Research Article |
Corresponding author: Javier Martínez-Fort ( thy2mus@yahoo.es ) Academic editor: Anthony R. Magee
© 2019 Javier Martínez-Fort, Maela León, Maria P. Donat-Torres.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Martínez-Fort J, León M, Donat-Torres MP (2019) A new subspecies of Peucedanum officinale L. subsp. album (Apiaceae) from the eastern part of the Iberian Peninsula. PhytoKeys 131: 37-55. https://doi.org/10.3897/phytokeys.131.32173
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We describe Peucedanum officinale subsp. album Martínez-Fort & Donat-Torres subsp. nov., in which we grouped the thermomediterranean populations scattered along the eastern part of the Iberian Peninsula. The characters that differentiate this new subspecies from other infraspecific taxa in Peucedanum officinale are its canaliculated leaflet, the inflorescences much branched and lack of dominant terminal umbels, the umbels are few rayed, sometimes sessile and lateral, the petals are white and the fruit pedicels short, the same or shorter in length than the fruit. We provide here a full description of the new subspecies based on herbarium specimens and field measurements, as well as providing dichotomous keys to the subspecies within P. officinale. In addition, we provide a comparison of the ITS sequences of nrDNA with the most closely related taxons.
Umbelliferae, Spain, taxonomy, subsp. nov., thermomediterranean, nrITS
Peucedanum L. 1753 is one of the most complex genera in the Apiaceae family. Based on the morphological characteristics of the fruit, according to the traditional classification systems, it is characterised by a strong dorsal compression and winged side ribs as in
On the other hand, phylogenetic studies, based on the ITS rDNA sequences, define “Peucedanum sensu stricto clade including taxa that are very similar with respect to their ITS sequences and they are very related in habit, sharing not only fruit characters but also vegetative features, like linear-filiform leaf lobes” and it has been regrouped in the Tribe Selineae Spreng (
The morphological differences between some populations from the central and eastern coastal parts of the Iberian Peninsula have long since been evidenced with the description and citation of several P. officinale subspecies (
The first distinction from the Iberian populations in P. officinale was made with the description of P. stenocarpum Boiss. & Reut. ex Boiss., 1844 (
Subsequently,
Another name proposed was P. stenocarpum var. catalaunicum Pau in
Afterwards, P. stenocarpum was included as a subspecies of P. officinale (viz. P. officinale subsp. stenocarpum Font Quer 1950) and included all the populations in the eastern half of the Peninsula. Its morphological characteristics were summarised as: extended (up to 15 cm) and very narrow (from 0.3 to 2 mm) leaf segments, with between 10 and 64 umbel rays and a ratio of fructiferous pedicel length to fruit length from 0.5 to 1.5 (Font Quer 1954). Font Quer measured some of the populations that we have included in the present study and which have always been assigned the extreme value in the description of his study. Along with the subspecies officinale, they are the only two taxa in the group recognised in the Peninsula at this time. Peucedanum paniculatum and P. longifolium Waldst. & Kit. 1812 were reduced to subspecies of Peucedanum officinale due to their ternate leaves, which were divided into linear or linear-lanceolate segments by
The last subspecies in the group, described in the Iberian Peninsula, Peucedanum officinale subsp. brachyradium García-Martín and Silvestre 1991 specifically in the province of Málaga (Spain), was indicated as an edapho-endemism on peridotites. That description derives from a population with only two individuals and is morphologically distinguished from the subspecies type and from P. stenocarpum by the characteristics of its inflorescence and by the dimensions of its fruit and pedicels.
The latest review undertaken of the genus for the Iberian Peninsula (
While conducting fieldwork during the previous years, we located the population of an umbelliferous species in eastern spurs of the Serra Grossa mountain range, which lies in the southern part of the Valencian province (eastern part of the Iberian Peninsula). This species is always found at the bottom and top of rocky areas, on the edges of paths, always in cracks in rocks and on rocky soil. It is characterised by possessing ternatisect leaves, with linear and canaliculated leaf divisions; inflorescences with sessile lateral umbels and with few rays; white-petalled flowers; elliptic fruits with a strong dorsal compression and prominent dorsal ribs and winged marginal ribs, borne on short pedicels. As a result, we assigned it to the genus Peucedanum sensu stricto.
To further identify this taxon and compare all of the subspecies of P. officinale throughout its area of distribution, we reviewed the herbarium specimens of the genus deposited in the VAL and MA herbaria. We studied all the bibliography about the P. officinale subspecies cited and described in the Iberian Peninsula and the monograph on the genus by
We noted that the population under investigation here had been identified to date as P. stenocarpum, widely considered as a synonym of P. officinale subsp. officinale, despite the fact that it has a series of morphological characters that are not accommodated within any of the subspecies of P. officinale (Figures
Population | Mountain range | Province/Towns | Local name | YWGS84 coordinates |
---|---|---|---|---|
Po1 | Extreme north, Serra Grosa | V: Carcaixent, Alzira | Hort de Soriano-Font de la Parra, Molló de Miramar | 39°3.88'N, 0°23.694'W |
Po2 | Spurs, Sierra de Corbera | V: Llaurí, Favara, Tavernes de la Valldigna | Font de la Granata, Pic Masalari | 39°5.43'N, 0°17.238'W |
Po3 | Spurs, Sierra de Corbera | V: Alzira, Corbera, Llaurí | Serra de la Murta, Creu del Cardenal | 39°7.61'N, 0°20.856'W |
Po4 | Sierra Picarcho | V: Tuejar, Cu:Talayuelas | Umbría del Picarcho | 39°49.83'N, 1°13.002'W |
Po5 | Serra Mondúver | V: Gandia, Xeresa | Cima Mondúver | 39°0.45'N, 0°15.834'W |
Po6 | Desert de les Palmes | Cs: Benicasim | La Comba | 40°3.94'N, 0°3.144'E |
Po7 | South, Sierra de Guadarrama | M: El Escorial | El Escorial | 40°35.09'N, 4°7.62'W |
Character | Units | Code | |||
---|---|---|---|---|---|
Stem | Size | Plant height | m | T1 | |
Ramification | First ramification height | m | T2 | ||
T1/T2 ratio | T3 | ||||
Diameters | Diameter at the base | cm | T4 | ||
Diameter at the first ramification | cm | T5 | |||
T4/T5 ratio | T6 | ||||
Leaves | Basal rosette | Blade | Leaf length, including petiole | mm | H1 |
Number of leaf divisions (ternate) | H2 | ||||
Leaf width at the base of the last division | mm | H3 | |||
Leaf width at the apex of the last division | mm | H4 | |||
Last leaf division length | mm | H5 | |||
H5/H3 ratio | H6 | ||||
H4/H5 ratio | H7 | ||||
Angle between contiguous leaf divisions of the last ternate | degrees | H8 | |||
Angle between lateral leaf divisions of the last ternate | degrees | H9 | |||
Petiole | Length | mm | H10 | ||
Diameter at its basis | mm | H11 | |||
Diameter before first division (ternate) | mm | H12 | |||
First on stem from the basal rosette | Blade | Number of leaf divisions (ternate) | H13 | ||
Last leaf division length | mm | H14 | |||
Petiole | Length | mm | H15 | ||
Diameter of the petiole at its basis | mm | H16 | |||
Diameter before the first division (ternate) | mm | H17 | |||
Last on stem | Blade | Number of leaf divisions (ternate) | H18 | ||
Petiole | Length | mm | H19 | ||
Diameter before the first division (ternate) | mm | H20 | |||
Diameter at its basis | mm | H21 | |||
Sheath | Sheath length | mm | H22 | ||
Inflorescence | Umbel | Bract length | mm | I1 | |
Bract width | mm | I2 | |||
Number of bracts | I3 | ||||
I1/I2 ratio | I4 | ||||
Number of rays | I5 | ||||
Length of internal rays | mm | I6 | |||
Length of external rays | mm | I7 | |||
Umbellule | Number of raylets | I8 | |||
Length of internal rays | mm | I9 | |||
Length of external rays | mm | I10 | |||
Number of bracteoles | I11 | ||||
Fruit | Size | Length | mm | F1 | |
Width | mm | F2 | |||
Wing width of the marginal ribs | mm | F3 | |||
Shape | F3/F2 ratio | F4 | |||
F1/F2 ratio | F5 | ||||
Raylet | Fructiferous raylets length | mm | F6 | ||
F1/F6 ratio | F7 | ||||
F6/F1 ratio | F8 | ||||
F2/F6 ratio | F9 |
Measurements were taken in the field and in the laboratory and always carried out with fresh plants. We used a CD-20DCX digital vernier caliper and a metal tape measure and saved measurements on spreadsheets and databases. We created the figure of P. officinale subsp. album using the holotype VAL 223161 to scan, as well as from the photos taken in the field with a Canon EOS 550D camera of inflorescences and fruits. Other photos were taken with a Leica stereomicroscope MZ 9.5 and a Leica DFC 320 digital camera of the cross-sectional cuts of fruits and leaf details. We obtained the flowering and fructification data while conducting fieldwork and according to the date of the specimens and phenologies from the consulted herbarium sheets. All the other figures are photos of herbarium sheets and pictures of scanned herbarium sheets. We compared and statistically analysed the taken measurements with the Statgraphics Centurion XVI software. For morphological characters, we followed the standardised terminology of
Taking measurements was limited by the small proportion of individuals which were flowering or under fructification in all the populations; indeed, flowering and fructification did not even exceed 4% of the individuals in the largest populations. In the specimens that presented no flowering, measurements were taken from the basal rosette leaves. In all, we took measurements from 31 specimens, of which 19 belonged to populations of the newly proposed taxon and 12 to the type subspecies. In all, 1,316 measurements and ratios were taken and calculated.
The examined herbarium specimens’ material at VAL and MA are listed below.
Peucedanum longifolium Waldst. & Kit.
YUGOSLAVIA. Dalmatia: Montes Biokovo, in rupibus calcareis sub cacumine montis Sv. Jure supra opp. Makarska, 1600 m alt. 29-VII-1979, F.Cernoch (MA 357267); ídem (MA 310966)
Peucedanum officinale L.
BULGARIA. Regio Sophiensis: distr. urb. Sophia, inter fruticeta supra, 20-IX-1979. N. Andreev, Z. Cerneva & P. Gerginov (MA 309655)
GERMANY. Maingebaeit: Würzburg, woodland between Gerbrunn and Kottendorf, 00-VII-1881, G. Evers (MA 713330)
SPAIN. Aragón:(MA 88547); Palau (MA 88581). Alava: Labastida, Salinillas de Buradón, 12-X-1990, P.M. Uribe-Echebarria & P. Urrutia (MA 523402); Labastida, Salinillas de Buradón, cerro calizo con matorral mediterráneo, 30TWN1320, 5-IX-1998, P.M. Uribe-Echebarría (VAL 144623); Labastida, Salinillas de Buradón, 02-X-1997, M. L. Gil Zúñiga (MA 616760); ídem (VAL 106191) Albacete: Molinicos, valle del río Mundo, entre Mesones y la fuente de la Plata, 30SWH587602, 900 m alt., 18-VIII-2002, M.J. Tohá & V.J. Arán (MA 703748); Molinicos, Valle del río Mundo, entre Mesones y la Fte. de la Plata, Laderas calcáreas con Cinar, 30SWH587602, 18-VIII-2002, V.J. Arán & M.J. Tohá (VAL 144146). Barcelona: Berga a Labaello, 16-VII-1911, Fre. Sennen (MA 88550); Al lado de la Ermita de S. Jerónimo, IX-1914, Caballero (MA 88584); Montserrat, IX-1905; Marcet (MA 88545). Castellon: Benicàssim (La Plana Alta), La Comba, 31TBE43, 07-IV-1990, J. Tirado & C. Villaescusa (VAL 26141); La Pobla Tornesa (La Plana Alta), Bartolo cresta, 31TBE44, 17-IX-1989, J. Tirado & C. Villaescusa (VAL 26140). Cuenca: Huete, hacia Garcinarro, valle del arroyo de Valquemado, 13-IX-2003, V. J. Arán & M. J. Tohá (MA 711491); Huete, hacia Garcinarro, valle de arroyo de Valquemado, 10-X-2004, V.J. Arán (MA 751028); Huete, hacia Garcinarro, valle del arroyo de Valquemado, al pie de cerros yesosos, 30TWK2449, 6-VIII-2005, V. J. Arán (VAL 179661); Huete, hacia Garcinarro, valle del arroyo de Valquemado, al pie de cerros yesosos, 30TWK245490, 13-IX-2003, V. J. Arán & M. J. Tohá (VAL 149187); Talayuelas, VII-1979, G. Mateo (VAL 110252); Talayuelas, X-1980, G. Mateo (VAL 110251). Gerona: Maçanes, 80 m alt., Font Quer, 12-X-1948 (MA 152333); ídem (MA 382969); Pyrénées à Gombreny, coteaux calcaires, 900 m alt., VIII-1913 (MA 88562), ídem (MA 88561), ídem (MA 88560). Huesca: Ayerbe, 600 m alt., 31-VIII-1973, A. Segura Zubizarreta (MA 359384); Arro, 26-IX-1979, P. & G. Montserrat (MA 357236); ídem (MA 311455). Lérida: La Granadella (Garrigues), hacia El Solerás, pr. riera de Vall de les Olives, junto a la carretera, 31TCF0486, 365 m alt., 08-IX-2008, V.J. Arán & M.J. Tohá (MA809441); ídem (VAL 196084). Logroño: Briones, Monte Lara, 1925, Hno. H. Elias (MA 88558). Madrid: Chozas, Cutanda, IX, (MA 88552); Entre Villalba y las Zorreras, en la Sierra del Guadarrama, 08-IX-1947, Rivas Goday & C. Pérez (MA 152463); ídem (MA 204879); ídem (MA 382966); Guadarrama, IX-1841, Reuter (MA 88577); Escorial, Graells (MA 720356); Guadarrama, J. Isern. (MA 720260). Pontevedra: Santa Maria de Oya, 10 m alt., 20-VIII-1983, S. Silvestre (MA 316210). Salamanca: Saucelle, 16-VIII-1978. F. Amich (MA 309660). Tarragona: Sant Carles de la Ràpita, Serra de Montsià, 450 m alt., 01-IX-1999, V. J. Arán & J. Masip (MA 631766); Sant Carles de la Ràpita, Serra de Montsià, Font de Burgà, hacia el SE, laderas soleadas sobre la fuente, entre el matorral calcícola. 31TBF9301, 1-VII-1999, Arán & Masip (VAL 41688). Teruel: Cantavieja, hacia Mirambel, 30TYK29, 4-IX-1993, Fabregat & López Udias (VAL 81745); Olba, IX-1894 (MA 88548); Castellote, alrededores de las Cuevas de Cañart, 11-IX-1991, C. Fabregat (MA 502852); ídem (VAL 75992); Olba, Caserío de la Berdeja, Ribazos. 30TXK9844, 25-IX-2004. S. López Udias & C. Fabregat (VAL 204090); San Agustín, valle del Mijares, hacia Rubielos de Mora, 30TXK9445, 4-IX-2004, G. Mateo (VAL 151501); Villarluengo, barranco de los Degollados, márgenes de la carretera, 30TYL00, 11-IX-1993, Mercadal (VAL 81675). Valencia: Ayora, La Hunde, 30SXJ52, 00-VIII-1981, J. B. Peris (VAL 17838); Bicorp, Cuesta de la Caruma, 25-VIII-1915, C. Vicioso (MA 88549); Corbera de Alcira, 30SYJ23, J. Borja. (VAL 154996); Serra de Corbera, 30SYJ23, 00-X-1944, J. Borja (VAL 117840), Favara: Serra de Corbera, 30SYJ33, 5-IX-1986, G. Mateo & al. (VAL 117826); Sinarcas, 30SXK50, 6-VII-1992, García Navarro (VAL 105040); Sinarcas, Peña del Rayo, 30SXK50, 12-IX-1989, García Navarro, (VAL 102949); Tuéjar (Serrans), Altos del Picarcho, rodenos, 30SXK5311, 12-IX-2004, C. Torres Gómez, G. Mateo & J. Fabado (VAL 217597); Tuéjar a Talayuelas, umbría del Picarcho, 30SXK51, VIII-1980, G. Mateo (VAL 110250); Tuéjar (Serrans) Altos del Picarcho, rodenos, 30SXK5310, 24-IX-2005, C. Torres Gómez (VAL 216649). Zamora: Muelas del Pan, 13-VIII-1978, E. Rico (MA 309659); Río Esña, Riberos del pantano de Ricobayo, 00-VII-1972, Rivas Goday & Ladero (VAL 117836). Zaragoza: El Frasno, 30TXL26208190, 2-IX-1995, A. Martínez (VAL 216135); Moncayo, 3-VIII-2000, Vicioso (VAL 180380); Torrero, 00-VI-1947, P. Capell S.J. (VAL 180379).
FRANCE. Pyrénées-Orientales. Conflent. En allant de Ille-sur-Têt à Montalba a 2 km env. de Montalba, 02-X-1970. J. Vivant (MA 357223); La Garde Freinet, an der D. 48 nörlich La Trémoulêde,, 4-X-1963 (MA 626021); Languedoc-Roussillon, Aude, sur le versant nord du col d´Extrème entre Villeneuve-des-Corbières et Tuchan, 15-IX-2004. Philippe Rabaute. (MA 802614); Cher: IX-1890, A. Le Grand (MA 88575)
HUNGARY. Bács Bodrog, Bezdan, 10-IX-1909, J. Prodan (MA 88576)
Peucedanum paniculatum Loisel.
FRANCE. Haute Corse: Castagniccia, Col di Bigorno, 10-IX-1996, J. Lambinon (MA 628116); Ghisoni, 17-VIII-1899, R. Rotges (MA 88546); Ghisoni, Maquis peu touffu, 10-VIII-1929, Dr. C. Gabrel (MA 425140); Massif du Tenda, Col di Bigorno, mun. Bigorno, 07-VIII-1996, L. Serra & A. Bort (MA 623316).
We made a genetic comparison of the ITS regions of ribosomal DNA between the populations of P. officinale subsp. officinale and P. officinale subsp. album, by extending the field sampling to one of the populations close to the classical location where P. stenocarpum had been described, El Escorial (Madrid) (Population Po7 in Figure
The populations measured and the DNA samples taken. The details of these populations are shown in Table
Populations | Number of plants | ||
---|---|---|---|
Species | Population (Table |
Measurements taken | DNA sample |
P. officinale subsp. album | Po1 | 4 | |
Po2 | 4 | ||
Po3 | 9 | ||
Po5 | 1 | ||
Po6 | 1 | ||
Po8 | 2 | ||
P. officinale subsp. officinale | Po4 | 12 | 1 |
Po7 | 1 |
The obtained sequences were aligned with CLUSTALW from Bioedit 7.2.5. (
The pairwise genetic distances between sequences were calculated with MEGA, version X (
By taking the morphological differences observed in leaves, inflorescences and fruits, its habitat and distribution in the humid and sub-humid thermomediterranean bioclimatic types as a basis, we distinguished P. officinale subsp. album as a new subspecies, after its comparison with all the revised herbarium specimens and data provided in the bibliography (Table
Comparison between the results obtained from the field measurements that we took with data reported in other studies about P. officinale subspecies: subsp. officinale (Figure
Origin of data | Field data |
|
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---|---|---|---|---|---|---|
Organ | album | officinale | stenocarpum | longifolium | paniculatum | brachyradium |
Habit (cm) | (24)41–98 (130) | 60–140(200) | 120 | (37)70–150(360) | 60–100 (120) | 250 |
Leaf | canaliculated | flat | flat | crested | canaliculated | canaliculated |
Last division basal leaf length (mm) | (31.3)47.9–83.7(94.4) | (20)30–60(100) | (27)40–85(95) | (13)35–80(165) | (17)24–35(53) | 20–55 |
Last division basal leaf width (mm) | 0.3–0.6(0.7) | (0.7)1–2(2.7) | (0.9)1–1.8(2) | (0.5)1–1.8(2.3) | 0.5–0.8(1) | 0.8–1.3(2.5) |
Bracts of umbels | 0–1(2) | (0)1–10 | 0–1(4) | (0)1–10 | 0–1(2) | (0)1–2(5) |
Umbel rays | (3)5–9(12) | (12)17–35(58) | (18)20–37(41) | (14)16–32(49) | 9–14(17) | 12–18 |
Umbel ray length (mm) | (2.8)8–43.8(83.7) | (11)30–85(150) | (40)49–90(100) | (18)25–75(108) | (24)30–60(90) | 15–40 |
Umbellule raylets | (7)10–16(18) | (14)18–35(50) | (12)31–30(34) | (7)15–36 (44) | ?? | 9–16 |
Colour of petals | white | yellow | yellow | yellow | yellowish | yellow |
Fruit length (mm) | (5.1)5.8–6.9(7.1) | 5.5–9 | 05–7 | 5.5–7 | 5.5–6 | 7.6–9.7 |
Raylet length/Fruit length | (0.3)0.5–0.8(1) | 2–6 | 1.5–4 | 1 | 1–3 | 2 |
Regarding the genetic results obtained, the ITS sequences of the studied specimens that belong to the populations of Xeresa, Benicasim, Tuejar and El Escorial were identical to one another. The length of the obtained consensus sequence was 603 base pairs. It is deposited in GenBank http://www.ncbi.nlm.nih.gov/GenBank). The accession number is KP681852.
This sequence is identical to all the P. officinale sequences in GenBank and has a comparable length when using sets of sequences with both 319 bp and 640 bp (with gaps) (Suppl. material
The briefness and the small number of characteristics that have been analysed in the studies have meant that our determination of these populations was complicated. With the exhaustive comparison that we have carried out, we have been able to identify the characteristics (Table
Conversely, the leaflets shape characteristics of the last divisions; flat, folded-crested or canaliculated and the shape of inflorescences: with terminal, pedunculate umbels with 17–35 rays that are 30–85 mm long or sometimes with panicled inflorescences or with umbels, some of which are sessile and lateral, with 5–9 rays measuring 8–43 mm, were determining factors for separating the subspecies. Thus, the album, brachyradium and paniculatum subspecies share the combination of canaliculated leaflets and panicled inflorescences or with sessile umbels and fewer rays; as opposed to the officinale and longifolium subspecies, which have flat or grooved leaflets and inflorescences with terminal umbels and more rays.
These values obtained in the field have been used for the description of P. officinale subsp. album (Table
Code (Table |
Character (Table |
P. officinale subsp. album | P. officinale subsp. officinale |
---|---|---|---|
Stem | |||
T1 | Plant height | (0.24)0.41–0.98(1.3) m | (0.27)0.42–1.26(1.74) m |
T2 | First ramification height | (0.07)0.14–0.39(0.5) m | (0.12)0.15–0.52(0.73) m |
T3 | T1/T2 ratio | (1.6)2–3.5(3.9) | 1.3–3.6(4.6) |
T4 | Diameter at the base of stem | 2.1–4.6(6.3) mm | (2)2.7–6.1(7.3) mm |
T5 | Diameter at the first ramification | 2–4.5(6.2) mm | (1.7)2.3–6.4(8.6) mm |
T6 | T4/T5 ratio | (0.8)0.9–1.1(1.2) | 0.9–1.2(1.3) |
Basal rosette leaf | |||
H1 | Leaf length, including petiole | (190)257–538(630) mm | (280)335–505(570) mm |
H2 | Number of leaf divisions (ternate) | 4–5(6) | 4–5(6) |
H3 | Leaf width at the base of the last division | (0.4)0.5–0.8(0.9) mm | (0.5)0.6–1(1.1) mm |
H4 | Leaf width at the apex of the last division | 0.3–0.6(0.7) mm | (0.6)0.7–1.1(1.4) mm |
H5 | Leaf division length | (31.3)47.9–83.7(94.4) mm | (45.6)47.6–76.7(85.6) mm |
H6 | H5/H3 ratio | (50.5)70.3–125.8(166.2) | (57.1)62–96.3(108) |
H7 | H4/H5 ratio | (3.2)4.1–10.7(17.2) ×10-3 | (10.2)11.4–18(19.4) ×10-3 |
H8 | Angle between contiguous leaf divisions of the last ternate | (30)32–66(90) degrees | (30)31–45 degrees |
H9 | Angle between lateral leaf divisions of the last ternate | (45)111–180 degrees | 45–138(180) degrees |
H10 | Length petiole leaf | (42)53–171(250) mm | (55)87–197(240) mm |
H11 | Diameter at its basis petiole leaf | 1.5–2.8(3.4) mm | (2.1)2.2–3.4(3.7) mm |
H12 | Diameter before first division (ternate) leaf | (1.5)1.9–3.5(4.4) mm | (2.7)2.8–4.4(5.2) mm |
Cauline leaves: First on stem from the basal rosette leaf | |||
H13 | Number of leaf divisions (ternate) | 3–4(5) | (2)3–5 |
H14 | Last leaf division length | (1.8)6.5–36.4(40.8) mm | 5–88.8(147.3) mm |
H15 | Length | (1.2)4.3–61.1(87.3) mm | 9.5–73.4(95.2) mm |
H16 | Diameter of the petiole at its basis | (0.8)0.9–2.2(2.6) mm | (0.7)1.2–2.9(3.1) mm |
H17 | Diameter before the first division (ternate) | (0.8)1–2.2(2.7) mm | (0.7)0.9–4.2(5.8) mm |
Cauline leaves: Second on stem leaf | |||
H18 | Number of leaf divisions (ternate) | 1–3(4) | 2–4 |
H19 | Length | 3.2–73.8(120) mm | 41–67.5(70) mm |
H20 | Diameter before the first division (ternate) | (0.5)0.7–1.7(2.1) mm | 1–2.4(2.5) mm |
H21 | Diameter at its basis | 0.9–2 mm | 1–2.1(2.2) mm |
H22 | Sheath length | (2)6.5–22.6(28.3) mm | 22–29.5(31.4) mm |
Umbel | |||
I1 | Bract length | (0.6)0.7 mm | (0.5)0.6 mm |
I2 | Bract width | 1.1(1.3) mm | 1.7(1.8) mm |
I3 | Number of bracts | 0–1(2) | 0–2(3) |
I4 | I1/I2 ratio | 3.5–7.6(9.1) | (3.2)4.3–10.5(12) |
I5 | Number of rays | (3)5–9(12) | (13)15–32(35) |
I6 | Length of internal rays | (2.8)4–24.3(42.6) mm | (5)6.2–24.3(29.7) mm |
I7 | Length of external rays | (15.7)17.7–52.6(83.7) mm | (21.4)23.5–71.2(82.7) mm |
Umbellule | |||
I8 | Number of raylets | (7)10–16(18) | (7)12–24(29) |
I9 | Length of internal rays | 0.2–6.5(13.1) mm | 2.1–21.7(35.3) mm |
I10 | Length of external rays | 0.3–17.6(35.1) mm | 5.2–43.6(71.2) mm |
I11 | Number of bracteoles | (4)6–9 | (6)7–9(10) |
Fruit | |||
F1 | Length | (5.1)5.8–6.9(7.1) mm | (3.7)5.9–8.1(8.5) mm |
F2 | Width | (2.6)3.3–4.7(5) mm | (2.6)3.3–4.4(4.8) mm |
F3 | Wing width of the marginal ribs | (0.2)0.4–1(1.1) mm | (0.4)0.6–1.1 mm |
F4 | F3/F2 ratio | (0)0.1–0.3 | 0.1–0.2 |
F5 | F1/F2 ratio | (1.2)1.3–1.9(2) | (1.4)1.7–2(2.1) |
F6 | Fructiferous raylet length | (1.8)2.8–5.3(6.1) mm | (3.8)5.8–10.4(11.7) mm |
F7 | F1/F6 ratio | (1)1.2–2.3(3.2) | (0.6)0.7–1(1.2) |
F8 | F6/F1 ratio | (0.3)0.5–0.8(1) | (0.8)1–1.3 |
F9 | F2/F6 ratio | 0.6–1.6(2.8) | (0.3)0.4–0.5(0.7) |
Values with statistically significant differences. Code refers to Table
Code | Organ | subsp. album | subsp. officinale |
---|---|---|---|
I5 | Number of rays (u.)* | (3) 5–9 (12) | (13) 15–32 (35) |
I8 | Number of raylets (u.)* | (7) 10–16 (18) | (7) 12–24 (29) |
F1 | Fruit length (mm)* | (5.1) 5.8–6.9 (7.1) | (3.7) 5.9–8.1 (8.5) |
F6 | Fruit raylet length (mm) * |
(1.8) 2.8–5.3 (6.1) | (3.8) 5.8–10.4 (11.7) |
F7 | Fruit length/ Raylet length * | (1) 1.2–2.3 (3.2) | (0.6) 0.7–1 (1.2) |
F8 | Raylet length/Fruit length ** | (0.3) 0.5–0.8 (1) | (0.8) 1–1.3 |
F9 | Fruit width/ Raylet length * | 0.6–1.6 (2.8) | (0.3) 0.4–0.5 (0.7) |
H7 | Last basal leaf division: width/length*** | (3.2) 4.1–10.7 (17.2) 10-3 | (10.2)11.4–8 (19.4) 10-3 |
H9 | Largest angle for external leaf divisions, last ternate (degrees) *** | (45) 111–180 | 45–138 (180) |
Peucedanum officinale subsp. album can be morphologically distinguished by the canaliculated leaflets, inflorescences without dominant terminal umbels, which are often sessile and lateral, with few rays (3) 5–9 (12), umbellules with (7) 10–16 (18) raylets, white-petalled flowers and fruits with a fructiferous raylet as long as or shorter than the length of the fruits.
Valencia, Tavernes de la Valldigna, eastern spurs of the Serra de les Agulles mountain range, on both ascents to Pic Massalari. 39,09°N, -0,284°W. 300 m alt. Rocky calcareous soil. 23-IX-2012. J. Martínez-Fort (Holotype: VAL 223161! Figure
Perennial plant with stem (0.24) 0.41–0.98 (1.3) m high and 2.1–4.6 (6.3) mm in diameter at the base, branching from the lower 1/3–1/2, striate, glabrous. Basal leaves 4- or 5- (6-) ternate, triangular in outline, (190) 257–538 (630) mm in length; petioles cylindrical, (42)53–171(250) mm long, sheathing at the base; linear terminal leaflets, (31.3) 47.9–83.7 (94.4) mm × 0.3 to 0.6 (0.7) mm, canaliculate, length/width ratio of (50) 70–126 (166) range, angle between the closest divisions of the terminals (30) 32–66 (90) degrees and (45) 111–180 degrees between the outermost leaf divisions. Cauline leaves, decreasing in size towards the apex of stems, but with enlarged sheaths. The first leaf on stem from the basal rosette leaf 3–4 (5) times ternate, with terminal divisions of (1.8) 6.5–36.4 (40.8) mm of length and canaliculated. The uppermost ones reduced to the sheath.
Inflorescence without a primary or dominant umbel, umbels arranged along the axis of inflorescence, sometimes sessile and lateral. Umbels compound, rays (3) 5 to 9 (12), inner rays shorter (2.8) 4- 24.3 (42.6) mm, outer rays (15.7) 17.7–52.6 (83.7) mm, bracts 0 or 1 (2), linear-triangular, (0.6) 0.7× 1.1 (1.3) mm. Umbellules with (7) 10 to 16 (18) raylets, bracteoles (4) 6 to 9, linear, gradually widened towards base. Flowers hermaphroditic, sepals 5 triangular, inconspicuous; petals 5, white, inflexed tips; stamens 5, alternate; stylopodium conical, similar in length to the styles; styles parallel at anthesis, becoming divergent in fruiting. Fruits dorsally compressed, elliptical, size (5.1)5.8–6.9 (7.1) × (2.6) 3.3–4.7 (5) mm, apex slightly off emarginated; mericarps homomorphic; median and lateral prominent ribs, apex slightly off emarginated; marginal ribs prominently winged, wings (0.2) 0.4–1 (1.1) mm, wide; commissural vittae 2; vallecular vittae 4; commissure very broad, from wing tip to wing tip. Raylets (1.8) 2.8–5.3 (6.1) mm long. The raylets length/fruit length ratio ranging from (0.3) 0.5–0.8 (1).
It is dispersed in the thermomediterranean sub-humid bioclima (
Flowering July to October.
SPAIN. Valencia: Alzira. Serra de la Murta, 30SYJ2934; on rocky outcrops, 25-IX-2011; J. Martínez-Fort (VAL 223162!, Figure
1 | Canaliculated leaflets (Figure |
2 |
– | Non-canaliculated leaflets (Figure |
4 |
2 | Habit of up to 1 (1.3) m; petals white or pale yellow; fruit length < 7 mm | 3 |
– | Habit of up to 1.5 m; petals yellow; fruit length > 7 mm | subsp. brachyradium |
3 | Umbels with 5 to 9 (12) rays, fructiferous raylets shorter than or equal to fruit length; petals white | subsp. album |
– | Umbels with 9 to 14 (17) rays, fructiferous raylets longer than fruit length, up to 3 times the length; petals pale to medium yellow | subsp. paniculatum |
4 | Fructiferous raylets equal or shorter than fruit length; folded or crested (concave) leaflets | subsp. longifolium |
– | Fructiferous raylets longer than fruit length; flat leaflets | subsp. officinale |
Types of bundle leaf A Flat-bundle leaflet of Peucedanum officinale subsp. officinale B Canaliculated-bundle leaflet of Peucedanum officinale subsp. album. A comparison of the other subspecies of P. officinale C Peucedanum officinale subsp. officinale D P. officinale subsp paniculatum. E P. officinale subsp. longifolium. Vouchers: C C. Pérez. (MA 152463) D L. Serra & A. Bort, (MA 623316) E, F Cernoch, (MA 310966).
Figure S1. Measured characters
Data type: species data
Explanation note: The figure shows the characters measured on the plants.
Table S1. Sequences obtained from GenBank to calculate pairwise genetic distances.
Data type: molecular data
Explanation note: Accession numbers of 85 ITS sequences downloaded from GenBank. The sequences from this study are in bold letters.