Abstract
The studied taxon has been attached since its discovery in Tunisia to Ophrys migoutiana H. Gay, a plant described in Algeria, which remains little known to botanists. This was already a net progress, ending the historical confusion that mixed many taxa under the name “O. fusca”. However, the recent observation of the true O. migoutiana near Medea in Algeria allowed us to distinguish the Algerian and the Tunisian taxa, the latter being a priori not yet formally described. The Tunisian taxon shares some similarities with oriental taxa of the Ophrys cinereophila / Ophrys creberrima group, and is named here as a new species Ophrys pseudomigoutiana. Its phenotypic variability is illustrated by several photographs and colour plates with fresh dissected flowers. Its chorology in Tunisia is detailed and suggests that it could also be present in eastern Algeria.
A taxonomic history of Ophrys fusca Link in Tunisia
Bonnet and Barratte (Citation1896) first mentioned “Ophrys fusca Link (syn. O. iricolor Desf.)” [sic!] in Tunisia, with a very long flowering season (“February–May”). Cuénod, Pottier-Alapetite, and Labbe (Citation1954) and Labbe (Citation1954) distinguished “Ophrys atlantica subsp. hayekii (Fleschm. et Soo) Maire et Weill.” and “Ophrys subfusca (Rchb.) Murb. (syn. O. funerea Batt.)” from O. fusca, but their illustration of “Ophrys fusca Link” actually represents Ophrys iricolor sensu lato. Vallès and Vallès-Lombard (Citation1988) renamed the main Tunisian O. fusca as “O. fusca subsp. iricolor (Desf.) O. Schwarz” with several distinct “forms”. Two of them, “fusca forme 1” and “fusca forme 2”, were later formally named, respectively as “Ophrys gazella Devillers-Tersch. et Devillers” (Devillers and Devillers-Terschuren Citation2000) and “Ophrys vallesiana Devillers-Tersch. & Devillers” (Devillers and Devillers-Terschuren Citation1994), whereas another was identified as “Ophrys migoutiana Gay” by Devillers and Devillers-Terschuren (Citation1994). Amich et al. (Citation2009) made some clarifications on the taxonomy of the main Tunisian species, but did not study O. migoutiana. Le Floc’h, Boulos, and Véla (Citation2010) and Martin et al. (Citation2013) maintained the recognition of O. migoutiana as a Tunisian species, whereas O. fusca sensu stricto was definitely excluded from Tunisian flora, because the type is from Portugal and the nearest similar plants are known from the surroundings of Skikda in Algeria (De Bélair et al., Citation2005). For the very first time, we proposed (in Martin, Ouni, and Véla Citation2015) the distinction between the true and the “false” O. migoutiana, and suggested the epithet “pseudomigoutiana” hereafter used for this new species (Table ).
Table 1. Historical taxonomy of Pseudophrys species in Tunisia.
The true Ophrys migoutiana and its affinities
The recent rediscovery of the O. migoutiana in its locus typicus near Médéa (Gay Citation1889) in central-northern Algeria (Rebbas and Véla Citation2013; Kreutz et al. Citation2013), 500 km west of the Tunisian border, provided new elements for a better characterization of this taxon originally typified in Algeria, and for a re-evaluation of the widely distributed Tunisian populations called so far O. migoutiana sensu Devillers & Devillers-Terschuren, non Gay.
Ophrys migoutiana s.s. showed unexpected morphological affinities (i) with the Aegean group formed by Ophrys blitopertha H.F. Paulus, pollinated by a coleopteran from the genus Blitopertha Reitter, 1903, and Ophrys persephonae H.F. Paulus, the pollinator of which is unknown (Rebbas and Véla Citation2013), (ii) as well as with Ophrys laurensis Geniez & Melki from Sicily (Melki and Geniez Citation1992), pollinated by the Hymenoptera Andrena schulzi Strand, 1921 (Delforge Citation2001a). Until now, pollinators of the Algerian O. migoutiana s.s. still remain unknown. These four species share long petals, the latter around 80% of the sepals’ length, and an excavated labellum base. Within sect. Pseudophrys Godfery, this combination of characters excludes them from both the O. fusca / Ophrys lutea Cav. aggregate (species of this group have short petals) and the Ophrys omegaifera Desf. / Ophrys atlantica Munby aggregate (characterized by a flat labellum base), so that we have placed them in a separate group (Martin, Ouni, and Véla Citation2015).
In the genus Ophrys, “divergent selection by pollinators can bring about strong reproductive isolation via changes at few genes of large effect” (Xu and Schlüter Citation2015), a pre-zygotic isolation mainly driven by floral chemical fragrance (Xu et al. Citation2011). Hence, coloration, shape and size of labellum are the most variable characters within a species, and are the first to derivate in the case of speciation by a pollinator shift (Schlüter et al. Citation2007, Citation2009, Citation2011). At a higher taxonomic level, form and length of petals on the one hand, architecture of stigmatic cavity and basis of labellum on the other hand, are good criteria to identify species groups, in other words “macro-species”, within the “sphegodes complex”, i.e. section Araniferae (Devillers and Devillers-Terschuren Citation1994; Véla Citation2007). Petal length and labellum basis were also successfully used for identifying the “omegaifera complex” (Devillers and Devillers-Terschuren Citation1994). Curiously, these criteria are not used by the same authors and followers (Devillers and Devillers-Terschuren Citation1994; Delforge Citation2001b) nor by authors with independent conceptions (e.g. Kreutz Citation2004; Faurholdt and Pedersen Citation2009) for identifying groups / macro-species within the “lutea-fusca-iricolor complex”. As a result, putative groups / macro-species are strongly unrealistic and unusable, if not incomprehensible. Our analyses (Rebbas and Véla Citation2013; Tison, Jauzein, and Michaud Citation2014; Tison and De Foucault Citation2014; Martin, Ouni, and Véla Citation2015) based on a morphological approach coherent with previous information, results in “morpho-species” completely different from the grouping of Devillers and Delforge. In this sense, length and form of petals combined with architecture of labellum basis allow us to separate easily O. migoutiana and allies (O. blitopertha, O. laurensis, etc.) from the “fusca-”, “subfusca-” and “lutea” groups.
Affinities of the Tunisian taxon
In the central-western Mediterranean context, the Tunisian “false migoutiana” shows only distant affinities with the true O. migoutiana from Algeria, O. laurensis from Sicily and to a lesser extent with the western Mediterranean O. funerea / O. obaesa aggregate (cf. Martin, Ouni, and Véla Citation2015). However, in a broader perspective, the Tunisian “false migoutiana” shows strong morphological affinities (long petals, excavated labellum basis, labellum convexity) with the Aegean Ophrys cinereophila H.F. Paulus & Gack / Ophrys creberrima H.F. Paulus, two closely related species respectively pollinated by Andrena cinereophila Warncke, 1965 and Andrena creberrima Pérez, 1895 (cf. Paulus Citation1998). Here again, the pollinator of the Tunisian “false migoutiana” remains unknown.
Pseudophrys species with long petals from central Mediterranean
In Tunisia and the central Mediterranean area, five Pseudophrys species have long petals (> 75% of sepal length) with margins often undulate (Table , and see Supplementary Data S1), if we do not consider the O. funerea Viv. aggregate with relatively less long petals (between 60 and 75% of sepals) not undulate (Martin, Ouni, and Véla. Citation2015):
| (1) | Ophrys atlantica, with a diagnostic labellum concavity, is very localized in central-western Tunisia, but extends westwards until southern Spain. | ||||
| (2) | Ophrys omegaifera subsp. hayekii (H.Fleischm. & Soó) Kreutz [= O. mirabilis], a regional vicariant belonging to O. omegaifera s.l. with a flat labellum basis and a more or less convex labellum, is localized in northern Tunisia, but extends eastwards to Sicily and westwards to Algeria. | ||||
| (3) | Ophrys laurensis: this remarkable endemic species from southeastern Sicily has got a characteristic and diagnostic very large and deep excavation at labellum basis. | ||||
| (4) | Ophrys migoutiana: although the locus typicus is the only one locality known in Algeria (localized in the central north of the country); another locality with big flowers and a relatively flat labellum is known in central-western Tunisia and is assigned the name “O. cf. migoutiana (?)”. | ||||
| (5) | Ophrys “pseudomigoutiana” nom. provis.: widespread throughout northern Tunisia, this species has a diagnostic small labellum (as O. laurensis) combined with a weakly excavated labellum basis (as O. migoutiana) and a global labellum convexity (Figure 1). | ||||
Table 2. Comparative morphology of Ophrys pseudomigoutiana and other Pseudophrys with long petals from Central Mediterranean area.
Ophrys pseudomigoutiana, a name for the Tunisian taxa
Name. Ophrys pseudomigoutiana R.Martin, Véla & Ouni, sp. nov.
Figure 1. Isotype of Ophrys pseudomigoutiana, R. Martin, Aïn Jemala (Governorate of Beja), 27 March 2008.
Etymology. “pseudo” = false, and “migoutiana” = dedicated to Mr Abel Migout (born 1830, high school teacher and author of Flore du département de l'Allier et des cantons voisins in 1890), because the species looks like O. migoutiana H. Gay with which it was confused for 20 years.
Diagnosis. Ophrys sect. Pseudophrys with petals very long (± 80% as sepals). It differs from O. migoutiana H. Gay by shorter sepals, shorter petals and above all a shorter labellum (11.5–15 versus 15–17.5 mm), more or less convex, with a thinner yellow border (0.6 –1.2 versus 1.5–2 mm). Its labellum differs from that of O. laurensis Geniez & Melki in having a less deeply excavated base and a thinner yellow border.
Type specimen (see Supplementary Data S2). E. Véla n°0010 [holotype AIX (upper right flower); isotypes AIX (same sheet, all other flowers)], Tunisia, Aïn Jemala (governorate of Beja), legit R. Martin 27 March 2008.
Other specimen (paratypes): E. Véla n°0011 [MPU (all flower except the last down)], Tunisia, Aïn Jemala (Governorate of Beja), leg. R. Martin 19 March 2010; E. Véla n°0012, (MPU), Tunisia, Bejaoua (Governorate of La Manouba), legit R. Ouni 29 April 2013; E. Véla n°0013 [MPU (all flower except the last down)], Tunisia, Jebel Sidi Zit (Governorate of Zaghouan), leg. R. Martin 13 March 2008; E. Véla n°0014 [MPU (all flower except the last down)], Tunisia, Oued Zarga PK 14,5 (Governorate of Beja), leg. R. Martin 21 March 2008; E. Véla n°0015 [MPU (all flower except right up and left down)], Tunisia, Jebel Amar – Jebbes (Governorate of Ariana), leg. R. Martin 25 March 2008; E. Véla n°0016 [MPU (flower at left)], Tunisia, Le Krib – La Chapelle (Governorate of Siliana), leg. R. Martin 27 March 2008; E. Véla n°0017 (MPU [all flowers]), Tunisia, Kesra I (Governorate of Siliana), leg. R. Martin 03 April 2008.
Ecology. Flowering from March to April, growing under Aleppo Pine (Pinus halepensis Mill.) in open forests. Pollinators still unknown.
Chorology. Throughout northern Tunisia, especially the Central Dorsal Mountains, from Cap Bon to the Algerian border (Figure ). It is expected in eastern Algeria, in the surroundings of Souk-Ahras and Tebessa.
Figure 2. Distribution map of Ophrys pseudomigoutiana in Tunisia (modified from Martin, Ouni, and Véla Citation2015). Its presence has not yet been reported in Algeria.
Notes on contributors
Errol Vela is an assistant professor at the University of Montpellier, where he teaches environmental expertise and botany. He was a naturalist expert in France for 5 years, and he is now a scientific advisor at an international level. In addition to his researches on biogeography and phylogeography, he is interested in taxonomy and flora in France, North Africa and the Near East. Contribution: Errol Vela initiated the taxonomic work, has delimited the taxon presented here and has written the scientific argumentation.
Roland Martin, a retired and amateur naturalist, has been responsible for the mapping of orchids in Vaucluse (France) for over 20 years. President of the Mediterranean Society of Orchidology, he is the author of several articles and books on wild orchids from France and Tunisia. Born in Kef, Tunisia, he began in 2000 to update the inventory of orchids from Tunisia and initiate a never made work: their mapping. Contribution: Roland Martin made the main mapping inventories in the field, photographs and specimen preparation.
Ridha Ouni is a self-educated naturalist and regular contributor to Tunisian universities and international organizations. He is also co-founder of the Tunisian Wildlife Society, and he is an international consultant in the fields of ornithology, herpetology, entomology, etc. Contribution: Ridha Ouni participated in numerous prospective field work and continues to increase the database.
Supplemental data
Supplementary Data S1. Flowers of the five Pseudophrys spp. with long petals from the Central Mediterranean area. Upper line, from left to right: Ophrys atlantica from Tunisia (Kalaat Esnan, 04.05.2005), Ophrys omegaifera subsp. hayekii from Tunisia (Nefza, 21.04.2008), Ophrys laurensis from Sicily (Monte Lauro, 18.04.2011); middle line, Ophrys migoutiana s.s. from Algeria (Ouzera, 28.03.2011), Ophrys cf. migoutiana from Tunisia (Ksar el Guellat, 20.04.2005), Ophrys pseudomigoutiana from Tunisia (Aïn Jemala, 27.03.2008); bottom line, examples of morphological variation in O. pseudomigoutiana from three localities: Oued Mellegue, 12.04.2005 (left), Oued Zarga, 23.03.2008(center) Zaghouan 21.03.2006 (right). All photographs by R. Martin except O. migoutiana s.s. by E. Véla.
Supplementary Data S2. Fresh material illustration of type specimen deposited in AIX, leg. R.Martin, Aïn Jemala (governorate of Beja), 27.03.2008, the first flower up and right is the holotype, other flowers of the plate are isotypes. Scale: please note that millimetre scale was reduced at 95% on paper; it means unit cells of the grid are 950 µm and not 1 mm.
1099468_Supplementary_data.zip
Download Zip (17.1 MB)Acknowledgements
Field trips in Tunisia were partially realized with the financial support of the “Fondation Eco-Med” (Marseille, France) to the Société Méditerranéenne d’Orchidologie. Field trips in Algeria benefited from the logistic assistance of Khellaf Rebbas (University of M’Sila). The authors are grateful to Jean-François Molino for his pertinent comments, to Sophie Gachet for her help in language corrections, and to the editorial board and an anonymous reviewer.
Related Research Data
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