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Molecular Data Helps Traditional Taxonomy: Re-evaluation of Reseda collina (Resedaceae), and a New Record for Europe

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Reseda collina is included in Reseda section Phyteuma, a monophyletic and morphologically well-defined group that is also characterized by taxonomic problems regarding species circumscription. The status of this species has been controversial, usually being subordinated to the widespread R. phyteuma. A study of R. collina and allied taxa based on morphological and molecular (nuclear ITS, plastid trnL-F) data supports the taxonomic identity of R. collina and its separation from R. phyteuma. An epitype is designated to support the holotype and prevent future nomenclature problems. In addition, our molecular and morphological analyses suggest that some problematic southeastern Iberian populations, previously identified as R. phyteuma, belong to R. collina, which implies a new European record for this species that was previously considered as a northwestern African endemic. Following this finding, biogeographic relationships between the southern Iberian Peninsula and northwestern Africa are also discussed.
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Molecular Data Helps Traditional Taxonomy:
Re-evaluation of Reseda collina (Resedaceae),
and a New Record for Europe
Santiago Martín-Bravo &Pedro Jiménez-Mejías
#Institute of Botany, Academy of Sciences of the Czech Republic 2009
Abstract Reseda collina is included in Reseda section Phyteuma, a monophyletic
and morphologically well-defined group that is also characterized by taxonomic
problems regarding species circumscription. The status of this species has been
controversial, usually being subordinated to the widespread R. phyteuma. A study of
R. collina and allied taxa based on morphological and molecular (nuclear ITS,
plastid trnL-F) data supports the taxonomic identity of R. collina and its separation
from R. phyteuma. An epitype is designated to support the holotype and prevent
future nomenclature problems. In addition, our molecular and morphological
analyses suggest that some problematic southeastern Iberian populations, previously
identified as R. phyteuma, belong to R. collina, which implies a new European
record for this species that was previously considered as a northwestern African
endemic. Following this finding, biogeographic relationships between the southern
Iberian Peninsula and northwestern Africa are also discussed.
Keywords Arid regions .Ibero-North African endemic .Molecular systematics .
Reseda section Phyteuma .Taxonomy.Typification
Introduction
The genus Reseda L. is composed of ca. 65 species distributed in temperate areas of
the Old World, with a center of diversity around the Mediterranean basin. Many of
its species grow on basic soils of arid habitats, others are ruderal weeds, and a few
are confined to high mountains. The taxonomy of the genus is complex, and mature
individuals are required to identify most species. Apart from the numerous
treatments in local floras, the most comprehensive taxonomic accounts of the
Resedaceae were published by Müller Argoviensis (1857,1868), and Abdallah and
Folia Geobot (2009) 44:399421
DOI 10.1007/s12224-009-9052-y
S. Martín-Bravo (*):P. Jiménez-Mejías
Department of Molecular Biology and Biochemical Engineering, Pablo de Olavide University,
ctra. de Utrera km 1, 41013 Sevilla, Spain
e-mail: smarbra@upo.es
de Wit (1978). In addition, a study of phylogenetic relationships and biogeography
of the family based on molecular markers (ITS, trnL-F) was recently published
(Martín-Bravo et al. 2007).
Reseda sect. Phyteuma Lange is the second largest section of the genus (14 species,
Table 1), after sect. Reseda. Most species are restricted endemics distributed in
northwestern Africa or the southeastern Mediterranean area (Table 1). This section
constitutes a monophyletic group, as revealed by previous molecular phylogenetic
analyses (Martín-Bravo et al. 2007). Morphologically, Reseda sect. Phyteuma is one of
the most homogeneous and well-defined sections of the genus (Abdallah and de Wit
1978), and is characterized by pendulous ripe capsules and relatively large seeds
generally with undulate-rugose testa (although papillose testa is found in R.
stenobotrys and rarely also in R. jacquinii). However, this morphological homogeneity
is also reflected in the small degree of differentiation between the taxa, which has
caused problems in species circumscription. In addition, hybridization processes seem
to exist between distinct species, as shown by the fact that one taxon (R. odorata L.) is
probably of hybrid origin (Abdallah and de Wit 1978;Martín-Bravoetal.2007).
Reseda collina J. Gay is a species included in Reseda sect. Phyteuma,andhassofar
been considered a northwestern African endemic (northern Algeria and Morocco). It
was originally described together with R. duriaeana J. Gay (sect. Reseda), in an
illustration with analysis (Fig. 1) included in the Atlas volume of Exploration
Scientifique de lAlgérie (Bory and Durieu 1849). Although the plate did not include
any written description or diagnosis, both species were effectively and validly
published under the rules of the International Code of Botanical Nomenclature (ICBN,
Art. 44.1, McNeill et al. 2006). Soon afterwards, Müller Argoviensis (1856,1857,
1868) performed the first descriptive studies on R. collina, based on a few specimens
collected around Oran (Algeria), the only place with known populations at that time.
He stressed the affinity of R. collina to the widespread and morphologically variable
R. phyteuma L., and reported some diagnostic characters for their distinction (shape of
petals, staminal filaments, calyx accrescence). Subsequently, Durand and Schinz
(1898)combinedR. collina as a subspecies of R. phyteuma (R. phyteuma subsp.
collina (J. Gay) Durand & Schinz). This subspecific rank has generally been preferred
by subsequent authors (Jahandiez and Maire 1932; Quézel and Santa 1962;Maire
1976; Abdallah and de Wit 1978;IbnTattou1999; Valdés 2002;MateosandValdés
2003). In addition, these studies expanded the range of the taxon to some regions of
northern Morocco. Recent molecular data, however, suggested the separation of R.
collina from R. phyteuma (Martín-Bravo et al. 2007).
The presence of R. collina in the Iberian Peninsula has barely been discussed to
date. The species has not been recorded either in Flora Europaea (Yeo 1964,1996)or
in Flora Iberica (Valdés Bermejo 1993). Previously, Lange (1880)includeditunder
the list of species of possible but not confirmed presence. Abdallah and de Wit (1978)
indicated the presence of this taxon (subordinated to R. phyteuma) in Spain, but based
on a single collection in northeastern Spain (Aragón, prope Castelserás, 15 Apr 1875,
F. Lo sc os 1 5 (W)), which belongs to R. phyteuma s. str. (Aránega and Pajarón 1990).
To our knowledge, the first reliable reference to the presence of the species in Spain is
in a collection belonging to Hno. Jerónimo in southeastern Spain (Almería, Sierra de
Gádor, Berja, 21 Jun 1934 (MA)), labelled as R. collina Gay, Nouveau pour
lEspagne. This chorological novelty was published as a doubtful record (Sennen
400 S. Martín-Bravo, P. Jiménez-Mejías
1936; nr. 9107), although it did not become widely known, probably because of the
doubtful taxonomic value of the species at that time. Afterwards, some authors saw
Jerónimo´s exsiccate and subsequently listed the taxon (as R. phyteuma subsp. collina)
in their catalogues of vascular plants of the Almería province (Losa España and Rivas
Goday 1974;Sagredo1987), although they did not perform any taxonomic study.
Finally, Valdés Bermejo, in the observations about R. phyteuma in Flora Iberica
(Valdés Bermejo 1993), quoted the existence of material from the Almería province
that showed anomalous characters (e.g., non-dilated staminal filaments) and did not
match R. phyteuma morphology. He also wrote such taxonomic comments on some
problematic sheets from Almería housed in the MA herbarium (MA 323143, MA
329711). However, the small amount of material that he studied was that of immature
individuals. He preferred to not draw taxonomic conclusions on these anomalous
populations, but adopted a synthetic approach, identifying them as the common and
widespread R. phyteuma.
The taxonomic revision of Resedaceae for the Flora de Andalucía Oriental
project (Martín-Bravo 2009) resulted in the finding of several sheets from the
southeastern Iberian Peninsula (Spain, Almería province), previously identified as R.
phyteuma, but morphologically similar to R. collina. The presence of R. collina in
Spain would constitute a new record for Europe. This fact, together with the current
taxonomic controversy, encouraged us to perform a molecular and morphological
study to elucidate the taxonomic status of R. collina, and to confirm its presence in
the southeastern Iberian Peninsula.
Material and Methods
Molecular Study
Twenty species of Resedaceae were included in the molecular study (Table 1). We
sequenced 34 accessions for the ITS region of nrDNA, and 31 accessions for the
trnL-F intergenic spacer of cpDNA. Twelve out of 14 species of Reseda sect.
Phyteuma were analyzed. We were unable to obtain material from the very rare and
restricted endemics R. balansae (southern Turkey) and R. stenobotrys (southwestern
Morocco). Five populations of R. collina were sampled and analyzed, two from
northwestern Africa (Morocco), and three from the southeastern Iberian Peninsula
(Table 1). We failed to amplify the trnL-F sequence of one Iberian population
(population 5 in Table 1). Accessions of the genera Caylusea and Sesamoides were
used as outgroups. Most sequences (29 ITS, 22 trnL-F) presented here were taken
from the previous phylogenetic study (Martín-Bravo et al. 2007). Here we expanded
the sampling of Reseda sect. Phyteuma by obtaining five additional ITS and nine
trnL-F sequences for seven species (Table 1), including one species previously not
sampled (R. tymphaea). These new sequences were obtained from silica gel-dried
material collected in the field or from herbarium specimens (GDA, HUAL, RNG,
UPOS, UPS; see Holmgren et al. 1990 for herbarium abbreviations). Procedures of
DNA extraction, amplification and sequencing followed those described in Martín-
Bravo et al. (2007). The sequences obtained were deposited in the GenBank
database (Table 1).
Taxonomic re-evaluation of Reseda collina 401
Table 1 List of plant material included in the molecular study
Taxon Distribution Locality Voucher GenBank ITS/
trnL-F accession number
Outgroup
Caylusea hexagyna (Forssk.) M.L. Green N Africa, SW Asia Cape Verde, Sal Island, Santa Maria H. Rustan 2653 (O) ITS (DQ987225)
trnL-F (DQ987069)
Sesamoides interrupta (Boreau) G. López Pyrenees, S France, Italy, Corsica,
Sardinia
Andorra, Lacs du Pesons J.M. Marín 12304JMM (UPOS) ITS (DQ987214)
trnL-F (DQ987065)
Oligomeris Cambess.
O. linifolia (Vahl) J.F. Macbr. N Africa, SW Asia, SW North America Yemen, Arhab J.R. Wood 2057 (BM) ITS (FJ212195)
trnL-F (FJ212275)
Reseda L.
sect. Glaucoreseda DC.
R. complicata Bory S Spain (Sierra Nevada) Spain, Granada, Sierra Nevada S. Martín-Bravo 62SMB04 (UPOS) ITS (DQ987172)
trnL-F (DQ987046)
sect. Leucoreseda DC.
R. undata L. subsp. undata C, E Spain Spain, Madrid, Rivas S. Martín-Bravo 39SMB04 (UPOS) ITS (DQ987203)
trnL-F (DQ987056)
sect. Luteola Dumort.
R. luteola L. Mediterranean Greece, Crete S. Martín-Bravo 391SMB05 (UPOS) ITS (DQ987160)
trnL-F (DQ987010)
sect. Reseda L.
R. elata Coss. ex Müll. Arg. SW Morocco Morocco, Agadir, Ait Abdallah C. Blanché s.n. (MA) ITS (DQ987172)
trnL-F (DQ987046)
R. viridis Balf. f. Socotra Island Yemen, Socotra Island A.N. Gifri 8701 (UPS) ITS (DQ987130)
trnL-F (DQ986996)
sect. Phyteuma Lange
R. alopecuros Boiss. SE Mediterranean Region Israel, Golan A. Liston s.n. 1984 (HUJ) ITS (DQ987139)
trnL-F (DQ987028)
R. arabica Boiss. N Africa, SW Asia Morocco, Errachidia, Erfoud (1) J. Lambinon 95/Ma/265 (RNG) ITS (DQ987131)
trnL-F (FJ477077)#
402 S. Martín-Bravo, P. Jiménez-Mejías
Table 1 (continued)
Taxon Distribution Locality Voucher GenBank ITS/
trnL-F accession number
Kuwait, Khabrat Al-Awazen (2) R. Halgawy 1183 (UPS) ITS (DQ987132)/
trnL-F (DQ987029)
R. armena Boiss. Turkey, Armenia Turkey, Mut M. Nydegger 47327 (MSB) ITS (DQ987143)
trnL-F (DQ987023)
R. collina J. Gay N Morocco, N Algeria, SE Spain Morocco, Beni-Snassen, Taforalt (1) S. Martín-Bravo 12SMB04 (UPOS) ITS (DQ987135)
trnL-F (FJ477074)#
Morocco, Kassita (2) S. Martín-Bravo 17SMB04 (UPOS) ITS (DQ987136)
trnL-F (DQ987031)
Spain, Almería, Sierrecilla (3)* E. Giménez s.n. (HUAL) ITS (FJ477066)#
trnL-F (FJ477075)#
Spain, Almería, Dalías (4)* A. Belén s.n. (GDA) ITS (FJ477067)#
trnL-F (FJ477076)#
Spain, Almería, Sierra de Gádor (5)* E. Giménez s.n. (HUAL) ITS (FJ477068)#
R. diffusa (Ball) Ball Morocco Morocco, Ourika (1) L. Jonsell 5413 (UPS) ITS (DQ987140)
trnL-F (FJ477073)#
Morocco, Tiznit (2) D. Podlech 48412 (MSB) ITS (DQ987141)
trnL-F (DQ987033)
R. inodora Rchb. SE Europe Rumania, Dobrudza F. Cernoch 24622 (BRNM) ITS (DQ987142)
trnL-F (DQ987030)
R. jacquinii Rchb. NE Spain, SE France France, Ardèche C. Simon s.n. 1963 (HBG) ITS (DQ987144)
R. media Lag. W Iberian Peninsula, NW Morocco,
Azores, Madeira
Spain, Cádiz, El Aljibe (1) P. Jiménez-Mejías 106PJM05 (UPOS) ITS (DQ987149)
Portugal, Beira Litoral, Foz de Arouce (2) R. Svensson 756 (UPS) ITS (DQ987150)
trnL-F (FJ477070)#
Spain, Huelva, Doñana (3) S. Martín-Bravo 156SMB05 (UPOS) ITS (DQ987151)
trnL-F (DQ987022)
R. odorata L. cultivated, probably native in
Lybia and Crete
Germany, Bayern (cultivated) (1) Eschelmüller s.n. 1998 (M) ITS (DQ987133)
trnL-F (DQ987026)
Ireland, Dublin, (cultivated) (2) D. Flynn s.n. 1999 (DBN) ITS (DQ987134)
trnL-F (DQ987027)
Lybia, Ras Hilal (3) Davis 50170 (RNG) ITS (FJ477069)#
trnL-F (FJ477078)#
R. orientalis (Müll. Arg.) Boiss. E Mediterranean Cyprus, Paphos (1) E. Julin s.n. 1972 (UPS) ITS (DQ987137)
trnL-F(DQ987025)
Taxonomic re-evaluation of Reseda collina 403
Table 1 (continued)
Taxon Distribution Locality Voucher GenBank ITS/
trnL-F accession number
Cyprus, Nicosia (2) J. Lambinon 92/Cy/406 (MSB) ITS (DQ987138)
trnL-F (DQ987024)
R. phyteuma L. W, C Mediterranean, C Europe Spain, Seville, Casariche (1) S. Martín-Bravo 26SMB04 (UPOS) ITS (DQ987145)
trnL-F (FJ477071)#
Morocco, Chefchaouen (2) S. Martín-Bravo 23SMB04 (UPOS) ITS (DQ987146)
trnL-F (DQ987032)
Spain, Valencia, Chelva (3) M. Nydegger 35720 (MSB) ITS (DQ987147)
trnL-F (DQ987034)
Spain, Murcia, Lorca (4) D. Podlech 25374 (MSB) ITS (DQ987148)
trnL-F (DQ987035)
R. tymphaea Hausskn. S Greece, NW Turkey Turkey, Çankiri S. Martín-Bravo 154SMB06 (UPOS) ITS (FJ477065)#
trnL-F (FJ477072)#
Not sampled species
R. balansae Müll. Arg. SW Turkey
R. stenobotrys Maire & Sam. SW Morocco
Herbarium abbreviations follow Holmgren et al. (1990). Localities of R. collina marked with asterisks represent new records from southeastern Spain. Multiple populations of a
given species are numbered (numbers in brackets). GenBank accession numbers with the symbols # indicate new sequences obtained in this study; otherwise they were taken
from the study by Martín-Bravo et al. (2007). Not sampled species of Reseda sect. Phyteuma are also listed
404 S. Martín-Bravo, P. Jiménez-Mejías
Maximum parsimony (MP) and Bayesian Inference (BI) analyses were
performed, with the settings as given in Martín-Bravo et al. (2007). Two matrices
were assembled, a matrix of ITS alignment (34 sequences) and of trnL-F alignment
(31 sequences). Gaps were treated as missing data and coded as additional
characters. Only informative indels were coded; mononucleotide repeat units
Fig. 1 Holotypes of Reseda duriaeana J. Gay (left) and R. collina J. Gay (right) published in Exploration
Scientifique de l´Algérie (Bory and Durieu 1849)
Taxonomic re-evaluation of Reseda collina 405
(poly-T and poly-A) were excluded due to their high homoplasy (Kelchner 2000).
When employing BI analyses, the F81 model of sequence evolution was used for the
indel characters, following the manual of MrBayes version 3.0b4 (Ronquist and
Huelsenbeck 2003). Congruence of ITS and trnL-F data sets was assessed using the
Hompart test for matrices, and the Kishino-Hasegawa and Shimodaira-Hasegawa
tests for topology (1,000 replicates each), as implemented in PAUP* version 4.0b10
(Swofford 2002).
Morphological Study
Representative herbarium material of R. collina (66 specimens; see Appendix 1), R.
phyteuma (264 specimens; not listed) and allied species of Reseda sect. Phyteuma
was studied (ALME, AMD, BM, DBN, E, GDA, GH, HUAL, IBF, K, MA, MGC,
MPU, O, RNG, S, SEV, UPOS, UPS). We made a special effort to examine the most
important morphological characters for the taxonomy of sect. Phyteuma (Müller
Argoviensis 1856,1857,1868; Abdallah and de Wit 1978; Valdés Bermejo 1993):
calyx accrescence (size of calyx in flower vs in fruit), shape of petals (especially with
respect to the laciniae of the lateral lobes of superior petals), staminal filaments
(distal part widened or not) and seeds (length, ornamentation of testa, presence of
apical carunculoid tissue, color when ripe, shininess). A Nikon SMZ645 stereo-
scopic microscope and a Hitachi S3000-N electron microscope were used to study
the herbarium specimens. Gold coating was applied to the samples examined using
scanning electron microscopy (SEM). The distribution of both taxa was revised,
using TDWG geographical codes at level 3 (Botanical countries) for specifying the
ranges (Brummit 2001).
Results
Phylogenetic Analyses
The ITS sequence length in the studied accessions of Reseda sect. Phyteuma was
627634 base pairs (bp); the length of trnL-F sequences was 704808 bp. For the
ITS matrix, 84 variable sites were detected within sect. Phyteuma (one variable site
in R. collina sequences), of which 73 were parsimony-informative (none in R.
collina). For the trnL-F matrix, 24 sites were variable (two variable sites in R.
collina), and 12 of them were parsimony-informative (none in R. collina). Twelve
informative indels of 14 bp length were coded as additional characters in the ITS
matrix, of which three deletions were found within R. collina sequences. Five
informative indels of 573 bp length were coded in the trnL-F matrix, of which no
deletion was found within R. collina sequences.
Phylogenetic reconstructions of the ITS matrix (aligned length 662 bp; 12 coded
indels) based on MP resulted in 19 trees of 364 steps (CI excluding uninformative
characters (e.u.c)=0.66; RI=0.82; RC=0.60; trees not shown). MP analysis of the
trnL-F matrix (aligned length 838 bp; five coded indels) resulted in a single tree of
120 steps (CI e.u.c=0.75; RI=0.88; RC= 0.76; tree not shown). Phylogenetic
analyses of the plastid (trnL-F) and nuclear (ITS) matrices generated significantly
406 S. Martín-Bravo, P. Jiménez-Mejías
different topologies (Fig. 2; Kishino-Hasegawa and Shimodaira-Hasegawa tests,
P=0.025), and the Hompart test showed that the data sets were incongruent
(P=0.01). Therefore, we did not perform phylogenetic analyses based on a
combined ITS+trnL-F matrix.
The hierarchical Likelihood Ratio Test (hLRT) and the Akaike Information
Criterion (AIC), as implemented in MrModeltest version 1.1b (Nylander 2002),
retrieved different models of sequence evolution for the ITS 1 (SYM +G; AIC =
2735.05), ITS 2 (HKY+G and GTR+G, respectively; AIC =2083.35), and the 5.8 S
region (JC; AIC=515.34). The models F81 + G and GTR +G (AIC =3761.48) were
retrieved for the trnL-F matrix by the hLRT and the AIC, respectively. The tree
topologies and clade supports obtained with the different models as suggested by
hLRT and AIC, however, were very similar for both ITS and trnL-F matrices. The
trees shown here are those based on the evolutionary models selected by the AIC,
following the recommendations of Posada and Buckley (2004).
Bayesian majority rule consensus trees obtained from the phylogenetic analyses
of the single matrices with coded indels (ITS, trnL-F; Fig. 2) were mostly consistent
with the corresponding strict consensus trees of the MP analyses (not shown). When
performing MP and BI analyses of the same ITS and trnL-F matrices but without
coded indels, the tree topology was similar but less resolved, and the clade support
was slightly lower, due to the lower number of informative characters (trees not
shown). Reseda sect. Phyteuma was resolved as a monophyletic group with low to
moderate support in both ITS and trnL-F analyses (91%100% posterior probability,
PP; 65%66% bootstrap support, BS). These results are congruent with previous
molecular studies (Martín-Bravo et al. 2007). Resolution and clade support within
sect. Phyteuma were lower in the trnL-F analysis when compared with the ITS
analysis, due to the lower number of informative characters. Our results confirm the
distinction of R. collina from R. phyteuma, because they are placed in different
clades within sect. Phyteuma in both analyses (Fig. 2). ITS sequences of the
southeastern Iberian populations of R. collina grouped together with those of the
Moroccan R. collina, forming a strongly supported clade (100% PP; 94% BS).
Monophyly of R. collina was thus strongly supported by the ITS data, but not by the
trnL-F data, in which the corresponding sequences appeared in an unresolved
polytomy, together with the accessions of R. diffusa and R. arabica.
Taxonomy
We present here an identification key to the eight species of Reseda sect. Phyteuma
that occur in the western Mediterranean basin. Detailed morphological descriptions
of R. phyteuma and R. collina are given (see below), considering also the variation
of southeastern Iberian populations of R. collina. The most important morphological
characters distinguishing these two species are calyx accrescence, shape of superior
petals, staminal filaments and inflorescence apex, color of the corolla, and
ornamentation and presence of apical carunculoid tissue in seeds (Table 2; Fig. 3).
Both species also show different distribution patterns. While R. phyteuma is widely
distributed in the western and central Mediterranean region and in Central Europe, R.
collina is restricted to the western Mediterranean, with a disjunct distribution in
southeastern Spain and northwestern Africa (northeastern Morocco, northwestern
Taxonomic re-evaluation of Reseda collina 407
Fig. 2 Comparison of the two majority rule consensus tree obtained from the Bayesian analyses of ITS
and trnL-F matrices with coded indels. Bayesian posterior probabilities are given above the branches;
bootstrap values from the corresponding maximum parsimony consensus trees are given below the
branches. Asterisks indicate samples for which we failed to amplify the plastid trnL-F region. Reseda
collina samples from northern Africa and southeastern Spain are highlighted in separate boxes
408 S. Martín-Bravo, P. Jiménez-Mejías
Algeria; Fig. 4). In addition, a nomenclatural survey was performed, including the
designation of the epitype of the name R. collina, to prevent future nomenclatural
problems (see Discussion).
Discussion
Taxonomic Status of Reseda collina
The case of Reseda phyteuma and R. collina is an illustrative example of taxonomic
problems common to the Reseda sect. Phyteuma; while these species appear to be
clearly separated from a phylogenetic point of view (Fig. 2; Martín-Bravo et al. 2007),
their morphological differentiation is low and diagnostic characters are difficult to
observe. In addition, both species often coexist (e.g., Oran, Algeria; Beni-Snassen
mountains, northeastern Morocco), but they do not seem to hybridize, which is
congruent with their relatively distant phylogenetic positions. A compilation of
diagnostic characters that can be used to discriminate between the two species is given
in Table 2and Fig. 3, based on the study of representative herbarium material.
Reseda collina in the Iberian Peninsula
The comparative taxonomic study of specimens of the North African R. collina
and the material from southeastern Iberian Peninsula showed that some Spanish
populations indeed belong to R. collina. The ITS sequences of the southeastern
Iberian populations clustered with those from Morocco with a strong support,
consistently suggesting that they are conspecific (Fig. 2). The unresolved position
of R. collina sequences in the trnL-F analysis was due to the low number of
informative characters in the plastid data set, rather than due to incongruence.
Nonetheless, certain topological discordances between nuclear and plastid
phylogenies reflect the lack of congruence between both data sets and may be
Table 2 Diagnostic morphological characters of Reseda phyteuma and R. collina
Character Reseda phyteuma Reseda collina
Distal part of the
inflorescence
More or less lax, inflorescence apex blunt Generally dense, inflorescence
apex acute
Calyx accrescence Sepals usually conspicuously accrescent
in fruit, as long as or longer than the
half of ripe capsule
Sepals slightly accrescent
in fruit, as long as or shorter
than the half of ripe capsule
Corolla White or yellowish when dry Creamy yellow, sometimes
with orange dots
Superior petals shape Laciniae generally linear Laciniae with apex generally
spathulate
Staminal filaments Distinctly widened in the distal part Not widened in the distal part
Seeds Testa with more or less sharp ridges,
apical carunculoid tissue absent
or minute
Testa with blunt ridges, apical
carunculoid tissue usually
conspicuous, subconical
Taxonomic re-evaluation of Reseda collina 409
Fig. 3 Detailed photographs of some diagnostic characters distinguishing Reseda phyteuma (left column
a, c, e, g) and R. collina (right column b, d, f, h), taken with a digital camera (a, b), a stereoscopic
microscope (c, d) or an electron miscroscope (eh). a, b calyx and capsule; c, d superior petal; e, f
stamen; g, h seed. Arrow in picture hindicates the apical carunculoid tissue typical of R. collina. Scale
bars 5mmina, b; 1 mm in c, d; 0.5 mm in eh. Specimens used are listed in Appendix 2
410 S. Martín-Bravo, P. Jiménez-Mejías
due to hybridization processes within Reseda sect. Phyteuma (Fig. 2; Martín-Bravo
et al. 2007).
The presence of R. collina in the southeastern Iberian Peninsula is a new record of
this taxon for Europe, which was previously considered to be a northwestern
African endemic (Müller Argoviensis 1857,1868; Durand and Schinz 1898;
Jahandiez and Maire 1932; Maire 1976; Valdés 2002), and it considerably increases
the species range (Fig. 4). Thus, the species should be considered an Ibero-North
African endemic. The Ibero-North African floristic element is well represented in the
flora of the southern Iberian Peninsula, and accounts for 10%13% of the vascular
flora of Andalusia (Blanca et al. 1999). Southern Iberia and northern Morocco show
a close biogeographical relationship (Médail and Quézel 1997) with more than 500
endemic species (Quézel 1978) and about 75% of 3,500 species shared (Valdés
1991). The two regions were connected from the end of the Miocene to the upper
Pliocene due to the partial desiccation of the Mediterranean Sea following an
increase in aridity (Messinian crisis, 5.54.5 million years ago; Hsü et al. 1977;
Duggen et al. 2003), allowing contact between the floras of the two continents. The
southeastern Iberian region, where the populations of R. collina are located, is the
most arid part of Europe, and shares climatic and ecological affinities with subdesert
areas of North Africa. This biogeographical relationship is also reflected in the
abundance of Ibero-North African species in southeastern Spain (Cueto et al. 1991;
Giménez Luque and Gómez Mercado 2002; Peñas de Giles et al. 2006). Interestingly,
two other congeneric species included in sect. Reseda (R. stricta Pers., R. lanceolata
Lag.), are also Ibero-North African endemics, which show not only a similar range to
R. collina, but also close ecological requirements (arid basic soils).
The polytomies, in which the accessions of R. collina are embedded (Fig. 2),
prevent us from establishing phylogenetic relationships between the Iberian and
African populations. Further work and more variable markers (e.g., AFLPs) are
needed to investigate the biogeography and evolutionary history of R. collina.
Typification Problems
As stated above, R. collina was described in 18461847 (cf. Abdallah and de Wit
1978) by J. Gay in a detailed color plate that included analytical drawings but no
Fig. 4 Geographical distribution of Reseda collina. Solid dots depict the localities of the studied
herbarium specimens; the empty dot depicts a population reported by Mateos and Valdés (2003)
Taxonomic re-evaluation of Reseda collina 411
Fig. 5 Reseda collina sheet (K 230987) used as the model for part of the original description by J. Gay
412 S. Martín-Bravo, P. Jiménez-Mejías
written diagnosis (Fig. 1). This illustration must be considered the original
description, and therefore the holotype (cf. Cheek 1989;ICBN, Arts. 42.3, 42.4,
44.1; McNeill et al. 2006). There is a specimen from Algeria deposited in Kew
herbarium (Oran, collines pierreuses de l´Ouest, 27 Apr 1842, Durieu (K 230987);
Fig. 5), which was used as the model for part of Gays plate, specifically for the
detailed drawings of petals, calyx and ovary. This fact probably motivated M.
Abdallah to label the sheet as the type of R. collina when he revised it in 1967
(Fig. 5). However Gays plate was subsequently explicitly cited as the only type
material in the monograph (Abdallah and de Wit 1978). Sheet K 230987 (Fig. 5)
does contain a R. collina specimen, but it is a markedly immature individual, and
therefore the plant with mature capsules represented in the plate (Fig. 1) was most
likely drawn from another currently unknown specimen. In our opinion, the plant
represented in the plate is not unequivocally assignable to R. collina, and could be
identified as an allied species such as R. phyteuma, because not all necessary
characters used to distinguish among them (Table 2) are clearly illustrated. Whereas
a superior petal is depicted in an analytical drawing, other important characters such
as petal color, inflorescence apex, staminal filaments and seeds are not represented in
detail (Fig. 1). The latter two characters were illustrated in the analysis by Abdallah
and de Wit (1978) in their monograph of the family (Fig. 6).
Fig. 6 Analytical drawings of superior petal, stamen and seed of Reseda phyteuma and R. collina, taken
from Abdallah and de Wit (1978)
Taxonomic re-evaluation of Reseda collina 413
Because the holotype may be ambiguous regarding reliable identification, it is
desirable to designate an epitype to ensure the precise application of the species
name (ICBN, Art. 9.7; McNeill et al. 2006). This would allow better definition of
diagnostic characters, and prevent problems with the identity of the species. Sheet K
Fig. 7 The epitype of Reseda collina J. Gay designated here (UPOS 3354)
414 S. Martín-Bravo, P. Jiménez-Mejías
230987 (Fig. 5), which was the model for part of the plate, appears to be an
inappropriate epitype due to its immature inflorescence. Because this sheet does not
help to determine the application of the species name (ICBN, Rec. 8A1; McNeill et
al. 2006), we propose the specimen UPOS 3354 (Morocco, Beni-Snassen mountains,
Taforalt, 20 Apr 2004, S. Martín Bravo; Fig. 7), as the epitype. This specimen shows
all the morphological characters that help to discriminate R. collina from R.
phyteuma and other allied species (Table 2; Fig. 3). In addition, its identity has been
confirmed in our molecular study, because a sample from this population was
included in the analyses (population 1; Table 1).
Identification Key, Nomenclature Survey and Morphological Descriptions
The following identification key helps to identify the species of Reseda sect.
Phyteuma that are present in the western Mediterranean basin.
1a Lateral lobes of superior petal lunate or falcate, with entire to palmatifid margin.
.....................................................R. jacquinii
(region around Gulf of Leon: NE Spain and SE France)
1b Lateral lobes of superior petal palmatisect, deeply laciniate. . . . ............2
2a Seeds with papillose testa, up to 1.5 mm; capsules (3)3.54.5 mm wide. . . . . .
....................................................R. stenobotrys
(SW Morocco)
2b Seeds with undulate-rugose testa, 1.5 mm or longer; capsules 410 mm wide. . . . 3
3a Staminal filaments persistent in fruit; ripe capsule sessile. . . . . . . . . . R .arabica
(N Africa, SW Asia)
3b Staminal filaments deciduous in fruit; ripe capsule generally estipitate. . . . . . 4
4a Superior petals 23 mm, shorter than stamens; seeds glossy........R. diffusa
(SW Morocco)
4b Superior petals 36 mm, usually longer than stamens; seeds dull. ..........5
5a Ripe capsules shorter than 10 mm; seeds 1.51.8mm............R. odorata
(cultivated, rarely escaping from culture, widespread; probably native in Libya
and Crete, where it is apparently wild)
5b Ripe capsules generally longer than 10 mm; seeds 2 mm or longer. .........6
6a Sepals accrescent in fruit, as long as or longer than half of the ripe capsule;
laciniae of superior petals usually linear; staminal filaments generally distinctly
widened in the distal part. ................................R. phyteuma
............................(W,CMediterranean region, S, C Europe)
6b Sepals not or slightly accrescent in fruit, as long as or shorter than half of the
ripe capsule; laciniae of superior petals with apex generally spathulate; staminal
filaments not or scarcely widened in the distal part. .....................7
7a Sepals oblong or spathulate; petals creamy yellow, sometimes with orange dots;
top of the inflorescence generally dense, with acute apex. . . . . . . . . . R. collina
(SE Spain, NW Africa)
7b Sepals linear or linear-lanced; petals white; top of the inflorescence more or less
lax, with blunt apex. . . . . . . . ................................R. media
(W Iberian Peninsula, Azores, Madeira, restricted in NW Morocco)
Taxonomic re-evaluation of Reseda collina 415
Reseda collina J. Gay in Bory & Durieu, Expl. Sc. Alger. t. 71. f. 2. 184647.
Ind. loc.: Algeria (not explicit).
Holotype: J. Gay, Expl. Sc. Alger. t. 71. f. 2. 184647.
Epitype (designated here): Marruecos, Beni-Snassen, subiendo hacia el puerto de
Taforalt, bosque de Tetraclinis articulata sobre calizas, taludes junto a la carretera,
34º5015′′ N, 2º2442′′ W, 495 m, 20 Apr 2004, S. Martín Bravo 12SMB04 (UPOS
3354; Fig. 7). Isoepitypes: G, K, MA, RNG.
Icon.: Gay in Bory & Durieu, Expl. Sc. Alger. t. 71, f. 2. 184647 (holotype);
Abdallah and de Wit in Meded. Landbouwhogesch. Wageningen 78(14), Atlas,
fig. 71. 1978 (sub R. phyteuma subsp. collina); Maire in Fl. Afrique N. 14: 189
(1976) (sub R. phyteuma subsp. collina).
Reseda phyteuma subsp. collina (J. Gay) Durand & Schinz, Consp. Fl. Afr. 1(2):
184. 1898.
Morphological description: Perennial herb, 1050 cm tall. Stems ascending, rarely
erect, branching from the base, glabrous or papillose in the ribs. Basal leaves entire
or subentire, sometimes rosette-like; stem leaves alternate, usually 3- or 5-lobed,
terminal lobe sometimes 3-lobed in turn; glabrous or papillose. Inflorescence
spiciform, generally dense at the top, with acute apex. Pedicels 410 mm in flower,
up to 15(18) mm in fruit. Flowers bisexual, 6(7)-merous. Calyx dialysepalous,
sepals 24(6)×0.51.5 mm in flower, persistent, slightly accrescent, reflexed in
fruit, 36(8)×0.51.5(2.5) mm, as long as or shorter than the half of the ripe
capsule, oblong or spathulate. Corolla dialypetalous, heteromorphic, creamy yellow,
sometimes with orange dots; superior petals 34(5) mm long, unguiculate, trisect,
lateral lobes palmatisect, each deeply divided in (4)57(8) spathulate or linear-
spathulate laciniae, rarely linear, central lobe linear or linear-spathulate, shorter than
lateral lobes; lateral and anterior petals reduced. Stamens 1924, filaments
deciduous, not widened in the distal part, connate at the base to form an
asymmetrical nectariferous disc. Capsules 3-carpelled, 818× 510 mm, estipitate,
pendulous when ripe, obovate-truncate to subglobose, glabrous, sometimes papillose
in the ribs, capsule teeth 0.51.5 mm long. Seeds 22.5 mm, reniform, brownish
when ripe, dull, testa undulate-rugose, with blunt ridges, apical carunculoid tissue
usually conspicuous, subconical.
Ecology: Basic arid soils, usually on gypsum-rich or loamy substrates; 01,000 m.
Distribution: ALG MOR SPA. Ibero-North African endemic (SE Iberian Peninsula,
NE Morocco, NW Algeria; Fig. 4).
Phenology: IIIVI.
Reseda phyteuma L., Sp. Pl. 1: 449. 1753.
Ind. loc.:Habitat in Gallia, Italia.
Lectotype:8Phyteuma(LINN photo!) (Designated by Jafri in Jafri and El-Gadi,
Fl. Libya 34: 14. 1977).
Icon.: Jacquin, Fl. Austriac. 2, tab. 132. 1774; Fiori and Paoletti, Iconogr. Fl. Ital. 1:
150, n. 1305. 1899; Reichenbach, Icon. Fl. Germ. Helv. 2, lám. XCIX, n. 4445.
1838; Abdallah and de Wit in Meded. Landbouwhogesch. Wageningen 78(14),
Atlas, fig. 70. 1978.
416 S. Martín-Bravo, P. Jiménez-Mejías
=R. confusa Pomel, Nouv. Mat. Fl. Atlant. 224. 1875.
Ind. loc.:Terrains de Sebkha: Saint-Louis près dOran, Télamine
Lectotype (designated here): Oran, St. Louis, Pomel s.n.(MPU 005087
photo!). Isolectotypes: MPU, P.
R. phyteuma var. confusa (Pomel) Batt. in Batt. & Trab., Fl. Algérie (Dicot.)
84. 1888.
=R. phyteuma var. rupestris Lange in Willk. & Lange, Prodr. Fl. Hispan. 3: 895.
1880.
Ind. loc.: in rupibus montis Sierra de Alfacar fl. Granat.
Lectotype (designated here): Sierra de Alfacar prope Granatam, in rupibus, 5
Aug 1876, Hackel 54(C, photo!).
R. phyteuma subsp. rupestris (Lange) Aránega & Pajarón in Anales Jard. Bot.
Madrid 47: 60. 1989.
=R. aragonensis Loscos & J. Pardo in Willk. (ed.), Ser. Inconf. Pl. Aragon: 14. 1863.
Ind. loc.: Provenit frequens in Aragonia centrali atque in ditione el
Maestrazgo
Lectotype:Aragonia austr. circa Castelseras, JunSep 1863, F. Loscos 72
(UPS, photo!) (Designated by Abdallah and de Wit in Meded. Landbouwhoo-
gesch. Wageningen 78(14): 306. 1978). Isolectotypes: UPS, W.
R. phyteuma var. aragonensis (Loscos & J. Pardo) Nyman, Consp. Fl. Eur. 69.
1878
R. phyteuma f. aragonensis (Loscos & J. Pardo) Rouy & Foucaud, Fl. France
2: 244. 1895
R. phyteuma subsp. aragonensis (Loscos & J. Pardo) Bonnier, Fl. III France
2: 20. 1913
R. phyteuma subsp. aragonensis (Loscos & J. Pardo) Rivas Mart., Itinera
Geobot. 15: 707. 2002, isonym.
R. phyteuma f. aragonensis (Loscos & J. Pardo) Sennen in Bol. Soc.
Aragonesa Ci. Nat. 10 (710): 134. 1911, isonym.
= R. phyteuma var. integrifolia Texidor in Revista Progr. Ci. Exact. 18(9): 597. 1869.
Ind. loc.: es c. en el partido de Olot, no r. en toda Cataluña, Madrid y
Aranjuez
Lectotype: probably in BCN (cf. Guàrdia and Allès 2008)
R. phyteuma f. integrifolia (Texidor) Maire, Fl. Afrique N. 14: 191. 1976.
R. phyteuma f. stenopetala Maire, Fl. Afrique N. 14: 191. 1976, nom. inval.
R. phyteuma subsp. eu-phyteuma Maire in Jahand. & Maire, Cat. Pl. Maroc 316.
1932, nom. inval.
Morphological description: Annual to perennial herb, 1050 cm tall. Stems
decumbent or ascending, rarely erect, branching from the base, glabrous, papillose or
hispidulous. Basal leaves entire or subentire, sometimes rosette-like; stem leaves
alternate, entire to 3(5)-lobed, terminal lobe rarely 3-lobed in turn; glabrous,
papillose or hispidulous. Inflorescence spiciform, usually more or less lax at the top,
Taxonomic re-evaluation of Reseda collina 417
with blunt apex. Pedicels 310 mm in flower, up to 20 mm in fruit. Flowers
bisexual, 6(7)-merous. Calyx dialysepalous, sepals 36× 0.52 mm in flower,
persistent, usually markedly accrescent, reflexed in fruit, (4)612×(0.5)14 mm,
oblong or spathulate. Corolla dialypetalous, heteromorphic, white, sometimes
yellowish when dry; superior petals 35(6) mm, unguiculate, trisect, lateral lobes
palmatisect, each deeply divided in (4)58 usually linear laciniae, central lobe
linear, shorter than lateral lobes; lateral and anterior petals reduced. Stamens 1520,
filaments deciduous, generally widened in the distal part, connate at the base to form
an asymmetrical nectariferous disc. Capsules 3-carpelled, 920 × 410 mm, estipi-
tate, pendulous when ripe, obovoid-cylindric to subglobose, glabrous, sometimes
papillose in the ribs, capsule teeth 0.51(1.5) mm long. Seeds 22.5 mm, reniform,
brownish when ripe, dull, testa undulate-rugose, with more or less sharp ridges,
apical carunculoid tissue reduced or absent.
Ecology: Wastelands, fields, slopes, occasionally rupicolous, on basic soils;
01,900 m.
Distribution: ALB ALG AUT CZE FRA GRC HUN ITA MOR POL POR ROM?
SPA SWI TUN? YUG. Western and central Mediterranean region, Central Europe.
Chromosome number:n=6; 2n=12, 24.
Phenology:IIIX(XII).
Acknowledgements The authors thank M. Míguez and F.J. Fernández for technical support; the curators of
ALME, AMD, BM, C, DBN, E, GDA, GH, HUAL, IBF, K, MA, MGC, O, RNG, S, SEV, UPOS and UPS
herbariafor the loanof specimens, granting permission for DNA extractions and providingphotographs of type
material. We are also grateful to M. Escudero, M. Luceño and to two anonymous reviewers for their critical
comments on the manuscript. The Royal Botanic Garden of Madrid kindly provided facilities for using the
electron microscope. This research was supported by the Spanish Ministry of Science and Technology and the
Andalusian Government through the projects CGL2005-06017-C02-02/BOS and P06-RMM-4128, respec-
tively. The experiments performed for this research comply with the current laws of Spain.
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Published online: 18 November 2009
Appendix 1
List of the studied herbarium specimens of Reseda collina
Algeria: Oran, collines pierreuses de l´Ouest, 27 Apr 1842, Durieu (K, Fig. 5);
Oran, 9 Apr 1856, E. Cosson (K, S); Oran, Jun 1844, Durieu (MPU, photo!); Oran, à
Gambetta, lieux incultes, 13 May 1909, A. Faure (AMD, BM); Circà Oran, 1849,
Boissier & Reuter (UPS, O); Oran, in arenosis, Apr 1851, Munby (BM, DBN); Oran,
in collibus arenosis maritimis, Apr 1856, Munby (BM); Oran, rochers des falaises,
10 Oct 1859, E. Bourgeau (GH); Oran, champs, 16 Apr 1881, O. Debeaux (BM);
Oran, in arvis siccis, 15 Apr 1884, O. Debeaux (MA); Oran, vallou de voiseux, 23
Mar 1934, R. Maire & E. Wilczek (MA); Coteaux incultes avoisinant la Batterie
espagnole, près d´ Oran, 23 Apr 1852, B. Balansa (MPU, photo!); Rochers des
falaises à Oran, 10 Apr 1856, E. Bourgeau (MPU, photo!).
Morocco: Marruecos, Beni-Snassen, subiendo hacia el puerto de Taforalt, bosque de
Tetraclinis articulata sobre calizas, taludes junto a la carretera, 34º50´ N, 2º24´ W,
495 m, 20 Apr 2004, S. Martín Bravo (UPOS, designated epitype, Fig. 7,isoepitypesG,
K, MA, RNG); Bou Idoudane El Hamman, hacia Ahrour, tomillares basófilos, 34º56´
N, 3º25´ W, 400 m, 14 Apr 1994, A.M. Romo et al. (BM, RNG); Ctra. a Al-Hoceima,
puerto de Kassita, taludes rocosos junto a la carretera, sobre margas áridas, 34º54´ N,
3º48´ W, 590 m, 22 Apr 2004, S. Martín Bravo & P. Vargas (UPOS); Beni Snassen,
Taforalt, pinar de Pinus halepensis, taludes rocosos al borde de la ctra., sobre areniscas,
34º47´ N, 2º24´ W, 870 m, 21 Apr 2004, S. Martín Bravo (UPOS); Imzourene, 6 km W
of Al Hoceima on road to Torres de Alcala, N to NE facing hillsides, dry clay/loam on
limestone, 35º13´ N, 3º59´ W, 28 Feb 1994, S.L. Jury et al. (RNG); Sidi Slimane,
wilaya d´Oujda, Monts des Beni-Snassen, Taforalt, petit affleurement calcaire au ras du
sol dans une friche, 34º47´ N, 2º24´ W, 860 m, 10 Mar 1995, J. Lambinon & G. Van
den Sande (RNG); South side of mounts SW of Berkane, near Taforalt, eroded bank,
800 m, 19 Apr 1971, Davis (E,RNG,mixedwithR. phyteuma); Targuist, ca. 40 km
WSW of Al Hoceima, near Peñon Vélez de la Gomera, E-facing steep limestone slope
near coast, 35º10´ N, 4º18´ W, 90 m, 28 Feb 1994, S.L. Jury et al. (RNG); Muley-
Rechid, talus de la route, 20 Jun 1931, Sennen & Mauricio (BM, MA); Oujda, between
Berkane and Taforalt, mountains of Beni-Snassen, Tetraclinis articulata forest and scrub
limestones, 34º50´ N, 2º20´W, 600 m, 30 May 1993, B. Valdés et al. (RNG, SEV);
Oued Selouane, hacia Irgana, pastos secos sobre terrenos calizos, 35º20´ N, 2º56´ W,
120 m, 6 Apr 1994, A.M. Romo et al. (SEV); Massif des Beni-Snassen, environs de
Taforalt, rochers et lieux rocailleux, 900 m, 28 May 1931, A. Faure (BM); Ulad-Settut,
à Muley-Rechid, coteaux, 22 May 1932, Sennen & Mauricio (MA); Muley-Rechid,
Ulad-Settut, talus, 12 Jun 1934, Sennen & Mauricio (MA); Nador, Kebdana, Azrou
Aujdiou, matorrales del Rosmarino-Ericion, 790 m, 7 Apr 1994, A.M. Romo et al.
420 S. Martín-Bravo, P. Jiménez-Mejías
(RNG); Melilla, Rostro Gordo, 10 Apr 1912, Caballero (MA); Melilla, Mazuza, 17 Apr
1932, Herbarium Pau (MA); Melilla, Mazuza, 26 Feb 1933, Pardo & Martí (MA).
Spain: Almería province: Loma de las Cabras, 2 May 1989, J.M. Sánchez Prados
(MGC); Boca de Berciel, 2 May 1989, J.M. Sánchez Prados (MGC); Almería, camino
de Dalías, 20 Apr 1989, J.M. Sánchez Prados (MGC); Almería, Berja, Sierra de Gádor,
1,000 m, 19 Apr 1951, Hno. Jerónimo (ALME); Barranco del Caballar, 26 Apr 1951,
Hno. Jerónimo (ALME); Almería, Sierra de Gádor, Rambla Molinos, 30SWF2676,
400 m, 25 May 1996, E. Giménez & F. Gómez-Mercado (HUAL);SierradeGádor,
Énix, 30SWF3581, 730 m, 11 Apr 1996, E. Giménez & F. Gómez-Mercado (HUAL);
Sierra de Gádor, Rambla del Boquerón, 30SWF0977, 400 m, 1 Apr 1996, E. Giménez
& F. Gómez-Mercado (HUAL); Sierra de Gádor, cara sur, 950 m, 20 Jun 1991, J.
Cabello et al. (HUAL); Sierrecilla, 600 m, 30 Feb 1996, E. Giménez & F. Gómez-
Mercado (HUAL); Barranco del Caballón, Apr 1960, Muñoz Medina (GDA); Sierra de
Gádor, May 1974, Hno. Jerónimo (GDA); Campos de Dalías, Matagorda, bordes de
invernadero, 30SWF1464, 31 Mar 1983, A. Belén Robles & A. Ortega (GDA); entre
Pechina y Los Baños de Sierra Alhamilla, tomillar semiárido sobre enclave de margas
yesíferas y yesos del Mioceno superior, 30SWF517882, 200220 m, 18 May 1983, R.
Lázaro Suau (ALME); entre Berja y Ugíjar, ruderal, 600 m, 15 Feb 1979, Molero Mesa
& Pérez Raya (MA); de Alhama de Almería a Enix, pasado el Marchal de Antón López,
15 May 1986, Fernández et al. (MA); Barranco del Caballar, roquedos calizos, laderas
margosas, 110 m, 6 Mar 1982, A. Barra et al. (MA); Dalías, A.M. Fernández & J.J.
González (MA); El Ejido de Dalías, Jan 1970, J. Fernández Casas (MA); Sierra de
Gádor, Berja, 21 Jun 1934, Hno. Jerónimo (MA); Almería, 2,000 ft, R. Clemente (MA);
Roquetas, 20 Mar 1921, Gros (MA); La Roqueta, 26 Mar 1883, N.H. Nilsson (UPS).
Appendix 2
List of the herbarium specimens used for Fig. 3
Reseda phyteuma. Photo a, Spain, Almería, prope pagum Viator, in arenosis, sol.
schistoso, 60100 m, May 1890, Porta & Rigo (IBF). Photo c, Spain, Málaga,
Ronda, alrededores del pueblo, pinar de Pinus pinea, suelos arcillosos profundos,
36º45N, 5º10W, 670 m, 8 Apr 2006, S. Martín Bravo (UPOS). Photo e, Spain,
Madrid, Valdemoro, cunetas de la N-IV, 40º08N, 3º39W, 622 m, 19 Apr 2004,
P. Vargas (UPOS). Photo g, Spain, Sevilla, entre Casariche y Badolatosa, taludes de
la carreteras sobre suelos margosos, 37º18N, 4º44W, 331 m, 28 May 2004,
S. Martín Bravo (UPOS).
R. collina. Photo b, Morocco, ctra. a Al-Hoceima, puerto de Kassita, taludes rocosos
junto a la carretera, sobre margas áridas, 34º54N, 3º48W, 590 m, 22 Apr 2004,
S. Martín Bravo & P. Vargas (UPOS). Photos d, f, Morocco, Beni Snassen, Taforalt,
pinar de Pinus halepensis, taludes rocosos al borde de la ctra., sobre areniscas,
34º4748′′ N, 2º2400′′ W, 870 m, 21 Apr 2004, S. Martín Bravo (UPOS). Photo h,
Spain, Almería, camino de Dalías, 20 Apr 1989, J.M. Sánchez Prados (MGC).
Taxonomic re-evaluation of Reseda collina 421
... Phyteuma Lange (1880: 894) (Abdallah & de Wit 1978). Seven taxa out of the 18 that form sect. Phyteuma (Martín-Bravo & Jiménez-Mejías 2009, 2013 are currently present in Turkey: R. minoica Martín-Bravo & Jiménez-Mejías (2013: 57), R. orientalis Boissier (1867: 427), R. armena Boissier (1843: 5), R. coodei Huber-Morath (1967: 312), R. balansae, R. malatyana Yıldırım & Şenol (2014Yıldırım & Şenol ( : 1014, and R. anatolica (Abdallah & de Wit 1978: 336) Snogerup & Snogerup (2002: 300), of which the five latter are Turkish endemics. Reseda balansae has been distinguished from other taxa in sect. ...
... The latter species is distributed in Central Europe and the Western and Central Mediterranean region, and is probably absent from Turkey. This illustrates the taxonomic complexity of section Phyteuma in the Mediterranean (Martín-Bravo & Jiménez-Mejías 2009, 2013. ...
... indumentum, calyx accrescence, number of stamens, shape of petals and staminal filaments, capsule and seed size; cf. Coode 1965;Abdallah & de Wit 1978;Martín-Bravo & Jiménez-Mejías 2009, 2013. For the evaluation of the conservation status of R. balansae at global level we followed criteria, categories and guidelines from IUCN (2012IUCN ( , 2017. ...
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Reseda balansae Müll. Arg. (Resedaceae) is a very restricted and endangered Turkish endemic species known only from Mersin province (South Turkey), in a hotspot of biodiversity. It was first collected in 1855, and subsequently described in 1857. Afterwards, it was only collected once again in 1896. In this study we report the rediscovery of these two populations of R. balansae in Turkey more than 120 years after its last known collection. We provide a detailed revised description of this poorly known species and comments about its taxonomy (including designation of a lectotype), distribution and ecology. We also perform the first conservation assessment of the species at a global scale under IUCN categories and criteria, resulting in the proposal of the critically endangered category for the species, which apparently persists in one single population, since the type population has recently been destroyed.
... Reseda balansae is a poorly known species that was described in the first comprehensive taxonomic monograph of Resedaceae published in 1857. It is included in section Phyteuma, a monophyletic group with a complex taxonomy characterized by problems regarding species circumscription, mainly due to limited morphological differentiation and hybridization processes (Martín-Bravo et al. 2007, Martín-Bravo and Jiménez-Mejías 2009, 2013. ...
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This is a highly restricted endemic species only known from two populations in the eastern Taurus range in southern Turkey. Its area of occupancy (AOO) is only 8 km2. One of the two only known populations has been recently largely destroyed by roadworks. There are probably fewer than 30 mature individuals remaining. Its main current threat is the destruction of its habitat due to infrastructure building. The species is listed as Critically Endangered (CR D).
... The evaluation of molecular and morphological data has uncovered errors in traditional taxonomy, and it has reconciled conflicts between traditional, morphologybased taxonomy and molecular phylogenies (e.g. Martín-Bravo & Jiménez-Mejías, 2009;Jiménez-Mejías, Martín-Bravo & Luceño, 2012;Vigalondo et al., 2016). ...
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... The evaluation of molecular and morphological data has uncovered errors in traditional taxonomy, and it has reconciled conflicts between traditional, morphologybased taxonomy and molecular phylogenies (e.g. Martín-Bravo & Jiménez-Mejías, 2009;Jiménez-Mejías, Martín-Bravo & Luceño, 2012;Vigalondo et al., 2016). ...
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Species delimitation in Carex section Rhynchocystis has remained relatively constant through its taxonomic history. The section is currently composed of five species distributed in the Western Palaearctic (C. microcarpa and C. pendula) and subSaharan Africa (C. bequaertii, C. mossii and C. penduliformis). Recent phylogenetic studies revealed that the monophyly of C. bequaertii and C. mossii was not well supported and that C. pendula comprises two divergent sister lineages. To evaluate the taxonomic significance of these unexpected results, we performed a rigorous statistical procedure based on morphometrics. We found morphological support for our molecular background, uncovering (1) characters that reflect the evolution of the group and were overlooked by traditional taxonomy and (2) the overlapping of some previously considered diagnostic characters. Our results suggest five species, but only C. microcarpa and C. penduliformis were supported in their traditional concepts. The two lineages of C. pendula corresponded to two morphologically distinct, biogeographically congruent groups: C. pendula s.s. in the western part of the range and C. agastachys in the eastern part. In congruence with the molecular results, C. bequaertii and C. mossii were weakly morphologically differentiated and thus better treated as subspecies of a single species. We propose a revised taxonomic treatment for the group.
... Combined approaches of both morphological and molecular data and statistical analyses of those data are currently the most frequent practice for species delimitation and new species descriptions in botany as well as in zoology (e.g. [50][51][52][53][54][55][56]). The combination of morphological and molecular data has been previously shown to be a powerful tool to resolve the taxonomy in Carex (e.g. ...
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... Its taxonomy is characterized by certain problems regarding species circumscription and the presence of hybridization processes (Martín-Bravo and Jiménez-Mejías 2009). In particular, R. arabica is probably one of the putative ancestors of the hybrid species R. odorata L., as inferred from the analysis of nuclear and plastid DNA sequences (Martín-Bravo et al. 2007;Martín-Bravo and Jiménez-Mejías 2009). Nonetheless, R. arabica is morphologically well-characterized and can be readily distinguished from closely related species by its persistent staminal filaments and the sessile or subsessile capsules (Martín-Bravo 2009). ...
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... To avoid further taxonomical problems, and to ensure the precise application of the species name, we propose the designation of an epitype (ICN McNeill et al. 2012, art. 9.7; see also Martín-Bravo & Jiménez-Mejías (2009) for further discussion in epitipification). We select an specimen with ripe fruits (Bourgeau 1353), which was cited by Cosson ("…in convalle Tamadaya (…) Bourgeau pl. ...
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— We studied sequence variation in 16S rDNA in 204 individuals from 37 populations of the land snail Candidula unifasciata (Poiret 1801) across the core species range in France, Switzerland, and Germany. Phylogeographic, nested clade, and coalescence analyses were used to elucidate the species evolutionary history. The study revealed the presence of two major evolutionary lineages that evolved in separate refuges in southeast France as result of previous fragmentation during the Pleistocene. Applying a recent extension of the nested clade analysis (Templeton 2001), we inferred that range expansions along river valleys in independent corridors to the north led eventually to a secondary contact zone of the major clades around the Geneva Basin. There is evidence supporting the idea that the formation of the secondary contact zone and the colonization of Germany might be postglacial events. The phylogeographic history inferred for C. unifasciata differs from general biogeographic patterns of postglacial colonization previously identified for other taxa, and it might represent a common model for species with restricted dispersal.