GIROS Notizie n. 53 – 2013 1 by C.A.J. Kreutz1 & J.M.I. Klaver2 Naturalis Biodiversity Center (section NHN), Biosystematics group, Wageningen University, Gen. Foulkesweg 37, 6703 BL Wageningen, The Netherlands - e-mail: c.kreutz@hccnet.nl 2 Dipartimento di Studi Internazionali, Università di Urbino, Piazza Rinascimento 7, 61029 Urbino - e-mail: jmi.klaver@uniurb.it Riassunto: gli autori discutono il concetto tassonomico usato da Pedersen & Faurholdt per il gruppo Ophrys argolica e propongono di separare da questa specie i taxa italiani e inserirli come sottospecie di O. crabronifera. Nel 2002 Pedersen & Faurholdt, in preparazione del loro libro sulle Ophrys d’Europa, hanno riordinato i taxa fino ad allora conosciuti del gruppo Ophrys argolica-crabronifera come sottospecie di Ophrys argolica. Nella loro definizione una specie è costituita da un gruppo di individui che, in condizioni naturali (reali o potenziali) possono incrociarsi per produrre discendenti vitali e fertili, e gli individui di una specie si distinguono da quelli di altre specie attraverso le loro caratteristiche morfologiche. Applicando questi criteri, distinguono quattro entità per il Mediterraneo orientale (O. argolica subsp. argolica, O. argolica subsp. lucis, O. argolica subsp. aegaea, O. argolica subsp. lesbis) e due per l’Italia (O. argolica subsp. crabronifera e O. argolica subsp. biscutella). Inquadrano Ophrys morisii nel complesso O. ×arachnitiformis, e identificano Ophrys pollinensis con O. argolica subsp. crabronifera H.Baumann & R. Lorenz, tuttavia mantengono O. crabronifera a livello di specie per i taxa italiani, di cui O. morisii e O. pollinensis sono inserite a livello di sottospecie di O. crabronifera, mentre O. biscutella è per loro un sinonimo di O. crabronifera subsp. pollinensis. In Orchidee d’Italia (GIROS 2009) gli autori riconoscono le combinazioni proposte da Pedersen & Faurholdt per O. crabronifera e O. biscutella come sottospecie di O. argolica, seguono la classificazione di Kreutz per O. argolica subsp. pollinensis, ma mantengono la proposta di Del Prete per O. exaltata subsp. morisii. Romolini & Souche attribuiscono il rango di specie a O. crabronifera, O. biscutella, e O. pollinensis, ma conservano O. morisii come sottospecie di O. exaltata. Pedersen & Faurholdt non concordano con la tendenza prevalente nella classificazione di Ophrys che ignora i ranghi infraspecifici e attribuisce il rango di specie anche a piccole popolazioni che sono morfologicamente solo leggermente diverse da popolazioni affini. Sottolineano che questa pratica tende a ignorare i diversi gradi di diversità nelle caratteristiche morfologiche, filogenetiche e geografiche. Anche se la riflessione di Pedersen & Faurholdt è condivisibile, noi crediamo, tuttavia, che nella loro classificazione siano andati all’estremo opposto, applicando un concetto di specie che è troppo limitato per essere utile in un tentativo di mantenere un concetto tassonomico chiaro. Organizzare tutti i taxa del gruppo Ophrys argolica-crabronifera presenti nell’Egeo e in Italia come sottospecie di Ophrys argolica aiuta sì a esprimere la sua probabile origine monofiletica, ma oscura le relazioni interne ai suoi rappresentanti orientali e occidentali. A questo proposito è istruttivo notare che proprio nella loro discussione dei taxa italiani del gruppo O. argolica Pedersen & Faurholdt percepiscono i limiti del loro concetto di specie e di sottospecie quando ammettono che non pretendono di essere stati in grado di affrontare tutte le numerose sfide in modo soddisfacente. Pertanto, in analogia con la distinzione tra O. oestrifera e O. scolopax proponiamo di mantenere O. argolica per i taxa del Mediterraneo orientale, e utilizzare O. crabronifera per i taxa italiani. Questa decisione si basa sia sulle affinità morfologiche che sulla distribuzione geografica dei taxa in questo gruppo, e prende anche in considerazione l’instabilità genetica di molti taxa alla periferia del gruppo O. argolica da una parte e il gruppo O. crabronifera dall’altra, questa essendo spesso il risultato di ibridazione e di reincrocio con altre specie. La nostra decisione tassonomica richiede due nuove combinazioni di taxa italiani: Ophrys crabronifera Mauri subsp. biscutella (O. Danesch & E. Danesch) J.M.I. Klaver & Kreutz comb. et stat. 1 Articoli The Italian taxa of the Ophrys crabronifera group Articoli GIROS Notizie n. 53 – 2013 nov.; Ophrys crabronifera Mauri subsp. virescens (Sommier) J.M.I. Klaver & Kreutz, comb. et stat. nov. Parole chiave: Orchidaceae, Ophrys crabronifera, Ophrys crabronifera subsp. virescens (Sommier) J.M.I. Klaver & Kreutz, comb. et stat. nov., Ophrys crabronifera subsp. biscutella (O. Danesch & E. Danesch) J.M.I. Klaver & Kreutz , comb. et stat. nov., Ophrys crabronifera subsp. pollinensis, Ophrys crabronifera subsp. morisii, Ophrys argolica, Italia. Abstract: the Authors argue for a rearrangement of the taxa of the Ophrys argolica group and propose to keep O. argolica for the eastern Mediterranean taxa, and to use O. crabronifera for the Italian taxa. This necessitates new combinations of two scientific names. Keywords: Orchidaceae, Ophrys crabronifera, Ophrys crabronifera subsp. virescens (Sommier) J.M.I. Klaver & Kreutz, comb. et stat. nov., Ophrys crabronifera subsp. biscutella (O. Danesch & E. Danesch) J.M.I. Klaver & Kreutz , comb. et stat. nov., Ophrys crabronifera subsp. pollinensis, Ophrys crabronifera subsp. morisii, Ophrys argolica, Italy. INTRODUCTION In 2002 Pedersen & Faurholdt, in preparation of their 2007 book on the bee orchids of Europe, revised the Ophrys argolica–crabronifera group, rearranging all the taxa which were then known from the central and east Mediterranean as subspecies under Ophrys argolica H. Fleischm. Considering a species as “all individuals that under natural conditions (in reality or potentially) can interbreed to produce consistently viable and fully fertile offspring,” and stressing the fact that “the individuals of one species are distinguished from those of other species by morphological features”, Pedersen & Faurholdt (2007, 56) distinguish four east-Mediterranean entities (O. argolica subsp. argolica, O. argolica subsp. lucis, O. argolica subsp. aegaea, O. argolica subsp. lesbis) and two Italian taxa (O. argolica subsp. crabronifera and O. argolica subsp. biscutella). They do not recognize the validity of O. icariensis from Ikaria and Naxos, which they see as hybrids with O. ferrum-equinum as one of its parents; furthermore they attribute the Corsican and Sardinian Ophrys morisii (Martelli) Soó to O. ×arachnitiformis. Ophrys pollinensis E. Nelson ex Devillers-Tersch. & Devillers is identified by them as a synonym of O. argolica subsp. crabronifera (Pedersen & Faurholdt 2007, 155-166). Baumann & Lorenz (2005, 719), however, keep O. crabronifera at species level for the Italian taxa, O. morisii and O. pollinensis at subspecies level and Ophrys biscutella O. Danesch & E. Danesch as synonymous with O. crabronifera subsp. pollinensis. GIROS (2009: 195-197, 230) recognizes the proposed combinations in Pedersen & Faurholdt (2002; 2007) for O. crabronifera and O. biscutella as subspecies of O. argolica, follows Kreutz (2004, 85-86) for O. argolica subsp. pollinensis, but maintains Del Prete (1984, 251) for O. exaltata subsp. morisii. Romolini & Souche (2012) distinguish O. crabronifera, O. biscutella, and O. pollinensis all at species level, but relegate O. morisii to subspecies status under O. exaltata. DISCUSSION Pedersen & Faurholdt (2002; 2007) argue against the prevailing tendency in 2 GIROS Notizie n. 53 – 2013 THE ITALIAN TAXA The following taxa in the Italian O. crabronifera group need to be distinguished: 1 – Ophrys crabronifera Mauri subsp. crabronifera This is the nominate species. It was first described by Ernesto Mauri in 1820 (42), based on plants from the province of Rome. Plants are typically 25 to 55 cm tall. Sepals whitish, pink or violet, strongly reflexed in flowers which are well-open. Labellum convex, longer than wide and trapezoid, reddish-brown often tending towards green at the base, protuberances absent or near absent, pseudo-eyes greenish, speculum generally reduced to two isolated eye-like parts on the lower half of lip. Ophrys crabronifera is endemic in central Italy. It is not uncommon in Tuscany south of Livorno and in the province of Grosseto, while it is one of the rarest orchids in the 3 Articoli Ophrys classification to disregard infraspecific ranks and conveying the rank of species even to small populations which are morphologically only slightly different from related populations. They underline that this practice has tended above all to efface the different degrees of the morphological, phylogenetic, and isolating characteristics of Ophrys taxa. Although Pedersen & Faurholdt’s (l.c.) reflection on the species question in Ophrys is definitely in place, we believe, however, that in their classification they have swung the pendulum to the other extreme by applying a species concept which is too limited to be helpful in an attempt to maintain a clear taxonomical concept. Arranging the Aegean and Italian taxa of the group under discussion here as subspecies of Ophrys argolica helps to express its probable monophyletic origin (cf. Delforge 2005, 536-538, 544-545) but obscures the internal relationships within its eastern and within its western representatives. In this respect it is instructive to note that it is in their discussion of the Italian taxa of the O. argolica group that Pedersen & Faurholdt (2002; 2007) sense the limitations of their species and subspecies concept when they admit that “we do not pretend that we have been able to deal with all of the many challenges in a satisfactory way” (Pedersen & Faurholdt 2007, 59). Therefore, in analogy with the distinction between O. oestrifera and O. scolopax (Kreutz 2004, 108-110) we propose to keep O. argolica for the eastern Mediterranean taxa, and use O. crabronifera for the Italian taxa. This decision rests on both the morphological affinities and geographical distribution of the taxa in this group. It also takes into account the genetic instability of many taxa at the periphery of the O. argolica group on one hand and of the O. crabronifera group on the other, this being often the result of hybridization and backcrossing with other species (cf. Delforge 2005; Pedersen & Faurholdt 2007). Our taxonomical decision necessitates two new combinations of the Italian taxa: Ophrys crabronifera Mauri subsp. biscutella (O. Danesch & E. Danesch) J.M.I. Klaver & Kreutz comb. et stat. nov.; Ophrys crabronifera Mauri subsp. virescens (Sommier) J.M.I. Klaver & Kreutz, comb. et stat. nov. Articoli GIROS Notizie n. 53 – 2013 Province of Pisa (Frigani 2011, 98, 163). In the confining regions Lazio, Marche, Umbria, Molise and Abruzzo it is not common. It has also been reported for the north of Campania, where it is rare. The species is often erroneously held to be endemic in a wide area along the Tyrrhenian coast (Del Prete & Tosi 1988, 34; Pedersen & Faurholdt 2007, 158), but Delforge (2005, 533) is correct in indicating a much larger area on mainland Italy stretching eastward into the Marche as far as the surroundings of Urbino (cf. Klaver 1989). Recent finds here include numerous inland locations in the Piobbico area, and a series of occurrences near Fossombrone, a mere 25 km from the Adriatic coast. 2 – Ophrys crabronifera Mauri subsp. virescens (Sommier) J.M.I. Klaver & Kreutz, comb. et stat. nov. Bas.: Ophrys exaltata Tenore var. virescens Sommier, L’Isola del Giglio e la sua flora: 77 (1900). This entity was described in 1900 by the Italian botanist Stéphen (Pietro Stefano) Sommier as Ophrys crabronifera var. virescens. His plants were collected 10 March 1897 from a population on the western promontory “Il Franco” on the island of Giglio in the Tuscan archipelago. He later reported the variety also on the island of Pianosa (cf. Baldini 1998, 391). The parsimonious Latin description he provided runs as follows: “Perigonii phyllis tribus exterioris laete, duobus interioribus sordid viridibus” (Sommier 1900, 77). In an unpublished note, preserved with the holotype (FI002752), Sommier however specifies: “Tepali esterni verde sbiadita […]. Interni marroni verdastri. Labello tutto nero peloso con delle piccolissime macchie lucide”, drawing attention to its green sepals and black labellum. This subspecies tends to produce slender plants with up to 6 flowers featuring bright green sepals and slightly darker petals, both resembling in colour those of O. sphegodes. The colour of the labellum is not as brightly reddish as in the nominate species, and tends to be dark or blackish brown, as in O. incubacea, and generally is not green at the base. The shape of the lip is proportionally less elongated and of more rounded aspect than in Ophrys crabronifera subsp. crabronifera; the pseudo eyes are dark, often black. As in the nominate species, the speculum is limited to the lower part of the labellum where it generally is made up of two dark violet eye-like drops. O. crabronifera subsp. virescens clearly is an endemic entity originating on the Tuscan archipelago (possibly Pianosa) where its geographically isolated status helps to keep it separate from the nominate species. Where, however, the areas of Ophrys crabronifera subsp. crabronifera and Ophrys crabronifera subsp. virescens overlap intermediate forms can be observed (Giglio, Argentario), often with typically luridly green-reddish sepals. 3 – Ophrys crabronifera Mauri subsp. biscutella (O. Danesch & E. Danesch) J.M.I. Klaver & Kreutz , comb. et stat. nov. Bas.: Ophrys biscutella O. Danesch & E. Danesch, Die Orchidee 21 (6): 358 (1970). This entity is restricted to the Gargano peninsula in Puglia. It seems closer to the 4 GIROS Notizie n. 53 – 2013 4 – Ophrys crabronifera Mauri subsp. pollinensis (E. Nelson ex O. Danesch & E. Danesch) H. Baumann & R. Lorenz Ophrys crabronifera subsp. pollinensis was originally described from plants on Mount Pollino. It is endemic for the south of Italy in an area that stretches from the south of Campania to the north of Calabria, including the Lattari Mountains, the Picentini Mountains, the Cilento, Lucania and the Pollino Mountains. This subspecies is a clearly less stable entity. A bewildering mixture of variety can be encountered. Lip shape is generally rounded rather than trapezoid, and is of a dark brownish colour. The speculum is generally much more developed than that of the nominate species and often connected to the base of the lip. The appendix is usually wellpronounced. In an early study of chromosome numbers in Ophrys Greilhuber & Ehrendorfer (1975, 135) drew attention to the chromosomically (2n=36) and morphologically stable nature of Ophrys crabronifera subsp. biscutella, while they found in Ophrys crabronifera subsp. pollinensis chromosome numbers oscillating between 2n=36 and 2n=38 (Greilhuber & Ehrendorfer l.c.: 133). Delforge (2005, 545) maintains that Ophrys crabronifera subsp. pollinensis is very likely of recent hybrid origin. Pedersen & Faurholdt (2007, 158) also write that “[p]articularly in Campania, a most confusing multitude of forms can be observed”. They believe that “extensive hybridization and backcrossing [with O. fuciflora subsp. fuciflora]” is taking place here (l.c., 59). Romolini & Souche (2012: 330), however, argue that these forms in Campania are intermediate between O. crabronifera and O. biscutella. 5 – Ophrys crabronifera Mauri subsp. morisii (Martelli) H. Baumann & R. Lorenz This entity is endemic for Corsica and Sardinia. Just like the subspecies above, Ophrys crabronifera subsp. morisii is a rather unstable form, which we believe to be of recent hybrid origin (cf. Delforge 2005: 545). Its morphological characters lead to inclusion in the O. crabronifera group rather than the O. exaltata Ten. group (cf. Kreutz 2004, 86; Baumann & Lorenz 2005, 719; Delforge 2012, 248), but its geographical isolation separates it well from the nominate species, from which it also differs in its pollinator: Anthophora sicheli (Paulus 2007: 278). BIBLIOGRAPHY Baldini R.M., 1998: Flora vascolare dell’isola Giglio (Arcipelago toscano): revisione tassonomica ed aggiornamento. ˗ Webbia 52(2): 307-404. Baumann H. & Lorenz R., 2005: Beiträge zur Taxonomie europäischer und mediterraner Orchideen. 5 Articoli Aegean O. argolica group than the other Italian entities and distinguishes itself from the nominate species by having a more rounded lip-shape, a smaller appendix, a stigmatic cave with narrower base, and a speculum which typically features a pair of spectacles rather than isolated drops or eye-like spots. While as pollinator Anthophora plumipes has been indicated for the nominate species, Anthophora retusa has been reported for subsp. biscutella (Paulus 2007, 278). Articoli GIROS Notizie n. 53 – 2013 ˗ J. Eur. Orch. 37(3): 705-743. Del Prete C. & Tosi G., 1988: Orchidee spontanee d’Italia: monografia e iconografia. Mursia, Milano. Del Prete C., 1984: The Genus Ophrys L. (Orchidaceae) in Italy: Checklist of the Species, Subspecies and Hybrids. ˗ Webbia 37: 247-255. Delforge P., 2005: Guide des orchidées d’Europe, d’Afrique du Nord et du Proche-orient. Delachaux et Niestlé, Paris. Delforge P., 2012: Guide des orchidées de France, de Suisse et du Benelux. Delachaux et Niestlé, Paris. Frigani F., 2011: Atlante delle Orchidee della Provincia di Siena. Edizioni Cantagalli, Siena. GIROS, 2009: Orchidee d’Italia. Guida alle orchidee spontanee. Il Castello, Cornaredo (MI). Greilhuber J. & Ehrendorfer F., 1975: Chromosome Numbers and Evolution in Ophrys (Orchidaceae). ˗ Plant Syst. Evol. 124: 125-138. Klaver J.M.I., 1989: Ophrys crabronifera Mauri in der italienischen Marche (mittleres Ostitalien) ˗ Die Orchidee 40(2): 55-56. Kreutz C.A.J., 2004: Kompendium der Europäischen Orchideen / Catalogue of European Orchids. Kreutz Publishers, Landgraaf. Mauri E., 1820: Romanarum Plantarum Centuria XIII. Typis de Romanis, Roma. Paulus H.F., 2007: Wie Insekten-Männchen von Orchideenblüten getäuscht werden – Bestäubungstricks und Evolution in der mediterranen Ragwurzgattung Ophrys. – Denisia 20: 255-294. Pedersen H.A. & Faurholdt N., 2002: Ophrys – Versuchsweise Definitionen der Kategorien Art, Unterart und Varietät in der Gattung und einige daraus resultierende taxonomische Änderungen. ˗ Die Orchidee 53(3): 341-346. Pedersen H.A. & Faurholdt N., 2007: Ophrys: The Bee Orchids of Europe. Kew Publishing, Royal Botanic Gardens, Kew. Romolini R. & Souche R., 2012: Ophrys d’Italia. Editions sococor, Saint-Martin-de-Londres. Sommier S., 1900: L’Isola del Giglio e la sua flora: con note geologiche. Clausen, Torino. 6
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