores that show the peculiar, almost reticulate ornamentation of the distal face described in Todaro’s protologue. Note – The absence of a velum, reported in the protologue of Isoetes gymnocarpa, and Gennari’s (1862) claim that his plant is distinct from I. histrix var. subinermis (as described by Gennari in the same paper) have led some authors to accept I. gymnocarpa as different from the latter, to which they refer as I. subinermis (Cesca & Peruzzi 2001; Ernandes 2011) or I. sicula Tod. (Troia 2005; Troia et al. 2011; Troia in Greuter 2012). In view of the scant taxonomic importance of velum presence (see above), we now agree with Arrigoni (2005) and Carta et al. (2009), who circumscribe I. gymnocarpa as including I. sicula and I. subinermis. However, a better definition of its morphological features is desirable, as some unresolved questions remain. For example, Troia et al. (2012), for I. sicula, report megaspores that are smaller than those of I. histrix, in contrast with what has been observed by Cesca and Peruzzi (2001) for I. subinermis. (8) Isoetes durieui Bory in Compt. Rend. Hebd. Séances Acad. Sci. 18: 1166. 1844 ; Isoetella durieui (Bory) Gennari in Comm. Soc. Crittog. Ital. 1: 115. 1862. Protologue locality: [Algeria] “Bab Azoum”. Lectotype (Motelay & Vendryes 1883: 393): “Alger, coteaux de Bab-Azoum”, 1 April 1844, Durieu (BORD? [n.v.]; isolectotype: P 00202838 [image!]). Note 1 – The original spelling of the epithet, “duriei”, must be corrected to durieui under the Code (McNeill et al. 2012: Art. 60.7(a,1), contrary to the opinion of Pichi-Sermolli (in Ferrarini et al. 1986). Note 2 – Several different chromosome numbers (2n ¼ 44, 55, 110) have been reported for this species (by Cesca & Peruzzi 2001; Giovannini et al. 2001; Peruzzi et al. 2003, respectively), which seems to be an Downloaded by [Universita di Palermo] at 02:59 06 February 2014 Critical conspectus of Italian Isoetes allopolyploid that may have originated independently in different places (cf. Bolin et al. 2011). (9) Isoetes todaroana Troia & Raimondo in Amer. Fern. J. 99: 238. 2010 (‘2009’). Holotype (Troia & Raimondo 2010): “Sicily, contrada ‘Critazzo’ near Mazara del Vallo, ca. 60 m a.s.l.”, 10 April 2009, Angelo Troia (PAL 85592!; isotypes: PAL 85593!, FI 001648!). (10) Isoetes iapygia Ernandes et al. in Pl. Biosyst. 144: 807. 2010. Holotype (Ernandes et al. 2010): “Italy, Apulia, Macchie Don Cesare, Salve (Lecce) in temporary pools on calcareous substrate, 105 m a.s.l.”, 15 March 2008, Ernandes & Beccarisi (LEC) [n.v.]. Note – For the distinction between I. iapygia and I. todaroana, see Ernandes & Marchiori (2012). Although we had not the opportunity to see any herbarium specimen of I. iapygia, and we are not perfectly convinced by the conclusions of the work made by Ernandes and Marchiori (2012), we maintain here the two species as provisionaly separate. Key for the identification of the Italian taxa For a correct identification, it is important to study several plants with mature spores. Descriptive terminology of megaspores and microspores follows Hickey (1986) and Musselman (2003), respectively. For a comparison between “phyllopodia” and scales, see Bolin et al. (2008). 1 Aquatic plants, normally submerged throughout the year . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1* Plants not permanently submerged (only during winter and early spring) or terrestrial, with a leafless resting phase in summer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 2 Stem 3-lobed in cross section; leaves long (usually . 30 cm); sporangium not covered by a velum; megaspores baculate; plants in running water . . . . . . . . . . . . . . . I. malinverniana 2* Stem usually 2-lobed in cross section, leaves shorter (,20 cm); sporangium partially or totally covered by a velum; megaspores not baculate; plants in stagnant water ... ... ... ... ... ... ... ... ... ...3 3 Megaspores 360–480 mm in diameter, their proximal face with straight spines; velum partially covering the sporangium; plants of temperate climates . . . . . . . . . . . . . . . I. echinospora 3* Megaspores 545–645 mm in diameter, their proximal face with appressed structures; velum 7 (almost) totally covering the sporangium; plants of mediterranean climates . . . . . . . . . . . . I. sabatina 4 Plants with numerous hardened, persistent, black “phyllopodia” on top of the corm and around the leaf bases . . . . . . . . . . . . . . . . . . 5 4* Plants without phyllopodia (but sometimes with seasonal non-hardened “scales”) . . . 7 5 Phyllopodia with 3 – 10 mm long lateral spine-like teeth, without (or with reduced) central tooth . . . . . . . . . . . . . . . . . . I. histrix 5* Phyllopodia with three irregular small teeth of similar length . . . ... ... ... ... ...6 6 Megaspore . 600 mm in diameter, with alveolate ornamentation . . . . . . . . . . . . I. durieui 6* Megaspore , 500 mm in diameter, with tuberculate ornamentation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. gymnocarpa 7 Plants terrestrial; scales with two lateral, rounded lobes; leaf with two air chambers . . . . . . . . . . . . . . . . . . . . . ... ...8 7* Plants submerged during winter and early spring; scales absent or lacking rounded lobes; leaf with four air chambers . . . . . . . . . . . . . . . 9 8 Scales black; microspores ca. 26 (23–28) mm long; plants of seasonally wet meadows . . . . . . . . . . . . . . . . . . . . . I. todaroana 8* Scales brown; microspores ca. 30 (26 – 35) mm long; plants of rock pans or on thin moss layer . . . . . . . . . . . . . . . . . . . . . I. iapygia 9 Plants robust, stem usually . 1 cm thick; leaf bases with wide (1 – 2 mm) membranous margin, extending up leaf for 2 cm at least; microspores echinate or aculeate . . . . . . . . . . . . . . . . . . . . . I. longissima 9* Plants slender, stem usually # 1 cm thick; leaf bases with narrow (,1 mm) membranous margin, narrowing apically; microspores smooth or echinate–aculeate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. tiguliana Acknowledgements We are grateful to the staff of the consulted herbaria (B, BORD, FI, PAD, PAL, RO, TO) for assistance in our search for original material, and especially to Brigitte Zimmer (B), Chiara Nepi (FI), Laura Guglielmone (TO). Thanks are due to the Herbarium, Dipartimento di Biologia Ambientale, Sapienza Università di Roma (RO), for the image of the lectotype of Isoetes malinverniana. Funding This study is part of the “Flora Critica d’Italia” project and as such was funded by the Società Botanica Italiana onlus, the Fondazione per la Flora 8 A. Troia and W. Greuter Italiana, and the International Foundation Pro Herbario Mediterraneo. 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