The Italian taxa of the Ophrys crabronifera group

GIROS Notizie n. 53 – 2013
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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: [email protected]
2
Dipartimento di Studi Internazionali, Università di Urbino,
Piazza Rinascimento 7, 61029 Urbino - e-mail: [email protected]
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.
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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
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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
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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.
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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
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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
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Baumann H. & Lorenz R., 2005: Beiträge zur Taxonomie europäischer und mediterraner Orchideen.
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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).
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GIROS Notizie n. 53 – 2013
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