Phelipanche cingularum (Orobanchaceae), a new species from southern France

Abstract

A new species of Phelipanche has been discovered in various steep places in southern France at meso-and supra-mediterranean levels. It is always parasitic on Brassicaceae, almost exclusively on Hesperis laciniata, a plant that had never been reported as host for any Orobanchaceae previously. The new species, named Phelipanche cingularum for its rocky ledge (cingula) habitat, is described and illustrated. Diagnostic characters against up to 16 presumably related species are given. We also present the results of molecular analyses well-supporting its independence. The distribution and the unusual ecology of the plant are considered and illustrated, and its conservation status is evaluated.

Full text

Phytotaxa 653 (1): 001–019
https://www.mapress.com/pt/
Copyright © 2024 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Gianniantonio Domina: 15 May 2024; published: 13 Jun. 2024
https://doi.org/10.11646/phytotaxa.653.1.1
1
Phelipanche cingularum (Orobanchaceae), a new species from southern France
THOMAS CROZE1, LUIS CARLÓN RUIZ2, JEAN-MARC TISON3, HENRI MICHAUD4, JAMES MOLINA4 &
GONZALO MORENO MORAL5
1Naturalia environnement, site Agroparc, 60, rue Jean Dausset, FR-84911 Avignon cedex 9 France
�
t.croze@naturalia-environnement.fr; https://orcid.org/0009-0001-2471-2032
2Jardín Botánico Atlántico, Avda. Jardín Botánico, 2230; 33203 Gijón-Xixón (Asturias) España
�
carlonluis@uniovi.es; https://orcid.org/0000-0003-3442-8710
34 Promenade du Decumanus, FR-38080 L’Isle d’Abeau France
�
jmltison@gmail.com; https://orcid.org/0009-0003-2801-2142
4Conservatoire Botanique National Méditerranéen de Porquerolles, avenue Gambetta, FR-83400 Hyères France
�
h.michaud@cbnmed.fr; https://orcid.org/0000-0001-7835-7366
�
j.molina@cbnmed.fr; https://orcid.org/0009-0004-3049-5267
5Santa Clara, 9-1 dcha., E-39001 Santander (Cantabria) España
�
gmorenomor55@gmail.com; https://orcid.org/0009-0001-7839-6506
Abstract
A new species of Phelipanche has been discovered in various steep places in southern France at meso- and supra-mediterranean
levels. It is always parasitic on Brassicaceae, almost exclusively on Hesperis laciniata, a plant that had never been reported
as host for any Orobanchaceae previously. The new species, named Phelipanche cingularum for its rocky ledge (cingula)
habitat, is described and illustrated. Diagnostic characters against up to 16 presumably related species are given. We also
present the results of molecular analyses well-supporting its independence. The distribution and the unusual ecology of the
plant are considered and illustrated, and its conservation status is evaluated.
Key words: Cévennes, conservation, Liguro-Provençal Pre-alps, Mediterranean, molecular phylogeny, Provence,
taxonomy
Résumé
Une nouvelle espèce de Phelipanche a été découverte en divers lieux escarpés du Midi de la France, aux étages méso-
et supra-méditerranéens. Elle parasite uniquement des espèces de la famille des Brassicaceae, et presque exclusivement
Hesperis laciniata (Brassicaceae), une plante qui n’avait encore jamais été signalée comme hôte d’une Orobanchaceae.
Cette espèce nouvelle, nommée ici Phelipanche cingularum en lien avec son habitat de vire rocheuse (cingula), est décrite et
illustrée. Des caractères diagnostiques concernant jusqu’à 16 espèces présumées apparentées sont donnés en comparaison.
Sont également exposés les résultats d’analyses moléculaires qui soutiennent nettement l’indépendance de ce taxon vis-à-
vis de ses congénères. La répartition, mais aussi l’écologie singulière de cette plante sont abordées et illustrées, et enfin son
statut de conservation est évalué.
Mots-clés: Cévennes, conservation, Préalpes Liguro-Provençales, Méditerranée, phylogénie moléculaire, Provence,
taxonomie
Introduction
The genus Phelipanche Pomel (1874: 102), often treated as Orobanche sect. Trionychon Wallr. (1822: 314), has been
very poorly known for as long as Orobanchaceae were studied in herbaria. Most authors of the second half of the XXth
century cited only 4 species in France, O. arenaria Borkh. (1794: 6) [= Ph. arenaria (Borkh.) Pomel (1874: 103)],
O. lavandulacea Rchb. (1829: 48) [= Ph. lavandulacea (Rchb.) Pomel (1874: 106)], O. purpurea Jacq. (1762: 252)
[= Ph. purpurea (Jacq.) Soják (1972: 130)] and O. ramosa L. (1753: 633) [= Ph. ramosa (L.) Pomel (1874: 103)], and

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less than 10 in Europe (see for instance Guinochet & Vilmorin 1975, Kreutz 1995, Tutin et al. 1972). Older French
authors having a best view on living plants also retained O. mutelii F.W. Schultz (1835: 353) and O. nana Noë ex Rchb.
(1842: n.° 1352) (see for instance Fournier 1940). Since the extreme end of the XXth century, new studies based on
the careful examination of living plants and hosts, lots of good photographs, and finally molecular data, allowed to
considerably increase the number of species, to 9–10 in France but excluding Ph. mutelii (F.W. Schultz) Pomel (1874:
106), and much more in southernmost European countries, most of the new taxa being formerly confused with Ph.
ramosa (Carlón et al. 2005, Carlón et al. 2008, Sánchez Pedraja et al. 2005).
Despite this, a striking Phelipanche, distinct from all the described French species and exclusively inhabiting the
foot and ledges of continental limestone cliffs, sporadically appeared in southern France. It was first recorded by J.
Molina in the Vallée du Gardon (Gard) in 1996, subsequently by J.C. Arnoux in the Gorges du Loup (Alpes-Maritimes)
in 2009, T. Croze in the Gorges de la Nesque (Vaucluse) in 2011, O. Peyronel in the Gorges de l’Ardèche (Ardèche)
in 2012, and H. Michaud, J. Molina, L. Garraud and J.-M. Tison on the foothills of Petit Luberon (Vaucluse) in 2013.
Although all the observers noted the unusual appearance and hosts of the plant, it remained undetermined at this time
and provisionally assigned to the group of Ph. nana (Noë ex Rchb.) Soják (1972: 140) and Ph. lainzii Gómez Nav.
et al. (2012: 17) (Tison & de Foucault 2014: 867). Finally, specific studies on calcareous cliffs by T. Croze and N.
Bianchin from 2014 to 2023 led to the discovery of several new localities in Provence and Ardèche, and a thorough,
conclusive study of the plant began in 2021.
Material & Methods
The morphological study was carried out on living plants in situ and mostly on digital photographs with scale in order
to check the colours and the value of measurements. Comparative measurements were also carried out on specimens
of morphologically related species from herbarium specimens and photographs. The haustoria of some specimens were
extracted, although the host was often quite apparent in situ due to the low plant cover of the rocky ledges and the close
spatial vicinity of Phelipanche with their host (unlike many Orobanche Linnaeus (1753: 632)). Phytosociological
surveys were carried out in some of the localities. Finally, attention was paid to the status of the populations and the
current and potential threats.
The extraction and amplification of DNA and the sequencing of the Internal Transcribed Spacer were performed,
following standard procedures, by Pablo Alvarado (www.alvalab.es). After inspection of the chromatograms (with
BioEdit 7.2.6, version of 2017, Hall (1999)) to rule out any ambiguities, new sequences were aligned (with MUSCLE,
Edgar (2004)) with a set of sequences obtained from GenBank (https://www.ncbi.nlm.nih.gov/genbank), the proposed
alignment having been afterwards manually checked.
The choice of sequences was guided by both a preliminary BLAST search and previous results, in order to ensure
the representation of similar taxa occurring in the same area and possibly overlooked ecologically similar taxa known
from other Mediterranean areas. Accessions collected from the loci classici, or as close as possible, were preferred
if available. The aligned matrix consisted of 495 bp. Phylogenetic analyses were performed with MrBayes 3.2.7
(Ronquist et al. 2012) with default settings. The consensus tree with the posterior probabilities of the retained clades
was edited with TreeViewer (Bianchini & Sánchez-Baracaldo 2024).
Taxonomic treatment
Phelipanche cingularum Croze, Carlón, J.-M.Tison, Michaud, J.Molina & Moreno Mor., sp. nov. (Fig. 1–6)
Type:—FRANCE. Alpes-de-Haute-Provence: Majastres, Gorges de Trévans, place L’Eirouelle, 23 May 2023, T.
Croze s. n. (Holotype AV!).
Diagnosis:—Plant up to 25 cm tall. Stem cream-coloured, sometimes slightly tinged with orange-brown at apex,
simple or with few short branches stemming from its lower third, densely covered with white, retrorse hairs; leaves
few, lanceolate to narrowly triangular; inflorescence long, at least the upper half of the stem, rather lax, the insertion of
flowers being clearly visible at adult stage. Flowers erecto-patent; bracts similar to the leaves, lanceolate or narrowly
triangular, much shorter than the adjacent flower, initially cream-coloured, then with a withered, brownish apex at
the end of anthesis; calyx weakly pigmented, cream-coloured to pale orange-brown, with short subulate teeth and
bracteoles, not reaching the half of the corolla; corolla 20–27 mm long, tubular-infundibuliform, densely hairy both

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internally and externally, with rather long glandular trichomes, light blue with darker longitudinal veins, dorsally
markedly bent at c. 1/3 of its length, then uniformly curved with no subapical hump, ventrally straight; upper lip
patent, bifid, with triangular, acute, irregularly dentate, conspicuously ciliate lobes; lower lip reaching 1/3 of the
corolla length, trifid, with poorly divergent, spathulate, dentate, often apiculate lobes, and two prominent, whitish,
densely hairy bumps between the lobes and the tube; stamens inserted at the level of the lower angulation of the
corolla, c. 1/3 of its length; stamen filaments basally yellowish and densely covered with long hairs, distally bluish-
white and sparsely hairy; thecae white, mostly glabrous, acute; ovary ovoid, cream-coloured, glabrous; style white,
increasingly bluish towards its apex, sparsely hairy throughout; stigma slightly bilobed, white.
FIGURE 1. Habitus. Photo by T. Croze.

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FIGURE 2. Phenological stages. Photos by T. Croze.

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FIGURE 3. Holotypus (in situ). Photo by T. Croze.

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FIGURE 4. Holotypus (herbarium sheet). Photo by T. Croze.

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FIGURE 5. Details of upper and lower lips of corolla. Photos by T. Croze and J.M. Tison.
FIGURE 6. Details of the flowers: calyx, corolla, stamens and gynoecium. Photos by J.-C. Arnoux.
Etymology:—the specific epithet is from a feminised late Latin form (genitive of cingula) derived of the classical
Latin cingulum (belt), having given place to several words applied in southern France to grassy fringes surrounding
limestone cliffs along its base or in ledges. It refers to the peculiar ecology of the plant grows on small flat rocky areas,
or « vires » in French.
French name:—Phélipanche des vires.
Host:—Parasitises species of the Brassicaceae family, mainly Hesperis laciniata All. (Fig. 7), although it proves
able to secondarily, but rarely, attack other Brassicaceae in the close vicinity of Hesperis, namely Biscutella lima,
Pseudoturritis turrita and Erysimum spp. None of these taxa has been recorded as a specific host of Orobanchaceae
until now (Sánchez Pedraja et al. 2005).
Habitat:—Sheltered limestone ledges on steep slopes and at the foot of cliffs, at the meso- and supra-mediterranean
level under temperate oceanic-submediterranean bioclimate (Rivas-Martinez & Rivas-Saenz 2011), elevation 170 to
1400 m for the known localities (Fig. 10, App. 1–3).
Phenology:—Flowering in May and early June.
FIGURE 7. View of the haustoria. Photos by T. Croze.

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Chorology and demography
Geographical distribution: Phelipanche cingularum is a rare, narrowly localized species known from thirteen places
in the Mediterranean limestone mountains of southern France (Fig. 8). It is mainly represented in Provence, rarer in
Languedoc (Tison et al. 2014). The localities are distributed in three regions, six departments and twelve municipalities.
The area of presence is approximately 14000 km², the area of occupancy 44 km² with a 4 km² grid, or 12 km² with a
1 km² grid.
Population data: From the available observations, the total number of mature individuals does not exceed 600
(App. 1). The generation length is unknown but the species seems perennial at least at haustorium stage. Population
trends are not well-known but most of them seem stable. The number of flowering individuals, however, has decreased
in the hottest and driest places (Rocher de Baude, Rocher du Cire, Les Hubacs) after 2020, certainly due to the dry
winters and springs, but this could be a temporary variation only limiting the expression of adult stages. More direct
threats such as rock reinforcement works, hiking trails, and predation by goats and chamois have been noted in situ.
The increasing development of climbing sites represents a potential threat through trampling and cliff purging. Most
of the populations are located inside regional natural parks (Mont Ventoux, Luberon, Sainte-Baume, Verdon, Préalpes
d’Azur) and included in the Natura 2000 network; only two of them are covered by prefectoral biotope protection
orders.
FIGURE 8. Distribution map of currently known localities of Phelipanche cingularum (made in december 2023).
Discussion
Morphological characters: Ph. cingularum displays a set of morphological characters not shared by the closely related
species (Tab. 1): stems rather stout, simple or barely branched, usually light coloured (yellowish brown); inflorescence
elongate, rather lax, with erect-patent flowers; calyces with short subulate teeth; corollas tubular-infundibuliform
without dorsal subapical hump, large (20–27 mm), pale blue to whitish-blue with darker longitudinal veins, mouth
taller than wide (H/L>1), lobes acute, slightly divergent, ciliate, irregularly toothed, usually patent, the throat opening
at about 100° on average; staminal filaments basally densely covered with long hairs; style sparsely glandular; stigma
white.

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In South France, Ph. cingularum should be compared with the closely related Ph. nana, which can grow in similar
habitats, sometimes together with it, and is able to parasitise many species, maybe including Brassicaceae.
All other species of Phelipanche have geographical areas and/or ecologies very distinct from those of Ph.
cingularum and their syntopia with the latter is virtually impossible. Table 1 summarizes the main differences between
Ph. cingularum and the 16 morphologically or ecologically most similar species.
Phylogenetic position: As shown in Figure 9, all the analysed accessions of Ph. cingularum constitute an
independent, strongly supported clade, a clear indication that the diagnostic traits described in the previous paragraph
and in Table 1 are not the result of mere phenotypical convergence of separate lineages induced by such a peculiar
ecology or by a common host, but synapomorphic traces of a long, independent evolutionary history. Furthermore, its
ability to succesfully parasitize Brassicaceae has evolved autonomously, as revealed by the distant positions occupied
by other parasites of that family (Tab. 1), most of which were included in our molecular analysis (as a minor shortfall
of our inquiry, it was finally not possible to obtain in time material of Ph. brassicae and Ph. chionistrae, whose
conspecificity with Ph. cingularum seems nonetheless almost unconceivable in terms of habitat and morphology).
TABLE 1. Diagnostic characters of presumably related species.
Considered
taxa Reasons for the comparison Main diagnostic features with regards to Ph.
cingularum
Ph. nana (Noë ex Rchb.) Soják
(1972: 140)
A rather common species across the area,
superficially similar, with a wide range of mostly
annual hosts.
Stems thinner; flowers smaller (12–18 mm),
more patent; corollas somewhat humped
subapically, less open, often more colourful,
with the corners of the mouth visible in lateral
view; lip lobes elliptic, entire; style usually
subglabrous.
Ph. lainzii J. Gómez Nav. et al.
(2012: 17)
Similar to Ph. nana but unknown in France, as
well as its host, Cleonia lusitanica (L.) L.
Smaller flowers (16–20 mm), with entire or
slightly erose lobes; stamen filaments basally
covered with short hairs.
Ph. oxyloba (Reut.) Soják (1972:
130)
Superficially similar; unclear relations with
western Mediterranean species.
Flowers smaller (15–19 mm); lower lip lobes
divergent and plicate, entire or sometimes
shallowly toothed; corolla less open, usually
less than 90º.
Ph. mutelii (F.W. Schultz) Pomel
(1874: 106)
Superficially similar; mainly parasitic on
Asteraceae, but reliably recorded on plants from
other families.
Stems thinner; flowers more erect; lower lip
lobes obtuse, rounded, somewhat erose, rarely
shallowly toothed; style usually subglabrous.
Ph. cernua Pomel (1874: 105)
Rather common species across the area,
occurring in similar environments, but seemingly
always parasitic on Lactuca.
Calyx with much longer segments; stigma
yellowish; stem taller, unbranched, more
pigmented; corollas grayish blue, strongly
curved dorsally; lower lip lobes elliptic, entire,
convergent; style and filaments glabrous.
Ph. brassicae (Novopokr.) Soják
(1972: 129)
A synanthropic plant, often attacking fields of
Brassicaceae.
Corollas ascending, narrower, straight, pale;
inflorescences dense; filaments subglabrous
except a few short hairs at base.
Ph. aegyptiaca (Pers.) Pomel
(1874: 107)
A synanthropic southern Mediterranean
plant, allegedly sometimes attacking fields of
Brassicaceae.
Plants often much-branched; corollae
ascending, much larger in regard of the calyx
(to 37 mm); corolla lip lobes entire.
Ph. ramosa (L.) Pomel (1874:
103)
A synanthropic plant, allegedly sometimes
attacking fields of Brassicaceae.
Plants often much-branched; flowers small;
corolla lip lobes entire.
Ph. gussoneana (Lojac.) Domina
et al. (2018: 209)
Described from limestone Mediterranean
mountains (Sicily), mistakenly recorded from
southern France.
Plants more pigmented throughout;
inflorescence denser, somewhat club-like;
corollas smaller (c. 15 mm); lower lip lobes
divergent, rhomboidal, entire.
......continued on the next page

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TABLE 1. (Continued)
Considered
taxa Reasons for the comparison Main diagnostic features with regards to Ph.
cingularum
Ph. nowackiana (Markgr.) Soják
(1972 : 130)
Parasitic on perennial Brassicaceae
(Odontarrhena) in limestone Mediterranean
mountains (Greece).
Very short plants with dense inflorescences;
corollas yellow; leaves and bracts much wider.
Ph. olbiensis (Coss.) Carlón et al.
(2008: 79)
Occurs in limestone scrub across Mediterranean
France, seemingly always parasitic on
Helichrysum.
Plant much more pigmented; stems seemingly
always unbranched; calyx-lobes falciform;
corolla dorsally deep violet-blue, ventally
whitish; stigma yellowish.
Ph. camphorosmae Carlón et al.
(2008: 35)
Occurs in southern France, seemingly always
parasitic on Camphorosma monspeliaca.
Stem unbranched; calyx segments longer, more
straight; lip lobes divergent, rounded, entire;
style subglabrous.
Ph. rosmarina (Beck) Banfi et al.
(2005: 235)
Occurs in southern France, seemingly always
parasitic on Salvia rosmarinus.
Stem stout, unbranched, often bearing flowers
for much more than half; corollas narrowed in
the middle, pale greyish-blue to greyish-violet,
sometimes slightly more pigmented at apex; lip
lobes entire; calyx teeth triangular; filaments
and style glabrous; stigma yellowish.
Ph. chionistrae Rätzel et al.
(2021: 78)
Mainly parasitic of perennial Brassicaceae
(Alyssum) in Mediterranean mountains (Cyprus).
Plant much more pigmented, very smaller;
calyx teeth triangular and short; much shorter
corollas (9–15 mm) with lip lobes entire; stigma
yellowish.
Ph. georgii-reuteri Carlón et
al. (2005: 17) (=? Ph. daninii
(Domina & Raimondo) Domina
(2017: 94)
Parasitic on perennial Brassicaceae (Vella,
Lepidium, Hormathophylla) in dry areas.
Plants stouter, woolly, more pigmented;
inflorescence dense, club-like; flowers smaller
(14–20 mm), deeper blue; lip lobes elliptic,
entire or rarely shallowly toothed.
Ph. schultzii (Mutel) Pomel
(1874: 107) Corolla lip lobes with toothed margin.
Can reach a larger size (up to 65 cm), with a
generally very elongated inflorescence and
many flowers. Very long calyx teeth (equal to
the length of the corolla tube) with a prominent
filiform apex. Base of staminate filaments
glabrous or subglabrous.
Interestingly, and despite its clear-cut external phylogenetic delineation, the clade of Ph. cingularum exhibits
a remarkable degree of internal diversity in comparison with other species like Ph. arenaria, Ph. lavandulacea, Ph.
mutelii, Ph. nana and Ph. rosmarina, much more ribotypically homogeneous even when sampled across much larger
areas (Fig. 9). The peculiar ecology of our species, by forcing a scattered, almost insular distribution, might suggest
a promising explanation for this divergence, but the apparent existence of intrapopulation diversity in Barouat and
Eirouelles brings such possibility into question.
A close relationship or even conspecificity of Ph. cingularum with Ph. gussoneana (a pretty reasonable hypothesis
as long as the latter was described from calcareous rocky areas in mostly montanous Mediterranean regions; in this
sense, Giannantonio Domina, a good expert on Ph. gussoneana, warns us—pers. comm., May 2024—that this species
frequents exactly the same habitats in Sicily) could be conclusively discarded thanks to a series of samples kindly
supplied by Corrado Marcenò, Vincenzo Ilardi and Alessandro Silvestre, including one from the very locus classicus
(Rocca Busambra. Sicily). This only confirms the clear morphological differences observable, both from the isotype
P04384573 and from the photographs taken at Rocca Busambra by G. Domina—Rätzel et al. (2018: 210)—and those
sent to us by C. Marcenò, V. Ilardi and A. Silvestre obtained there on 22-VI-2022. Compared to the quasi-monophagous
Ph. cingularum, the polyphagous Ph. gussoneana has significantly smaller and distincly infundibuliform corollas, with
entire lobe edges, and almost glabrous staminal filaments with some very short hairs; moreover, the inflorescences are
denser and shorter (somewhat club-like).

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FIGURE 9. Phylogram showing the robustness (Posterior Probability of 1) of the clade containing all and only the analysed accessions
of Phelipanche cingularum. Non-new accessions are identified by their GenBank codes (to full references of the new ones, see the text).
Only posterior probabilities above 0.8 are shown.
The taxonomic consequences of the molecular analysis of those samples will be fully discussed elsewhere, but for
now we can state our profound scepticism over the occurrence of Ph. gussoneana in France: the two records by Uhlich
& Rätzel (2021), having seen their vouchers, correspond plainly to Ph. nana (lax inflorescences with patent flowers,
somewhat humped dorsally and with elliptic, entire lower-lip lobes, etc.), a widespread species that we have personally
collected and photographed in one of those localities.
As a second corollary of our phylogenetic reconstruction, the full independence of Phelipanche sinaica (Beck)
Rätzel & Uhlich (2017: 302) could be proven by means of a couple of samples kindly sent to us by S. Rätzel and H.

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Uhlich after we shared with them our suspicion, inspired by the common host and the striking morphological similarity
(including the yellowish stigma), that it could be closely related to Ph. cernua. Those two samples correspond to the
populations cited by Rätzel et al. (2017).
Lastly, the new analysis evinces ongoing difficulties to achieve a satisfactory definition of Ph. ramosa, since
sequences corresponding to synanthropic plants presumably referable to the Linnean species appear in different
positions of the phylogram, suggesting that adaptation to agrosystems took place several times starting from different
lineages.
Ecology: Phelipanche cingularum is found inside rocky sites remarkably structured by the thick Mesozoic
carbonate series of the Jurassic (Tithonian-Kimmeridgian-Oxfordian) and Cretaceous (Urgonian) periods. These
limestone reliefs offer a wealth of geomorphological features such as cliffs and gorges that have been configured by
karstic and cryoclastic processes and water erosion. Sheltered ledges in contact with the walls (cf. App. 3), usually
south-facing, at elevation comprised between 100 and 1400 m, are the preferred habitats of Ph. cingularum.
These ledges are located on steep slopes, sometimes at the base or middle cliffs, often encircling escarpments
(cingulum). They form flat areas where rocky outcrops mingle with accumulations of rock remains (boulders, stones,
gravels) and organic matter. Depending on the depth of the limestone beds, these ledges can be relatively steep.
Accumulations of rocky material can also form small, more or less stabilized scree slopes. These soils, deeper and
richer in humic components than the adjacent steep slopes, offer opportunities for a whole range of local species.
Depending on elevation, exposure, topo-edaphic complexity, biotic interactions and trophic levels, it is possible
to find associated perennial species of pioneer dry grasslands of the Alysso alyssoidis—Sedetalia acris Moravec with
therophytes of the Stipo capensis—Trachynietea distachyae Brullo, and meso-eutrophilous hems of the Cardaminetea
hirsutae Géhu and Sisymbrietea officinalis Korneck, but also with the hemicryptophytes and chamaephytes of the
scoured lawns and low rocky scrublands of Lygeo sparti—Stipetea tenacissimae Rivas-Martínez and Festuco—
Brometea Br.-Bl. & Tüxen ex Klika & Hadač. Phanerophytes belonging to the subrupicolous matorrals of Pistacio
lentisci—Rhamnetalia alaterni Rivas-Martínez and Rhamno catharticae—Prunetea spinosae Tüxen or elements
linked to the saxicolous communities of Polypodion serrati Br.-Bl. in Br.-Bl., Roussine & Nègre, Asplenion glandulosi
Br.-Bl. & Meier in Meier & Br.-Bl. or Potentillion caulescentis Br.-Bl. in Br.-Bl. & H. Jenny can be found (cf. App.
2). This intricacy makes the phytosociological interpretation of these ledge vegetations particularly complex, as they
are still poorly studied and known in the French Mediterranean area. The notable feature of this vegetation is the
significant occurrence and the dynamism of nitrophilous elements, especially Hesperis laciniata, more plentiful here
than anywhere else. At higher elevation, when conditions are favorable, H. laciniata can form luxuriant fringes at the
edge of rock walls and subrupicolous matorrals, with others perennials such as Alliaria petiolata (M. Bieb.) Cavara &
Grande, Delphinium fissum Waldst. & Kit., Lamium garganicum L. These species are mixed with various nitrocline
and nitrophilous therophytes such as Anisantha sterilis (L.) Nevski, A. tectorum (L.) Nevski, Anthriscus caucalis M.
Bieb., Geranium purpureum Vill., G. rotundifolium L., Veronica hederifolia L. This is probably the best environment
for Phelipanche cingularum (Fig. 10).
The droppings of rock birds and especially ungulates like chamois and goats, which regularly use these ledges
as passage, grazing and resting areas, maintain trophic conditions favorable to these associations. Moreover, the
germination of H. laciniata is favored by a high concentration of nitrates in the soil (Yücel et al. 2008), and the
moderate effects of trampling by ungulates also help to maintain an open environment probably beneficial to Ph.
cingularum, regularly found on skinned soil, as Carlón et al. (2015: 49) and Pavon et al. (2015: 102) have noted for
other Orobanchaceae.
Depending on winter and spring rainfalls, H. laciniata and Ph. cingularum populations can fluctuate greatly
from one year to the next. A strong shortage of rainfall, as in 2023, drastically reduced their populations at low and
medium elevation. In this case, the pressure of ungulate browsing on these two species can considerably reduce their
reproductive capacity. Accordingly, low-elevation populations of Ph. cingularum may disappear temporarily, or occur
only on north-facing rocks.
Conservation: The effects of urbanization, such as the securing of risks associated with rock faces to protect
property and people, may have led to local damage. Urbanization strengthens the attractiveness of natural rocky sites
to tourists for sports activities (climbing, via ferrata, hiking) which can dramatically impoverish the plant associations.
The aridification of the climate may weaken the lowest populations of Ph. cingularum, making them more vulnerable
to other damages. However, due to the lack of information on the chorology and dynamics of the species, the IUCN
(2012, 2022) criteria of population decline (A), recent reduction in range (C) and risk of extinction in the wild over
100 years France are not applicable. Not is the criterion B since more than 10 populations exist. The only criterion
applicable for now is D1 with a total number of individuals estimated at less than 1000, placing the species in the

A NEW SPECIES OF PHELIPANCHE FROM FRANCE Phytotaxa 653 (1) © 2024 Magnolia Press • 13
vulnerable category « VU ». If new individuals and/or localities and/or a larger area of occupancy are found in the
future, the species will be transferred to the Near Threatened « NT » category.
FIGURE 10. Typical fringes of ledges with Hesperis laciniata, Lamium garganicum, Delphinium fissum and Phelipanche cingularum at
l’Eirouelle (elevation: 1140 m). Photo by T. Croze.
Conclusion
Phelipanche cingularum is a new species remotely related to the complex of Ph. ramosa, possibly limited to south-
eastern France, but to be searched from Catalonia to Piemont and Liguria. It lives only in steep, rocky environments,
usually far from hiking tracks and difficult to access. This is probably one of the reasons of its underobservation in a
well-known region such as Provence.
It must be emphasized, however, that the detection of such a species identifiable only in a living state, as well as
many cryptic Orobanche, Phelipanche, Allium etc., would have been almost impossible before the popularization of
digital photography. In this context, we emphasize the importance of the visit of loci typici and loci classici of formerly
described taxa, including presumed synonyms, which often leads to the rediscovery of overlooked species (Orobanche
balsensis (J.A. Guim.) Carlón et al. (2015: 39), O. centaurina Bertol. (1846: 430), O. grenieri F.W. Schultz (1845:
739), O. leptantha Pomel (1874: 110), O. loscosii Carlón et al. (2011: 93), O. lycoctoni Rhiner (1890–91: 253), O.
serbica Beck & Petrovič in Petrovič (1885: 146), Phelipanche cernua, Ph. georgii-reuteri...).

CROZE ET AL.
14 • Phytotaxa 653 (1) © 2024 Magnolia Press
Acknowledgments
We are grateful to our late friend Jean-Claude Arnoux for his floral illustrations, Jean-Pierre Roux and Nicolas Bianchin
for helpful discussions and bibliographical resources, Olivier Peyronel for his observation, Laurent Michel for his
research in Luberon montains, Teddy Baumberger for corrections to the manuscript, Holger Uhlich and Stephan Rätzel
for provinding useful materials and fruitful comments, and the Italian group: Corrado Marcenò, Vincenzo Ilardi and
Alessandro Silvestre for their notes, photographs and sampling of Phelipanche gussoneana and its hosts in its locus
classicus. This work was supported by the SV-23-GIJON-JBA project.
Conflict of interest
The authors declare to have no conflict of interest.
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16 • Phytotaxa 653 (1) © 2024 Magnolia Press
Appendix 1. Eco-geographical features.
Region Counties Municipality
Mountainous
massif /
Valley
Locality Observador / Date Elevation Exposition Geology Estimated
population Know threat Potentiel
threat
Regulatory
protection
zone (strong
protection)
Contractual
protection zone
(low protection)
Auvergne-
Rhône-Alpes Ardèche
Vallon-Pont-
d’Arc
Gorges de
l’Ardèche Le Tiourre
O. Peyronel 2012
N. Bianchin & T. Croze
2013
340 m S-E Cretaceous limestone
(Urgonian) 11–100 ind. Ungulate pressure Climbing /
Via ferrata None Site N2000
Saint-Didier-
sous-Aubenas
Vallée de
l’Ardèche
Rochers de
Jastres N. Bianchin 2023 321 m N-O Jurassic limestone
(Oxfordian) 1–10 ind. None None
Prefectural
biotope
protection order
Site N2000
Occitanie Gard Anduze Vallée du
Gardon
Porte des
Cévennes
J. Molina 1996
T. Croze 2023 170 m S-E / N
Cretaceous limestone
(Kimmeridgian-Tithonian)
Jurassic limestone
(Oxfordian)
11–100 ind. Securing rocky risks
Ungulate pressure
Climbing /
Via ferrata None Site N2000
Provence-
Alpes-Côte
d’Azur
Vaucluse
Robion Petit
Luberon
Rocher de
Baude
H. Michaud, J. Molina,
L. Garraud and J-M.
Tison 2013
T. Croze 2023
240 m S-O Cretaceous limestone
(Urgonian) 11–100 ind. Hiking
Securing
rocky risks
Climbing
Prefectural
biotope
protection order
Regional natural park
Site N2000
Monieux Gorge de la
Nesque
Rocher du
Cire
T. Croze 2011, 2013,
2014, 2015, 2022, 2023 730 m S-E Cretaceous limestone
(Urgonian) 1–10 ind. Ungulate pressure Climate
change
Prefectural
biotope
protection order
Regional natural park
Site N2000
Alpes-
Maritimes
Gourdon Gorge du
Loup
L’Oustaou
Dou Rey J.-C. Arnoux 2009 740 m S-E Calcaires jurassiques
(Oxfordian) Unknow Unknow
Securing
rocky risks
Climbing /
Via ferrata
None Regional natural park
Site N2000
Saint-Auban Haut-Estéron Barouat T. Croze 2014, 2022 1380 m S Cretaceous limestone
(Kimmeridgian-Tithonian) 11–100 ind. None Climbing /
Via ferrata None Regional natural park
Var
Pourrières Mont
Aurélien
Olympe T. Croze 2022 800 m N-E Cretaceous limestone
(Kimmeridgian-Tithonian) 1–10 ind. None Climbing /
Via ferrata None Regional natural park
Pourcieux Les Hubacs T. Croze 2022 835 m N-O Cretaceous limestone
(Kimmeridgian-Tithonian) 1–10 ind. None Climbing /
Via ferrata None Regional natural park
Alpes-de-
Haute-
Provence
Saint-
Vincent-sur-
Jabron
Vallée du
Jabron
Les Baumes
Rochers T. Croze 2014 1060 m S-E Cretaceous limestone
(Kimmeridgian-Tithonian) 1–10 ind. Ungulate pressure None None None
Majastres Gorges de
Trévans
Pont de Tuf T. Croze 2023 950 m S-E Cretaceous limestone
(Kimmeridgian-Tithonian) 1–10 ind. Ungulate pressure None None Regional natural park
Site N2000
L’Eirouelle T. Croze 2017, 2022,
2023
1140 m
1180 m S / S-E Cretaceous limestone
(Kimmeridgian-Tithonian) 11–100 ind. Ungulate pressure None None Regional natural park
Site N2000
Castellane Gorges du
Verdon Les Caires T. Croze 2023 1070 m S Cretaceous limestone
(Kimmeridgian-Tithonian) 1–10 ind. None Climbing /
Via ferrata None Regional natural park

A NEW SPECIES OF PHELIPANCHE FROM FRANCE Phytotaxa 653 (1) © 2024 Magnolia Press • 17
Appendix 2. Associated species
Porte des Cévennes
Rocher de Baude
Les Hubacs
Rocher du Cire
Pont de Tuf
Baumes Rochers
L’Eirouelle
Barouat
Freq. 170 m 239 m 835 m 730 m 950 m 1060 m 1140 m 1380 m
Hesperis laciniata All. (main host) 91 2 2 2 2 3 3 3
Phelipanche cingularum sp. nov. 9r 1 r r1 r 1 1
Species of nitrophilous vernal annual fringes (Cardaminetea hirsutae)
Geranium rotundifolium L. 7• • • •• • •
Geranium purpureum Vill. 6• • • •• •
Geranium lucidum L. 4 • • • •
Centranthus calcitrapae (L.) Dufr. 4 •• • •
Anthriscus caucalis M.Bieb. 3 • • •
Cardamine hirsuta L. 3 • • •
Veronica arvensis L. 2 • •
Arabidopsis thaliana (L.) Heynh. 2 • •
Cerastium brachypetalum Pers. 2 • •
Veronica hederifolia L. 2 • •
Myosotis ramosissima Rochel 2 • •
Campanula erinus L. 2 • •
Torilis africana Spreng. 2• •
Mercurialis huetii Hanry 2• •
Scandix stellata Banks & Sol. 2• •
Species of ruderal wasteland (Sisymbrietea officinalis / Papaveretea rhoeadis / Glechomo hederaceae - Urticetea dioicae)
Anisantha madritensis (L.) Nevski 4• • • •
Anisantha sterilis (L.) Nevski 4 • • • •
Urospermum picroides (L.) Scop. ex F.W.Schmidt 3• • •
Crepis foetida L. 3 • • •
Torilis leptophylla (L.) Rchb.f. 3 • • •
Carduus litigiosus Nocca & Balb. 3 • • •
Papaver dubium L. 2 • •
Sisymbrium orientale L. 2 • •
Lamium amplexicaule L. 2 • •
Lactuca serriola L. 2 • •
Alliaria petiolata (M.Bieb.) Cavara & Grande 2 • •
Brachypodium rigidum (Roth) Link 1•
Sonchus oleraceus L. 1 •
Avena barbata Pott ex Link 1 •
Hordeum murinum L. 1 •
Annual oligotrophic species (Stipo capensis - Trachynietea distachyae)
Arenaria leptoclados (Rchb.) Guss. 3 • • •
Sideritis romana L. 3•• •
Catapodium rigidum (L.) C.E.Hubb. 2 • •
Teucrium botrys L. 2 • •
Saxifraga tridactylites L. 1 •
Erodium cicutarium (L.) L’Hér. 1 •
Medicago coronata (L.) Bartal. 1 •
Pioneer grasslands on slabs and flats with rocky debris (Sedo albi - Scleranthetea biennis)
Sedum album L. 8• • • •• • • •
Petrosedum sediforme (Jacq.) Grulich 6• • • • • •
Lactuca perennis L. 5 • • • • •
Alyssum alyssoides (L.) L. 5 •• • • •
Antirrhinum majus L. 4• • • •
Arenaria serpyllifolia L. 4 • • • •

CROZE ET AL.
18 • Phytotaxa 653 (1) © 2024 Magnolia Press
Sedum dasyphyllum L. 3 • • •
Hornungia petraea (L.) Rchb. 3 • • •
Poa bulbosa L. 3 • • •
Medicago minima (L.) L. 3 • • •
Clypeola jonthlaspi L. 2 • •
Melica minuta L. 2 • •
Ziziphora acinos (L.) Melnikov 2 • •
Saxifraga fragosoi Sennen 1•
Sempervivum calcareum Jord. 1•
Petrosedum ochroleucum (Chaix) Niederle 1•
Rocky spurs on shady ledges (Polypodietea)
Polypodium cambricum L. 1 •
Asplenium ceterach L. 1 •
Asplenium trichomanes subsp. quadrivalens D.E.Mey. 1 •
Umbilicus rupestris (Salisb.) Dandy 1 •
Perennial and biennial species of basic scree (Noccaeetalia rotundifoliae)
Delphinium fissum Waldst. & Kit. 4 • • • •
Lamium garganicum L. 4 • • • •
Low-scrubby scrubland (Rosmarinetea officinalis / Festuco - Brometea)
Thymus vulgaris L. 6 • •• • • •
Iris lutescens Lam. 5• • • • •
Galium corrudifolium Vill. 5• • • • •
Stachys recta L. 5 • •• • •
Silene italica (L.) Pers. 4• • • •
Saponaria ocymoides L. 3 • • •
Dactylis glomerata subsp. hispanica (Roth) Nyman 3• • •
Helianthemum nummularium (L.) Mill. 3 • • •
Centaurea paniculata subsp. leucophaea (Jord.) Arcang 3 • • •
Genista cinerea (Vill.) DC. 3 • • •
Festuca cinerea Vill. 2 • •
Satureja montana L. 2 • •
Biscutella lima Rchb. (secondary host) 2• •
Centaurea stoebe L. 2• •
Euphorbia characias L. 2• •
Allium sphaerocephalon L. 2 • •
Euphorbia spinosa L. 1 •
Lavandula angustifolia Mill. 1 •
Lilium pomponium L. 1 •
Leucanthemum pallens (J.Gay ex Perreym.) DC. 1 •
Minuartia glomerata subsp. burnatii (Rouy & Foucaud) Favarger & F.Conti 1 •
Sub-rupicolous matorrals (Rhamno catharticae - Prunetea spinosae)
Amelanchier ovalis Medik. 5 • • • • •
Buxus sempervirens L. 4 • • • •
Cotoneaster nebrodensis (Guss.) K.Koch 3 • • •
Rhamnus saxatilis Jacq. 3 • • •
Quercus ilex L. 3• • •
Rhamnus alaternus L. 3• • •
Juniperus phoenicea subsp. phoenicea L. 3 • ••
Other species
Phelipanche nana (Reut.) Soják 3 • • •
Pseudoturritis turrita (L.) Al-Shehbaz (secondary host) 3• • •
Orobanche caryophyllacea Sm. 1 •
Orobanche centaurina Bertol. 1 •
Orobanche grenieri F.W.Schultz 1 •
Orobanche reticulata Wallr. 1 •

A NEW SPECIES OF PHELIPANCHE FROM FRANCE Phytotaxa 653 (1) © 2024 Magnolia Press • 19
Appendix 3. Phelipanche cingularum on rocky escarpments (yellow arrow). A. Barouat (elev. 1280–1380 m). B. l’Eirouelle (elev. 1140
m). C. Rocher du Cire (elev. 730 m). D. Rocher de Baude (elev. 240 m). Photos by T. Croze.