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The overlooked Orobanche balsensis (J.A. Guim.), Comb. Nov., and some remarks on O. Subbaetica Triano & A. Pujadas

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The overlooked Orobanche balsensis (J.A. Guim.), Comb. Nov., and some remarks on O. Subbaetica Triano & A. Pujadas

Abstract and Figures

In this paper a neglected taxon of Orobanche s. str. is shown to deserve specific rank for combining a set of correlated morphological features and a particular host (Carlina). As it had already been validly named at varietal and subva-rietal rank by J. A. Guimarães, it needn’t be validly described, but is here typified, raised to the rank of species, thoroughly described, illustrated with detailed drawings and colour photographs and demonstrated to occur well beyond its terra classica not only in Portugal and Spain, but probably also in southern France, Corsica and Sicily. In addition, some morphological, chorological and iconographical additions to the knowledge of the recently described Orobanche subbaetica will be done.
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Flora Montiberica 60: 38-53 (V-2015). ISSN: 1138-5952, edic. digital: 1998-799X
THE OVERLOOKED OROBANCHE BALSENSIS (J.A. GUIM.),
COMB. NOV., AND SOME REMARKS ON
O. SUBBAETICA TRIANO & A. PUJADAS
Luis CARLÓN1, Manuel LAÍNZ2, Gonzalo MORENO MORAL3
& Óscar SÁNCHEZ PEDRAJA4
1 Jardín Botánico Atlántico. Avenida del Jardín Botánico, 2230. E-33394 Cabueñes,
Gijón (Asturias) España. lcarlon77@gmail.com
2 Avda. Hnos. Felgueroso, 25. E-33205 Gijón (Asturias) España.
lainz@colegioinmaculada.es
3 Santa Clara, 9-1 dcha. E-39001 Santander (Cantabria) España.
4 E-39722 Liérganes (Cantabria) España. osanchez@farmalierganes.com
ABSTRACT: In this paper a neglected taxon of Orobanche s. str. is shown to
deserve specific rank for combining a set of correlated morphological features and a
particular host (Carlina). As it had already been validly named at varietal and subva-
rietal rank by J. A. Guimarães, it needn’t be validly described, but is here typified,
raised to the rank of species, thoroughly described, illustrated with detailed drawings
and colour photographs and demonstrated to occur well beyond its terra classica not
only in Portugal and Spain, but probably also in southern France, Corsica and Sicily. In
addition, some morphological, chorological and iconographical additions to the
knowledge of the recently described Orobanche subbaetica will be done. Key words:
Orobanchaceae, Mediterranean flora, Carlina, parasitic plants, biosystematics, bio-
geography, evolution.
RESUMEN: La subestimada Orobanche balsen sis (J.A. Guim.), comb. nov.,
y algunos comentarios acerca de O. subbaetica Triano & A. Pujadas. En este traba-
jo se dan pruebas de la pertinencia del rango específico para un taxon desatendido de
Orobanche s. str. en el que coinciden todo un conjunto de caracteres morfológicos y un
hospedante propio (Carlina). Puesto que J. A. Guimarães ya le había dado nombre vá-
lido en los rangos varietal y subvarietal, no es preciso describirlo como nuevo, y aquí
tan solo se lo tipifica, se lo eleva al rango de especie, se lo describe en detalle, se lo
ilustra con dibujos y fotografías en color y se muestra que su área de distribución reba-
sa la terra classica y se extiende, no solo por España y Portugal, sino muy probable-
mente por el sur de Francia, Córcega y Sicilia. Además, se hacen algunas adiciones
morfológicas, corológicas e iconográficas al conocimiento de la recién descrita Oro-
banche subbaetica. Palabras clave: Orobanchaceae, flora mediterránea, Carlina, plan-
tas parásitas, biosistemática, biogeografía, evolución.
INTRODUCTION
Both morphology (namely the crenate
corolla lip lobes and the deeply divided
calyx, with long and narrowly lanceolate,
often distally filiform teeth) and, more
recently, molecular analysis (SCHNEE-
WEISS & al. 2004) have recognised the
existence of a widespread and well-
characterised set of species typified by
Orobanche minor and categorised under
several taxonomic ranks Orobanche ser.
Minores Beck (1882) ["tribus" sensu BECK
(1890) = "Grex" sensu BECK (1930)], O.
38
L. CARLÓN, M. LAÍNZ, G. MORENO & Ó. SÁNCHEZ
subsect. Minores (Beck) Teryokhin (1993).
But the internal structure of this group,
obscured by the subtility of morphologi-
cal divergences as well as by a remarka-
ble molecular homogeneity, has been
controversial. Some authors have adopted
an extremely synthetic treatment, not ad-
mitting but a handful of species to which
the rest were synonymised or, at most,
infraspecifically subordinated, whereas
other have advocated very analytical
approaches, with dozens of taxa arranged
into complex hierarchical systems.
Such discrepant systematic views are
well apparent in the Iberian Peninsula,
which represents a remarkable hotspot for
this group. The seminal monograph by
BECK (1890) inspired the detailed studies
by GUIMARÃES (1904), who offered a
complex and extremely analytical taxo-
nomic treatment. On the opposite side,
lies the synthetic review by FOLEY (2001).
PUJADAS (2002) presents a more thorough
and balanced review of the Iberian diver-
sity of these plants, also summarising his
own noticeable contributions. Important,
comprehensive studies about the nature
and evolutionary roots of the diversity of
the group, partially referred to Iberian
plants, are those by THOROGOOD & al.
(2008, 2009).
The Minores include both broad host
range species, namely O. minor s. str. (pa-
rasite mostly on Asteraceae, Fabaceae and
Apiaceae, more rarely on other families
like Solanaceae, its real host range having
probably been exaggerated in literature as
a result of the misidentification of the
actual host and confusion among species),
and truly specialised, morphologically re-
cognisable and for the most part molecu-
larly supported species like O. ame-
thystea, O. artemisiae-campestris, O.
ballotae, O. hederae, O. picridis and O.
santolinae (see FRAJMAN & al., 2013: 6).
The evolutionary basis and significance of
this duality will be explored in some de-
tail in the discussion section of this paper.
Orobanche balsensis (J.A. Guim.) Carlón,
M. Laínz, Moreno Mor. & Ó. Sánchez,
comb. nov.
Orobanche loricata var. synomora J.A. Guim.
subvar. balsensis J.A. Guim. in Brotéria 3: 163[-
164] (1904) [basionym]. Ind. loc.:β. 2 (O.)
balsensis Guim.”. Portugal, “Algarve: Nas co-
llinas aridas proximo de Tavira (parasita da
Carlina corymbosa L., Welwitsch!)(cf. Gui-
marães, 1904: 164). Lectotype [designated here
(hic designatus)]: LISU 34658 (Flor. Algarb. Nº
780 / Orobanche / ad radices Carlina hispani-
cae parasitic / pr. Tavira in collinis aridis. / ... /
maio [1]847).
O. loricata f. balsensis (J.A. Guim.) Beck in Engl.,
Pflanzenr. 96: 200 (1930).
= O. loricata var. synomora J.A. Guim. subvar.
Synomora J.A. Guim. in Brotéria 3: 163[-164]
(1904). Ind. loc.:β. 1 ( O. ) synomora Guim.”.
Portugal, “Algarve: Serra de Monchique: Foia
(sobre a Carlina sp., Welwitsch!); na charneca
arenosa do Cabo de S. Vicente (fórma de tran-
sição para a O. Picridis Schultz, com as anthe-
ras mais curtamente mucronadas, Welwitsch!)”
(cf. Guimarães, 1904: 164). Lectotype [desig-
nated here (hic designatus)]: LISU 34657 [Flor.
Algarb. Nº 944 / Orobanche / In adition ... Foiae
/ in Carlina ... / ... / ... / 26 Jun (1)847].
= O. loricata f. synomora (J.A. Guim.) Beck in
Engl., Pflanzenr. 96: 200 (1930).
= O. lusitanica J.A. Guim., nom. nud., in schedis.
= O. pseudoloricata J.A. Guim., nom. nud., in
schedis.
?= O. minor subsp. neglecta var. strangulate f.
genuine J.A. Guim. in Brotéria 3: 174 (1904),
nom. illeg. Ind. loc.: γ I (O.) strangulata
Guim.”. Portugal, “Centro littoral:Otta (jun-
cto da Carlina sp., Welwitsch!) / Algarve:
Estoy (A. Guimarães!)”. Lectotype [designated
here (hic designatus)]: LISU 34683 (Orobanche
/ Tota sordida purpu-/reo violacea / auf Car-
lina / ... Otta junio [1]848) !)” (cf. Guimarães,
1904: 178).
?= O. minor subsp. Neglecta var. strangulate subvar.
Strangulate J.A. Guim. Ex Beck in Engl., Pflan-
zenr. 96: 212 (1930).
O. minor [race / ecotype] “on Carlina corymbosa
Thorogood, C.J., Rumsey, F.J., Harris, S.A. &
Hiscock, J. in Plant Syst. Evol. 282, Nos. 1-2:
37[-38] f. 4f (2009). Ind. loc.: “on Cape St Vin-
cent, Portugal”.
O. amethystea auct., non Thuill.
O. artemisiae-campestris auct., non Vaucher ex
Gaudin
O.castellana auct., non Reut.
Plants sharing with other Minores the
prolonged calyx teeth and the crenate
corolla lip margin, but readily distinct by
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Flora Montiberica 60: 38-53 (V-2015). ISSN: 1138-5952
The overlooked Orobanche balsensis (J.A. Guim.), comb. nov., and remarks on O. subbaetica
their slender habit, their lax and short
inflorescences (shorter than the rest of the
stem) and the markedly patent flowers
which are tubular in shape, a bit widened
distally and with a mostly straight dorsal
line (such a straight, patent, geniculated
corolla evokes O. amethystea, which is
nonetheless a much more robust, thick-
stemmed plant with more crowded in-
florescences and bracts exceeding the
flowers). Parasitic on Carlina (Astera-
ceae).
Description: Stem (14)17-34(37,5) ×
(0,16)0,30-0,42 cm, simple, usually slen-
der, purplish or creamy (somewhat pur-
ple-tinged towards the inflorescence),
glandular-hairy throughout (hair density
decreasing from the inflorescence towards
the base). Leaves 10-15 × 2,3-5 mm,
triangular-lanceolate. Inflorescence (3,8)
6,5-11,0(12,5) × (2,8)3,0-3,6(4,4) cm,
comparatively short -shorter than the rest
of the stem [ratio long. infl. / long. stem =
0,27-0,36(0,41)]-, oblong [ratio width
infl. / long. infl.= (0,26)0,33-0,48(0,60)],
lax (most of the flowers distant from each
other), dense only in the apex [(9)12-
26(31) flowers]; bracts (9)10-13,0(14) ×
(2,9)4-5,0 mm, lanceolate, commonly
shorter than the corolla [ratio long. bract /
long. corolla = 0,58-0,80(0,93)], purple
throughout, more rarely creamy in the
base, densely covered with glandular hairs
(yellowish gland). Calyx (9)11-12,5(14) ×
(2,1)2,5-3,5(4) mm, segments free, ± un-
equally two-toothed with the upper tooth
usually longer, pink-purplish (more deep-
ly in the middle nerves of the teeth and in
the upper 2/3 of the tube, sometimes ±
creamy towards the base), densely cove-
red with glandular hairs (yellowish
gland), specially in the teeth, which are
(7,0)9,0-9,5(11,5) mm, long, unequal,
narrowly lanceolate, acuminated to fili-
form in the apex and usually much longer
than the tube [ratio long. teeth / long. tube
= (1,75)2,0-4,0(5,6)]. Corolla (15,0)16-20
mm, ratio long. calyx / long. corolla=
(0,46)0,55-0,73,5(0,83), patent (forming
an angle of 57-75º with the axis of the
inflorescence), ± tubular, slightly broade-
ned distally, dorsal line geniculate at the
point where the stamens are insered and
then straight; outer surface, specially in
the dorsal area, densely covered with
glandular hairs (translucent hairs with ye-
llowish gland), whitish with purple ner-
ves, sometimes purplish-spotted distally;
upper lip shallowly bilobate, with its
lobes variably displayed (erect, bent for-
ward or even a bit deflexed) during the
anthesis, ± rounded in shape, with undu-
lated and crenate-denticulate margins;
lower lip trilobate, with deflexed, almost
overlapping or slightly distant lobes,
rounded to subquadrangular in shape and
with ± crenate-denticulate margins. Fila-
ments 6,5(8,5)-9,0(10,2) mm, insered at
(2,1)2,5-3,5(4) mm from the base of the
corolla, hairy only in their basal half
(traslucent and rather long glandless
hairs), whitish or somewhat yellowish and
more or less purple-tinged below the
anthers. Anthers 1,5-1,75 × 0,7-0,9 mm
(beak c. 0,3 mm), purple or brown-gre-
yish (beak white, conspicuous), glabrous
except along the sutures in the basal half.
Ovary whitish, glabrous. Style ± pure
white and glabrous in the base and in-
creasingly purplish and covered with
short glandular hairs towards the apex.
Stigma ± deeply purple.
Its only confirmed host is Carlina co-
rymbosa L., s.l. (Asteraceae), and its cu-
rrently known area covers the south-
western corner of the Iberian Peninsula
(Portuguese region of Algarve and
Spanish provinces of Cádiz and Málaga)
at elevations of 0-900 m a.s.l. As ex-
plained below, it’s possible and even pro-
bable that the species occurs as well in
other regions around the Mediterranean,
where it might be parasitic on other spe
cies of Carlina.
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Flora Montiberica 60: 38-53 (V-2015). ISSN: 1138-5952
L. CARLÓN, M. LAÍNZ, G. MORENO & Ó. SÁNCHEZ
Studied specimens other than the
above cited lectotypes:
POR, Algarve: Faro: 29SNA0197, Vila do
Bispo, Laginha, near Cabo de São Vicente,
not far from Sagres, 60 m, parasitic on (!) C.
corymbosa in the clearings of the scrub, on
stony soils, 29-IV-2013, G. Moreno Moral
MM0041/2013 (herb. Sánchez Pedraja
13841); 29SNB9001, Mata do Pontal, near
Gambelas, but in the concelho of Loulé, 25 m,
beside C. corymbosa in a grassy wayside, 1-
V-2014, G. Moreno Moral MM0007/2014 et
MM0008/2014 (herb. Sánchez Pedraja 13963
et 13964); 29SNB9102, Mata do Pontal,
South from Vale da Venda, above the bed of
stream Biogal, 30 m, parasitic on C. corym-
bosa in a shady slope under pine trees, 2-V-
2014, G. Moreno Moral MM0011/2014 (herb.
Sánchez Pedraja 13967).
ESP, Andalua: *Cádiz: 30STF8315,
San Roque, southern slope of Cerro del Águi-
la, Pinar del Rey, near San Roque, 110 m,
parasitic on C. corymbosa on sandy soils in
the clearings of a pine grove, 15-V-2011, J.A.
García Rojas, G. Moreno Moral, F. Muñoz
Secilla & G. Velasco Cintrano MM0033/2011
(herb. Sánchez Pedraja 13668); 30STF8468,
Benaocaz, above the Cortijo de la Fuentezue-
la, near Benaocaz, 860 m, beside C. corym-
bosa on soils shaded by holm oaks by the path
to the Tajo del Cabrero, 13-VI-2013, L.
Carlón & G. Moreno Moral MM0094/2013
(herb. Sánchez Pedraja 13882). *Málaga:
30STF8741, Gaucín, la Almuña, 400 m,
parasitic on C. corymbosa in sunny meadows
but under the shadow of scattered holm oaks,
23-V-2004, G. Gómez Casares & G. Moreno
Moral MM0097/2004 (herb. Sánchez Pedraja
11852 & 11853).
The molecular analysis by THORO-
GOOD & al. (2009) (see discussion section
of this paper) supports the singularity of
this plant, whose morphological distinc-
tiveness, that we had noticed already in
2004, was also neatly outlined there, but
without taking the step of proposing its
formal systematic recognition. Neverthe-
less, our bibliographical surveys in search
of a name for the species suggested that
the first in realizing the existence of this
well characterised Carlina parasite was
José d'Ascensão Guimarães during his
profound review of Portuguese Oroban-
chaceae. Intentional visits to some of the
Portuguese localities cited by Guimarães
strengthened that suspicion, which at last
could be fully confirmed. As pointed out
in the above nomenclatural list, it is un-
clear whether the type material of the
“var. strangulata” and that of balsensis
are taxonomically identical, and thus whe-
ther both names are synonyms: the flow-
ers of the “var. strangulata” are shorter
and with an intense apical tinge. We will
try to study the plant in the field as soon
as possible, but in the mean time it can’t
be discarded that it corresponds to a form
of the widespread and host-unspecific O.
minor s. str.
In order to prevent host-based misi-
dentifications, it must be kept in mind that
broomrapes other than O. balsensis para-
sitize on Carlina around the Mediterrane-
an. Carlina nebrodensis and C. Sicu-la,
for instance, are amongst the short-lived
Asteraceae on which O. canescens C. Presl
is known to grow. But this species de-
scribed from Sicily differs from O. Bal-
sensis by its much shorter, erecto-patent
and dorsally curved flowers, and is barely
distinguished from O. minor by the yel-
lowish-tinge in the top of the inflorescen-
ce and in the stigma (DOMINA & STEPA-
NEK, 2009).
This doesn’t mean necessarily that O.
balsensis is absent from Sicily herself,
from where we have seen pictures of
plants presumably parasitic on Carlina
and showing a strong resemblance to the
species [e. g. those taken in Monte Catal-
fano by C. Marcenò, those that A. La Ro-
sa took near Castelvetrano, and two pic-
tures taken in Castiglione di Sicilia by F.
Russo (viewable, sub O. crenata, at
https://www.flickr.com/photos/etnature)].
The case of Corsica, judging by the pic-
tures taken in La Rondinara by J.-P.
Chabert (http://kalliste.flora.free.fr/, sub
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Flora Montiberica 60: 38-53 (V-2015). ISSN: 1138-5952
The overlooked Orobanche balsensis (J.A. Guim.), comb. nov., and remarks on O. subbaetica
O. minor), results, not surprisingly, analo-
gous, as does that of Southern France at-
tending to the specimen P 04419134 -
herb. Grenier (“n.º2 / Cette [Sète, Hérault]
- Sur le Carlina / corymbosa / fleur bleu /
Godron 1851”); but this notwithstanding,
and despite our inability to see any reason
why this specialised Carlina parasite
should be restricted to the small area from
which we can so far confirm its occur-
rence, we prefer to be cautious and wait
for sounder proofs and in vivo studies be-
fore firmly assigning a larger geograph-
ical range to O. balsensis, and in fig. 1 we
show as still uncertain all of the non-
Iberian localities.
For the above stated reasons, it ap-
pears too soon to evaluate sensibly the
conservation status of O. balsensis. As-
suming the so far known data in an exer-
cise of ephemeral formalism, an applica-
tion of the IUCN (2013) criteria would
lead to consider the species vulnerable,
VU [B1ac(ii, iii,iv)+2ac(ii,iii,iv); D1]. Its
habitat is subject to some degree of hu-
man impact by means of animal husband-
ry, silvicultural activities and roadworks,
but the species is likely to overcome such
pressures by following its abundant host
from recently disturbed to abandoned
patches, and there is no reason to suspect
significant declines. The confirmation of
the French and Italian populations would
suffice to take the conservation status of
the species well into the Least Concern
(LC) category.
Fig. 1. Known distribution of Orobanche balsensis (red) and O. subbaetica (yellow). Stars in-
dicate not fully confirmed but very plausible records (see text). The map was plotted with QGIS
2.6.0 and uses as base a cross-blended hypsometric tint (www.naturalearthdata.com) correspond-
ing to an area approximately between 30 and 48ºN and between 10ºW and 20ºE. Each side of the
grid equals to 6 arc degrees.
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Flora Montiberica 60: 38-53 (V-2015). ISSN: 1138-5952
L. CARLÓN, M. LAÍNZ, G. MORENO & Ó. SÁNCHEZ
Fig. 2. Lectotype of Orobanche balsensis (LISU 34658)
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Flora Montiberica 60: 38-53 (V-2015). ISSN: 1138-5952
The overlooked Orobanche balsensis (J.A. Guim.), comb. nov., and remarks on O. subbaetica
Fig. 3. Orobanche balsensis: Portugal, Algarve, Faro, Vila do Bispo, Laginha, near Cabo de São
Vicente, not far from Sagres, 29-IV-2013, photo by G. Moreno Moral (herb. Sánchez Pedraja
13841). These plants inhabit a very sunny and windy spot, whereas in other localities the species
grows in less exposed places under the shade of trees. It is thus conceivable that the paler colour of
these plants is purely phenotypical (the marked, diagnostic dark veins of the corolla lobes remain,
however, as obviously do all the distinctive morphological features).
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Flora Montiberica 60: 38-53 (V-2015). ISSN: 1138-5952
L. CARLÓN, M. LAÍNZ, G. MORENO & Ó. SÁNCHEZ
Fig. 4. Orobanche balsensis: Spain, Cádiz, San Roque, southern slope of Cerro del Águila, Pinar
del Rey, near San Roque, 15-V-2011, photo by G. Moreno Moral (herb. Sánchez Pedraja 13688).
45
Flora Montiberica 60: 38-53 (V-2015). ISSN: 1138-5952
The overlooked Orobanche balsensis (J.A. Guim.), comb. nov., and remarks on O. subbaetica
Fig. 5. Orobanche balsensis: Spain, Málaga, Gaucín, la Almuña, 23-V-2004 by G. Moreno Moral
(herb. Sánchez Pedraja 11852 & 11853).
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Flora Montiberica 60: 38-53 (V-2015). ISSN: 1138-5952
L. CARLÓN, M. LAÍNZ, G. MORENO & Ó. SÁNCHEZ
Fig. 6. Orobanche balsensis, Ó. Sánchez Pedraja del., coram herb. Sánchez Pedraja 13841: a)
habit; b) flower, lateral view; c) corolla, frontal view; d) opened corolla showing stamens (middle
lobe of lower lip and its part of tube cut away); e) calyx segment and bract; f) pistil and stigma
(this from two different angles).
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Flora Montiberica 60: 38-53 (V-2015). ISSN: 1138-5952
The overlooked Orobanche balsensis (J.A. Guim.), comb. nov., and remarks on O. subbaetica
Orobanche subbaetica Triano & A. Puja-
das in Acta Bot. Malacitana 39: 275 (2014)
We will obviously omit our complete
description of this recently published and
very well characterized species, whose
publication as new had to be suppressed
from this very paper in the last moment
and whose chosen holotype, for which we
had even reserved a number in MA, had
been drawn in detail by one of us (fig. 7).
Such statement must be understood as a
way to highlight the soundness of the new
species by showing that it was inde-
pendently recognised in different locali-
ties by two groups of researchers. Never-
theless, we still find useful some descrip-
tive, diagnostic remarks and more im-
portantly some distributional additions.
We would summarise and somewhat
reinforce the protologue by saying that, in
addition to its peculiar host, O. subbaetica
is defined by the association of a) distinct
dorsally purplish-blue and ventrally whit-
ish corollas, b) stigma with distinctive,
deep-purple spherical lobes and c) rectan-
gle-shaped, short and divergent lower
corolla lip lobes. Other particular features
can be cited to distinguish it from other
Iberian thick-stemmed Minores: from
Orobanche castellana, specially by the
above referred lower corolla lip lobes and
stigma and by the purplish instead of
brown-reddish pigmentation; from O.
calendulae also by the denser indumen-
tum of longer hairs, the erecto-patent
flowers, the entire or strongly asymmetric
calyx and the uniformly curved dorsal line
of the corolla; from O. amethystea it can
be readily distinguished by the erecto-
patent non-geniculate flowers and the not
so longly filiform calyx teeth; from O.
santolinae, in addition to the already
mentioned traits, by the lack of the pul-
verulent tomentum (particularly apparent
in the bracts of dried plants) and by the
glabrescence of ovary, style and staminal
filaments; apart from by the unique ar-
rangement of the lower corolla lip and by
the stigma, it is distinguished from O. pi-
cridis by the uniformly curved and much
more extensively and intensely pigmented
corolla, and from O. artemisiae-campes-
tris by the lack of a straight segment in
the dorsal line of the corolla, the absence
of winish tones, the not so filiform calyx
teeth and the untwisted apical crown of
bracts.
In the populations by us studied (see
below) flowers are larger than indicated in
the protologue (18-22 mm), and anther
filaments are entirely glabrous or subgla-
brous (fig. 7). However, a comparative
glance at the protological pictures and
ours (fig. 8) suffices to dispel all doubts
about the conspecificity of all these Antir-
rhinum parasites, and don’t imply but a
somewhat larger morphological variabili-
ty within Orobanche subbaetica.
With regards to the ecology and bio-
geography of the species, we can add
Antirrhinum australe Rothm. as a host
and improve the knowledge of its distri-
butional area by indicating the following
localities, some 140 km to the southwest
from the locus classicus:
ESP, Andalua, *Cádiz: 30STF8865, Vil-
laluenga del Rosario, below Casa del Encinar,
km 5.900 of the road A-374 between Grazale-
ma and Villaluenga del Rosario, 840 m, para-
sitic on A. australe in a rocky calcareous road
bank, G. Moreno Moral MM0020/2012 (herb.
Sánchez Pedraja 13751); 30STF8563, Villa-
luenga del Rosario, base of the rocky, calcar-
eous, sunny slope of the Sierra de la Bandera,
above the Manga de Villaluenga, 880 m, close
to A. australe, 12-VI-2013, L. Carlón & G.
Moreno Moral (obs.); 30STF8970, Graza-
lema, below the town of Grazalema, km 51 of
the road A-372, 800 m, parasitic on A. austra-
le on limestone boulders at the foot of an east-
facing cliff, 12-VI-2013, L. Carlón & G.
Moreno Moral, MM0093/2013 (herb. Sán-
chez Pedraja 13881). *Málaga: 30STF9967,
Benaoján, between Benaoján and Montejaque,
580 m, parasitic on (!) A. australe, 16-V-
2011, G. Moreno Moral MM0040/2011 (herb.
Sánchez Pedraja 13675); 30SUF0069, Benao-
ján, near the Puerto de Tavizna pr. Mon-
48
Flora Montiberica 60: 38-53 (V-2015). ISSN: 1138-5952
L. CARLÓN, M. LAÍNZ, G. MORENO & Ó. SÁNCHEZ
tejaque, 730 m, parasitic on (!) A. australe,
16-V-2011, G. Moreno Moral MM0041/2011
(herb. Sánchez Pedraja 13676); 30SUF0069,
Montejaque, near Puerto de Tavizna, between
km 4 and 5 of the road MA-505, pr. Mon-
tejaque, 750 m, parasitic on A. australe in the
rocky road bank, 12-VI-2013, L. Carlón & G.
Moreno Moral, MM0092/2013 (MA 884663
ex herb. Sánchez Pedraja 13880); 30STF9867,
Montejaque, below Cerro Canchuelo, above
the path to the fountain of the Llanos de-
bar, in the outskirts of Montejaque, 730 m, pa-
rasitic on A. australe in the rocks lying at the
foot of the northern slope of the hill, 12-VI-
2013, L. Carlón & G. Moreno Moral (obs.).
Attending to the data and comments
provided by CARINE & al. (2006: 215) it
seems very likely that O. subbaetica
occurs as well on the other side of the
Strait of Gibraltar, in the climatically and
biogeographically analogous mountains
of the Western Rif, some 200 km to the
south (fig. 1). Further findings in the
Iberian Peninsula and North Africa, where
so many species of the host genus are
widespread, are to be expected.
DISCUSSION
The two above considered species are
specific parasites of perennial hosts and
presumably perennial themselves. Given
the plasticity that this crucial feature of
the life cycle shows within the subsect.
Minores of Orobanche to which both O.
balsensis and O. subbaetica undoubtedly
belong, putting their systematic and evo-
lutionary significance in its proper context
requires going to some length to under-
stand the roots of this revealing ecophysi-
ological divergence.
Generalist parasites, germinating in
response to unspecific signals, can thrive
in diverse, patchy, annual-dominated
landscapes, where, being the dispersal of
host and parasites independent, the off-
spring of successful parasites on certain
hosts is unlikely to reencounter those
hosts. It is probably no coincidence that
the most neatly unspecific Mediterranean
broomrapes carry names like Orobanche
minor and Phelipanche nana, their short
size minimising their risk of tapping to
death any annual host before completing
their own cycle. In more stable habitats
(like the steep slopes and cliffs where
Carlina occurs and the rocky outcrops
colonised by Antirrhinum) perennial, re-
liably available hosts prevail and tend to
induce the appearance of host-specific li-
neages (THOROGOOD & HISCOCK,
2010).
Population genetic evidence suggests
that host-driven diversification of Oro-
banchaceae depends on the comparative
resistance of parasites to inbreeding, giv-
en that their reduced physiology implies
fewer “fragile components” (WIC-KE &
al., 2013). A single mother plant which
results successful in a host can even self-
pollinate and found a population by pro-
ducing thousands of seeds inheriting the
ability to grow on that kind of host. Any
mutation linking the germination to the
particular exudates of that host will rapid-
ly prevail by preventing the useless pro-
duction of seeds attacking unreliable
hosts. As a result of their more complex
physiology, hosts are more vulnerable to
inbreeding depression, which slows down
their pace in the coevolutionary arms race
in such a way that comparatively resistant
strains of the host would spread only after
repeated genetic bottlenecks in the para-
site have sculpted homogeneous, recog-
nisable species. Gap-prone soils (e.g.
gypsum, scree, sand dunes, cracking clay)
further catalyse the speciation of broom-
rapes by facilitating the access of seeds to
the often compensatorily extensive root
systems of the hosts.
It remains uncertain whether broom-
rapes were at first unspecific parasites of
annuals or grew specifically on a limited
number of perennial hosts (SCHNEE-
WEISS, 2007). However, it appears more li
kely, given all the above considerations
49
Flora Montiberica 60: 38-53 (V-2015). ISSN: 1138-5952
The overlooked Orobanche balsensis (J.A. Guim.), comb. nov., and remarks on O. subbaetica
Fig. 7. Orobanche subbaetica: Sánchez Pedraja del., coram herb. Sánchez Pedraja 13880: a) habit;
b) flower, lateral view; c) corolla, frontal view; d) opened corolla showing stamens; e) calyx seg-
ment and bract; f) pistil and stigma.
50
Flora Montiberica 60: 38-53 (V-2015). ISSN: 1138-5952
L. CARLÓN, M. LAÍNZ, G. MORENO & Ó. SÁNCHEZ
Fig. 8. Orobanche subbaetica: Spain, Málaga, between Benaoján and Montejaque, 16-V-2011,
photo by G. Moreno Moral (herb. Sánchez Pedraja 13675).
51
Flora Montiberica 60: 38-53 (V-2015). ISSN: 1138-5952
The overlooked Orobanche balsensis (J.A. Guim.), comb. nov., and remarks on O. subbaetica
and the findings on other broadly Irano-
Mediterranean lineages (JABBOUR &
RENNER, 2011), that the perennial, pre-
sumably host-specific condition was first.
The two species here considered would
thus be two particular members of an ol-
der set of specific parasites from some of
which the unspecific O. minor evolved as
a response to the massive annualization of
the Mediterranean flora, rather than recent
specialised descendants of an annual, wi-
despread lineage. In this case, the lack of
molecular divergence within the Minores,
by comparison with other sections of the
genus containing as well species tightly
linked to specific hosts, would be a con-
sequence of the introgression with the
annual and widespread O. minor after this
species’ range was amplified by human
activities (THOROGOOD & al., 2009). Fur-
thermore, annualization and host-range
expansion within the Minores could arise
more than once, and the process of mo-
lecular blurring might have been mul-
tifront and thus relatively fast.
Were this the case, to their intrinsic
biological interest these modest wild
species would add a considerable agro-
nomical interest as snapshots of the evolu-
tionary background of actual or potential
pest species like O. crenata and O. minor
itself. In fact, the molecular analysis by
THOROGOOD & al. (2009), in agreement
with an obvious morphological similarity,
places O. balsensis in the very core of a
widespread generalist O. minor lineage,
although differentiated from it. Its long
branch in the phylogram is interpretable
in the light of the above considerations as
the result of successive population bottle-
necks through the adaptation to a specific
host, the homogeneity of O. minor and its
lesser divergence from their putative com-
mon ancestor being attribuable to the lack
of selective forces combined with larger
populations.
Acknowledgements
Our deepest gratitude goes to Ana Isabel
Correia, who tirelessly searched for us in LISU
the Welwitch’s materials, buried after more
than a century of fluctuant revisions, that made
possible the resurrection of the name Oroban-
che balsensis. The help in this task of Carlos
Aedo and Jorge Paiva must also be sincerely
acknowledged. Without the assistance of Juan
Antonio García Rojas and Felipe Muñoz Secil-
la, skillful botanists from San Roque (Cádiz),
it would have been impossible to find one of
the Spanish populations of what happened to
be O. balsensis. Alfonso La Rosa, Corrado
Marcenò and Francesco Russo, prominent
Sicilian botanists, kindly sent us suggestive
photographs: theirs is now the duty to check
whether O. balsensis does actually grow in the
“bella Trinacria”, and to what extent.
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Flora Montiberica 60: 38-53 (V-2015). ISSN: 1138-5952
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