zygodon (orthotrichaceae) in the iberian peninsula€¦ · co´ rdoba: sierra morena, camino de...

Zygodon (Orthotrichaceae) in the Iberian Peninsula

GRACIELA M. CALABRESE AND JESUS MUNOZ

Real Jardın Botanico, CSIC, Plaza de Murillo 2, 28014 Madrid, Spain

e-mail: [email protected]; [email protected]

ABSTRACT. The genus Zygodon in the Iberian Peninsula is revised and six species are

recognized: Zygodon catarinoi, Z. conoideus, Z. forsteri, Z. rupestris, Z. stirtonii and Z.

viridissimus. A key, descriptions, illustrations and maps of each taxon are given. Zygodon

stirtonii is reported for the Iberian Peninsula for the first time. Zygodon catarinoi and Z.

bistratus are conspecific, and the former predates Z. bistratus by two days.

KEYWORDS. Taxonomy, Zygodon, Orthotrichaceae, Spain, Portugal.

¤ ¤ ¤

Zygodon Hook. & Taylor includes some 90 species

mainly with tropical or subtropical distribution

(Calabrese 2006). After the world revision by Malta

(1926), recent detailed treatments have been

published for Europe (Dull 1985, 1992), Australasia

(Lewinsky 1990), Himalayas (Wilbraham & Long

2005), and southern South America (Calabrese

2006). Hill et al. (2006) accepted eight species and

one variety of Zygodon for Europe. Although not

particularly speciose in any European area, Zygodon

is more common and diverse towards the western

part of the continent. Thus, while only one species

grows in European Russia (Ignatov & Ignatova 2003),

Italy and Scandinavia harbor four species (Cortini

Pedrotti 2001, Lewinsky-Haapasaari 1998,

respectively), Germany and France five (Frahm &

Frey 1992, Husnot 1884–1894), and six each in Great

Britain (Smith 2004), the Iberian Peninsula (this

treatment) and northern Africa (Ros et al. 1999). In

the Iberian Peninsula the genus has received some

attention lately; Heras Perez et al. (1997) revised the

distribution and conservation status of Z. forsteri,

and two new species, Z. catarinoi and Z. bistratus,

were described (Garcıa et al. 2006, Calabrese &

Munoz 2006). These results mainly reflect the

growing interest in the rich and diverse epiphytic

flora of the Iberian Peninsula. They also demonstrate

the necessity of a revisionary study to resolve the

taxonomic and distributional problems encountered

around Zygodon in the area, which is the aim of the

present study.

Zygodon is considered in this treatment in a

broad sense, following Calabrese (2006). All Iberian

species belong to subg. Zygodon, except Z. forsteri,

which is included in subg. Codonoblepharon

(Schwagr.) Calabrese, characterized by having

smooth laminal cells and a double peristome. A

detailed description of the morphological variation

within the genus is provided by Malta (1926), and a

critical revision of taxonomic characters by Calabrese

(2006).

MATERIALS AND METHODS

This revision is based on field observations and

the study of 414 herbarium specimens from the

following herbaria: BCB, BM, BP, LISU, M, MA, MACB, MUB,

PAMP, S, SALA and VIT. Specimens were studied either in

water or mounted in Hoyer’s medium (Anderson

1954). Illustrations of leaves, cells, propagula and

capsules were made with a camera lucida on a light

microscope. All measurements were made from

moist material. Plant length refers only to the

gametophyte, costal width was measured in the lower

third, cell measurements include the walls, capsule

The Bryologist 111(2), pp. 231–247 0007-2745/08/$1.85/0Copyright E2008 by The American Bryological and Lichenological Society, Inc.

length excludes the operculum, and seta length

includes the portion embedded in the vaginula. The

species have been distinguished on the basis of

morphological characters. Terminology follows

Magill (1990). The regional distribution maps are

based solely on specimens studied by the authors.

KEY FOR IDENTIFICATION OF ZYGODON IN THE

IBERIAN PENINSULA

1. Leaf cells smooth; plants autoicous ................ 3. Z. forsteri

1. Upper and middle leaf cells papillose; plants dioicous .... 2

2. Leaf lamina bistratose, at least partially; leaf cells with

1–2(–3) tall, sometimes forked, papillae; peristome

absent ....................................................... 1. Z. catarinoi

2. Leaf lamina unistratose; leaf cells usually with 3–6 low,

simple papillae; peristome double or absent ............. 3

3. Propagula cylindrical, with only transverse septa; leaves

oval-lanceolate; peristome double ............... 2. Z. conoideus

3. Propagula fusiform, clavate or ovoid, either with only

transverse septa or with transverse and longitudinal septa;

peristome absent .................................................................. 4

4. Costa excurrent ......................................... 5. Z. stirtonii

4. Costa always ending below the apex .......................... 5

5. Leaves oval-lanceolate to lanceolate; propagula clavate to

ovoid, with transverse and longitudinal (straight or curved,

vertical or oblique) septa ......................... 6. Z. viridissimus

5. Leaves lanceolate to narrowly lanceolate; propagula fusi-

form-clavate or fusiform, only with transverse septa, rarely

with some longitudinal, oblique septa .......... 4. Z. rupestris

1. Zygodon catarinoi C. Garcıa, F. Lara, Sergio &

Sim-Sim, Nova Hedwigia 82: 248. 2006 [26 Feb].

TYPE: ‘‘Portugal, Baixo Alentejo: Barrancos, in

woodlands near Castelo de Noudar, 29SPC7027,

185 m, 13-04-2004, on trunks and branches of

Quercus rotundifolia Lam., C. Garcıa (LISU

196277).’’ Figs. 1, 9A

Zygodon. bistratus Calabrese & J. Munoz, The

Bryologist 109: 38. 2006 [1 Mar]. TYPE: ‘‘SPAIN.

BADAJOZ: La Morera, Sierra de Marıa Andres, pr.

La Morera, 460 m, MGRS Coord. 29SQC06,

corteza de Quercus ilex L. subsp. ballota Samp.,

Munoz s.n., 22-IV-1992’’ (holotype MA 20460,

isotype NY),’’ syn. nov.

Description. Plants to 0.6 cm tall, in loose turfs,

green above, reddish-brown to brown below. Stems

erect, sometimes branched. Rhizoids reddish-brown

to yellowish-brown, roughened to papillose, at the

base of stem. Leaves erect and somewhat contorted

when dry, erect-spreading when moist, increasingly

larger to the stem apex, lanceolate, keeled,

(0.82)1.0–1.47 3 0.20–0.32 mm, slightly decurrent;

apex acute, occasionally with small denticulations;

margins plane, entire, papillose throughout or

papillose above and smooth towards leaf base; lamina

partially or almost entirely bistratose in the upper

half, partially bistratose to unistratose in the lower

half. Upper leaf cells isodiametric, rounded-

hexagonal to subquadrate, thick-walled, 6–10 mm in

diam., towards the margins perfectly aligned,

papillose; papillae 1–2(–3) per cell, tall, subacute,

simple or forked. Basal cells subquadrate to short-

rectangular, shorter towards the margins, thick-

walled, sometimes nodulose, 10225(243) 3

7213 mm, usually smooth. Costa ending below the

apex, 25–38 mm wide in lower 1/3, dorsally smooth

in the lower half and papillose above, ventrally

smooth throughout; in cross-section with two ventral

guide cells and dorsal stereids. Propagula usually

abundant, fusiform to fusiform-clavate, walls hyaline

to yellowish-brown or brown, 32270 3

(152)20233 mm, with only transverse or somewhat

oblique septa. Dioicous. Perichaetial leaves

undifferentiated, slightly larger than vegetative leaves.

Vaginula pilose. Seta ca. 2.5 mm long, yellowish-

brown, dextrorse above, sinistrorse below. Capsule

erect, ellipsoidal, 1.0–1.2 mm long, with 8

longitudinal ribs; exothecial cells slightly

differentiated in longitudinal bands 3–4 cell rows

wide, with 5–6 rows of smaller cells around capsule

mouth; operculum conic at base, rostrate, beak

curved, yellowish, reddish at base; peristome lacking.

Spores papillose, 12–15 mm in diam., with weak

trilete mark. Calyptra not seen.

Ecology and distribution. Epiphytic on trunks

and bases of Quercus canariensis, Q. faginea, Q.

pyrenaica, Q. rotundifolia, Castanea sativa, Olea

europaea and Fraxinus angustifolia, usually mixed

with Zygodon rupestris; more rarely on Abies pinsapo,

and then only on the basal portion of the trunks. The

species is particularly abundant on isolated Quercus

rotundifolia trees in cleared woods, which in Portugal

are termed montados, and dehesas in Spain. The

species was only described in early 2006, but it has

proven to be the second most common species in the

Iberian Peninsula. It is also known from Morocco

(Garcıa et al. 2006), and is here reported for the first

232 THE BRYOLOGIST 111(2): 2008

Figure 1. Zygodon catarinoi. A. Habit when dry. B. Habit when moist. C. Leaves. D. Leaf apex. E. Leaf cross-sections: ei in the

middle of leaf, eii in the basal third of leaf. F. Basal leaf cells. G. Propagula. A, B from Cezon et al. s.n. (MA 31437); D, F, ci,eii, gi from

Munoz s.n. (MA 20460); cii, gii from Mateo & Cezon 95 (MA 29502); ei from Ederra s.n. (PAMP 7195).

Calabrese & Munoz: Iberian Zygodon 233

time from Greece (MUB 12186, base of Abies trunk),

which shows that it has a wider distribution in the

Mediterranean than previously realized.

Differentiation. Zygodon catarinoi is a small

taxon characterized by bistratose leaf laminae and the

tall, occasionally bifurcate papillae of the leaf cells. No

other species in the genus has bistratose leaf laminae,

not even partially, and thus this character at once

separates this taxon from any other Zygodon. The

species is macroscopically similar to Z. rupestris and Z.

viridissimus, which have the same stature and

distribution. However, besides the difference in leaf

stratosity, the latter species have (2–)3–5(–6) low and

usually blunt papillae per cell (Figs. 5, 8), while Z.

catarinoi has 1–2 tall papillae per cell (Fig. 1). Zygodon

catarinoi is also similar to the North American Z.

apiculatus Redfearn, but the latter has pluripapillose

laminal cells, oblong-lanceolate to oblong-ovate leaves

with recurved tips when moist (Redfearn 1967). Vitt

(2003) included Z. apiculatus and Z. rupestris under a

broadly defined Z. viridissimus.

Specimens examined.. PORTUGAL. ALTO ALENTEJO:

PNSSM, Porto Espada (Azinhal do Porto Espada), na

parte oeste, Sergio & al. 17.2 (LISU 165291a). SPAIN.

ALBACETE: Sierra del Calar del Mundo, lago de las

Truchas, Jimenez s.n. (MUB 1654); ALICANTE: Alcoy,

subiendo a la Font Roja, Casas s.n. (BCB 26740);

BADAJOZ: Barcarrota, Lara & Albertos s.n. (MA 35238);

CACERES: Villar del Pedroso, Barranco de la Venta,

base del Cerro Buen Amigo, Casas et al s.n. (BCB

49531); CADIZ: Sierra del Pinar, Grazalema, Guerra

s.n. (MUB 15837); CIUDAD REAL: Sierra Madrona, Pena

Rodrigo, pr. Fuencaliente, Fuertes s.n. (MACB 44420);

CORDOBA: Sierra Morena, camino de Trassierra a las

Ermitas, Oliva s.n. (BCB 11195); HUELVA: Encinasola,

arroyo Caba, cerca de Encinasola, 24-IV-1992 (BCB

31237, MA 20462, PAMP 7179); JAEN: Sierra del Pozo,

La Agracea, arroyo Agracea, Lopez-Soria s.n. (BCB

31621); MALAGA: Sierra de la Nieves, Guerra et al. s.n.

(MUB 18397); SALAMANCA: bajada a Valero, Elıas s.n.

(SALA 496); TOLEDO: Almendral de la Canada, Cezon et

al. s.n. (MA 31439); ZAMORA: Arribes del Duero,

encinar-enebral entre Marmoles y el rıo Duero,

Fernandez Mendoza 104 (MACB 081035).

2. Zygodon conoideus (Dicks.) Hook. & Taylor,

Musc. Brit. 71. 1818; Bryum conoideum Dicks.,

Pl. Cryptog. Brit. 4: 9, tab. 11, fig. 2. 1801. TYPE:

‘‘In Fagorum truncis, prope Inverary’’ (BM

918154!) Figs. 2, 9A


green above, dark green to brown below. Stems erect,

simple or branched. Rhizoids reddish-brown, usually

smooth, rarely slightly roughened, at the base of

stem. Leaves erect when dry, erect-patent when

moist, increasingly larger toward the stem apex, oval-

lanceolate, keeled, (0.582)0.7021.22 3

(0.192)0.2420.42 mm, slightly decurrent; apex

acute; margins plane, entire, papillose above and

smooth towards leaf base; lamina unistratose. Upper

leaf cells rounded-hexagonal to oval-hexagonal, 6

thick-walled, 10–15 mm in diam., papillose; papillae

(1–)2–3(–5) per cell, low, blunt, simple. Basal cells

hexagonal to short-rectangular, longer toward the

costa and shorter toward the margins, thick-walled,

the smaller 10–15 mm in diam., the larger 10228 3

10213 mm, usually smooth, rarely with 1–2 papillae.

Costa ending below the apex, 45–55 mm wide in

lower 1/3, dorsally smooth except near the apex

where short cells with few low papillae covering the

elongate cells, ventrally smooth throughout; in cross-

section with two ventral guide cells and dorsal

stereids. Propagula frequent, cylindrical, walls

hyaline, (622)1002150 3 (222)30235 mm, with

only transverse septa, filamentous structures of

2882305 3 20225 mm. Dioicous. Sporophytes not

seen from Iberian specimens; Merino (1916)

described it as having ovate capsules with a neck as

long as the urn, and peristome double. Albertos et al.

(2005) reported a specimen with sporophytes from

Lugo (northwestern Spain), without further detail.

The type specimen has setae 2–3 mm long, yellowish-

brown, dextrorse above, sinistrorse below. [Capsule

description after Lewinsky-Haapasaari 1998 and

Smith 2004.] Capsules ovoid to ellipsoid, striate,

sulcate when dry and empty; operculum obliquely

rostrate; peristome double, exostome teeth short,

fugacious, endostome segments 8, short, filiform,

sometimes rudimentary. Spores papillose, (16–)18–

20 mm in diam.

Ecology and distribution. In the study area the

species has been collected on Quercus robur and

Fraxinus excelsior in two widely separated areas in the

western and eastern extremes of the Cantabric coast.


Figure 2. Zygodon conoideus. A. Habit when dry. B. Habit when moist. C. Leaves. D. Leaf apex. E. Cells in the middle of leaf.

F. Leaf cross-sections. G. Basal leaf cells. H. Propagula. A–H from Toledo s.n. (PAMP 7000).


In all the Cantabric facade there are extensive

deciduous forests suitable for the species, the rarity of

which we think is due to undercollection.

The species has been reported from

northwestern Iberian Peninsula by Merino (1916,

Pontevedra, Spain; later recorded by Luisier 1918),

Allorge (1955, Navarra, Spain), and Sergio and

Schumacker (1992, Minho, Portugal), and from

central Portugal by Ade and Koppe (1942,

Estremadura). None was located for the present

study, although the identification of the former was

confirmed by Malta (1926).

Differentiation. Zygodon conoideus is a small

plant with broadly lanceolate leaves that are erect and

straight when dry, and cylindrical propagula of

chlorophyllose cells with hyaline walls, which make it

a distinctive species. Zygodon rupestris occasionally

has leaves of similar shape, although when dry they

are twisted and not regularly disposed like in Z.

conoideus (cf. Figs. 2, 5). Additionally, Z. rupestris is

a taller plant with fusiform propagula which usually

have colored walls. Zygodon conoideus has also been

confused in the past with Z. viridissimus, but the

latter has the leaves twisted and homomallous when

dry, and propagula with longitudinal septa.

Specimens examined. PORTUGAL. MINHO: Parque

Nacional de Geres, Mata do Ramiscal, Garcıa s.n.

(LISU 207639). SPAIN. LA CORUNA: rıo Ulla, coto Corero,

Casas & Brugues s.n. (BCB); NAVARRA: Lanz, Toledo s.n.

(PAMP), Irarzoz, de Miguel s.n. (PAMP 5380), Lizarrusti,

de Miguel & Ederra s.n. (PAMP 5381).

3. Zygodon forsteri (Dicks. ex With.) Mitt., Ann.

Mag. Nat. Hist. II, 8: 321. 1851; Bryum forsteri

Dicks., Pl. Cryptog. Brit. 3: 4, tab. 7, fig. 8. 1793,

nom. inval.; Bryum forsteri Dicks. ex With., Syst.

Arrangm. Brit. Pl., ed. 4, 3: 798. 1801;

Codonoblepharon forsteri (Dicks. ex With.)

Goffinet in Goffinet, A. J. Shaw, C. J. Cox,

Wickett & S. B. Boles, Monogr. Syst. Bot.

Missouri Bot. Gard. 98: 286. 2004. TYPE: ‘‘In

arborum truncis caesis, in Walthamstow, D.

Thom. F. Forster jun.’’ (BM). Figs. 3, 4, 9B

Note. The systematic position of Zygodon forsteri

has been debated since Goffinet and Vitt (1998)

recognized Codonoblepharon as a genus independent

from Zygodon. At that time they speculated that Z.

forsteri could have evolved from a Zygodon s.str.

ancestor, and explicitly excluded it from

Codonoblepharon. Later, this species was moved to

Codonoblepharon on the basis of new molecular

evidence (Goffinet et al. 2004). However, Calabrese

(2006) decided that Codonoblepharon should be

treated at the subgeneric rank within Zygodon.

Finally, Matcham and O’Shea (2005) excluded this

species either from Codonoblepharon or Zygodon on

the basis on its white rhizoids, wider cells, negative

KOH reaction of the laminal cell walls, and

temperate Northern Hemisphere distribution.

Description. Plants to 1 cm tall, in loose turfs,

dark green above, reddish-brown below. Stems erect,

usually branched. Rhizoids hyaline to greenish,

smooth, at the base of stem. Leaves erect and with

incurved apex when dry, erect-spreading when moist,

increasingly larger to the stem apex, lanceolate to

lingulate, strongly keeled, 1.4521.80 3

0.3720.55(20.75) mm, not decurrent; apex acute;

margins plane, entire; lamina unistratose. Upper leaf

cells isodiametric, rounded-hexagonal, 6 thick-

walled, (12–)15–27 mm in diam., smooth, towards

the margins perfectly aligned. Basal cells hexagonal,

rectangular to broadly ellipsoidal, longer to the costa

and shorter towards the margins, thin-walled, 25270

3 12230 mm, smooth. Costa ending below or at the

apex, 70–75 mm wide in lower 1/3, smooth

throughout; in cross-section with two ventral guide

cells and dorsal stereids. Propagula scarce,

cylindrical, walls hyaline, 4–5 cells, 852100 3

17220 mm, with only transverse septa. Autoicous.

Perichaetial leaves undifferentiated. Vaginula naked.

Seta 3–6 mm long, yellowish-brown, dextrorse

above, sinistrorse below. Capsule erect, ellipsoidal to

pyriform, ca. 1.7 mm long, with 8 longitudinal ribs;

exothecial cells differentiated in longitudinal bands

3–4 cell rows wide, with 1–2 rows of smaller cells

around capsule mouth; operculum conic at base,

rostrate; peristome double, exostome teeth in 8 pairs,

recurved and appressed to the capsule when dry,

dorsal surface (OPL) with papillae on a reticule in the

basal part and with papillae aligned or irregularly

disposed above, ventral surface (PPL) papillose;

endostome with basal membrane low, continuous,

striate-papillose, segments 8, dorsally keeled, PPL

smooth, IPL with papillose lines obliquely oriented at


Figure 3. Zygodon forsteri. A. Habit when dry. B. Habit when moist. C. Leaves. D. Capsule when dry. E. Propagula. A, B, ci, D

from Heras 79/94 (VIT 17432); cii from Mateo & Cezon 138 (MA 31435); ciii, E from Dismier s.n. (MA 3286).


the base, papillae longitudinally aligned at the central

part and irregularly disposed at the distal part. Spores

almost smooth, 10–13 mm in diam. Calyptra not seen.

Ecology and distribution. In the Iberian

Peninsula the species has been collected on trunks

and decaying wood of Quercus rotundifolia, Q.

faginea, Fagus sylvatica, Acer monspessulanum and

more rarely, terricolous at the bases of Abies pinsapo.

Differentiation. Zygodon forsteri has smooth leaf

cells, which readily distinguishes this species from

any other in the area.

Specimens examined. PORTUGAL. TRAS-OS-MONTES

E ALTO DOURO: Braganca, Rabal, Garcıa & al. MN2QrB

(LISU 198668). SPAIN. ALAVA: Boveda, hayedo karstico

del Leron, cara Norte, Heras 1071/84 (BCB 20428);

BARCELONA: Montnegre, a prop de can Mainou, Cros

(BCB 6379, BCB 9582); BURGOS: Huidobro, Fuertes (BCB

27934, MACB 35976); CANTABRIA: Vega de Liebana,

sobre Valmeo y bajo Porcieda, barranco de la Piedra,

Heras 79/94 (VIT 17432); CIUDAD REAL: Puebla de Don

Rodrigo, Cerro de Cornicabra, Cezon & Munoz s.n.

(MA 36144); HUESCA: Tella-Sin, barranco Irues-

Garona, Infante & Heras 391/02 (VIT 29301); LA RIOJA:

Ortigosa de Cameros, J. Martınez Abaigar s.n. (BCB

24469, PAMP 03128); MALAGA: Sierra de las Nieves,

Varo & Guerra s.n. (BCB 6380); TARRAGONA: Pauls,

Font de Sant Roc, Sergio et al. s.n. (BCB 12789);

TOLEDO: La Iglesuela, ‘‘Zoologico de Piedra’’, Mateo &

Cezon 138 (MA 31435).

4. Zygodon rupestris Schimp. ex Lorentz, Bryol.

Notizb. 32. 1865 & Verz. Europ. Laubm. 12.

1865, nom. nov.; Z. rupestris Schimp. ex Lorentz

& Molendo in Lorentz, Moosstud. 95. 1864,

nom. nud.; Z. viridissimus var. saxicola Lorentz &

Molendo in Lorentz, Moosstud. 95. 1864, hom.

illeg. TYPE: [Austria] ‘‘Rabenhorst, Bryotheca

europaea nu 626. Zygodon viridissimus var.

saxicola. Tirolis silvatica alpina pr.

Windismatrey, rupes valde adumbratae 34–

36009 S 1862 Molendo’’ (lectotype, M!).

Fig. 5, 6, 9C

Zygodon viridissimus f. borealis Correns, Unters.

Vermehr. Laubm. 117. 1899. TYPE: ‘‘a.

Buchenstamm bei Oberthal im wurttembergischen

Schwarzwald, zuerst VIII. 1896 gesammelt.

Herbarexemplare lagen mir von folgenden

Standorten vor : b. Westfalen, leg. H. Muller; c.

Harz, Bodethal, leg. Bertram; d. Neu-Ruppin, V.

1883 leg. Warnstorf (b, c, d im Herb. Univ. Tub.);

e. Hamburg, III. 1895 leg. Jaap, f. Vejle (Jutland),

V. 1882 leg. C. Jensen; g. Skane (Sudschweden),

VII. 1894 leg. H. Nilsson.’’ (none seen).

Zygodon viridissimus f. mediterranea Malta, Latvijas

Univ. Raksti 9: 119. 1924; Z. viridissimus var.

mediterraneus (Malta) Giacom., Ist. Bot. Reale

Univ. Reale Lab. Crittog. Pavia, Atti V, 4: 245.

1947; Z. baumgartneri f. mediterranea (Malta)

Cros, Lazaroa 5: 307. 1983. TYPE: Not indicated.

Zygodon baumgartneri Malta, Latvijas Univ. Raksti 9:

147. 1924; Z. viridissimus f. baumgartneri

(Malta) Malta, Latvijas Univ. Bot. Darza Darbi

1: 41. 1926; Z. viridissimus subsp. baumgartneri

(Malta) Dull, Bryol. Beitr. 8/9: 112. 1992, nom.

inval. TYPE: [Austria] ‘‘Obersteiermark, Hieflau,

im Gesause, an Tilia b. d. Kapelle unterm

Stalleck c. 550 m. 30 VI 1913 leg. J.

Baumgartner’’ (not seen).

Figure 4. Zygodon forsteri. A, B. Leaf apices. C. Leaf cross-

section. D. Cells in the middle of leaf. E. Basal leaf cells. A, C, D

from Heras 79/94 (VIT 17432); B, E from Dismier s.n. (MA 3286).


Figure 5. Zygodon rupestris. A. Habit when dry. B. Habit when moist. C. Leaf apices. D. Cells in the middle of leaf. E. Basal leaf

cells. F. Leaf cross-sections. A, B from Heras 5/96 (VIT 20300); ci, D, E from Molendo s.n. (M 63964); cii from Mateo & Cezon 213 (MA

31436); ciii from Guerra s.n. (MUB 2152); fi from Jensen s.n. (MA 6814); fii from Guerra s.n. (MACB 60913).


Figure 6. Zygodon rupestris. A–H. Leaf variability. I–N. Propagulum variability. A, J from J. M. Lopez & C. Lopez (MA 9418); B, N

from de Miguel & Viente s.n. (PAMP 5376); C from de Miguel s.n. (PAMP 5377); D, I from Viera s.n. (MA 5159); E, M from Fuertes s.n.

(MA 12796); F from Infante & Heras 866/00 (VIT 26676); G from Guerra s.n. (MUB 2152).



green above, reddish-brown below. Stems erect,

simple or branched. Rhizoids reddish-brown or

yellowish-brown, roughened or papillose, at the base

of stem. Leaves twisted when dry, erect-spreading

when moist, lanceolate to narrowly lanceolate,

keeled, (0.752)0.921.5 3 0.220.35 mm, slightly

decurrent; apex acute, sometimes long; margins

plane, entire, papillose above and smooth towards

leaf base; lamina unistratose. Upper leaf cells 6

isodiametric, rounded-hexagonal, thick-walled, 7–

10 mm in diam., papillose; papillae 3–5(–6) per cell,

low, blunt, simple. Basal cells subquadrate to

rectangular, shorter towards the margins, thick-

walled, transverse walls usually thicker than

longitudinal walls especially in the marginal cells,

7218 3 (72)10 mm towards the margins and 27233

3 10213 mm toward the costa, smooth. Costa

ending below the apex, 30–38(–50) mm wide in lower

1/3, dorsally smooth except at the tip where short

cells with few papillae cover the elongate cells,

ventrally smooth throughout; in cross-section with

two small ventral guide cells and dorsal stereids.

Propagula abundant, clavate, walls yellowish-brown

or hyaline, 572100 3 20235 mm, with transverse

septa, very rarely with few longitudinal oblique septa.

Dioicous. Perichaetial leaves undifferentiated, slightly

larger than vegetative leaves. Vaginula pilose. Seta ca.

3.5 mm long, yellowish-brown, dextrorse above,

sinistrorse below. Capsule erect, ellipsoidal,

6 1.2 mm long, with 8 longitudinal ribs; exothecial

cells slightly differentiated in longitudinal bands 3–5

cell rows wide, with 4–6 rows of smaller cells around

capsule mouth; operculum conical at base, rostrate;

peristome lacking. Spores papillose, 15–18 mm in

diam. Calyptra cucullate, smooth.

Ecology and distribution. Very common on

trunks of deciduous trees (e.g., Acer, Castanea,

Crataegus, Fagus, Fraxinus, Quercus, Ulmus), more

rarely on perennials like Buxus sempervirens, Laurus

nobilis or conifers (Abies pinsapo or Juniperus

thurifera), and on the latter mostly found on the

bases or rotten logs.

Differentiation. Zygodon rupestris is a variable

species, and the above description corresponds to the

type expression. Many taxa have been described on

the basis of the observed variability, and their

systematic position has also varied widely, either

considered at any infraspecific rank under Z.

viridissimus, or as independent species. Zygodon

viridissimus differs from Z. rupestris in having

propagula with many longitudinal septa, which are

only exceptional in the latter. Karttunen (1984)

resolved the nomenclatural difficulties around this

taxon.

The taxon described by Malta (1924) as f.

mediterranea from the Mediterranean mountains of

Europe differs in having distal cells slightly larger (to

13 mm in diam.) with thinner walls, basal cells

uniformly rectangular and with all the walls

uniformly thin, and more clearly fusiform propagula.

All these characters are well within the range of

variation of typical Z. rupestris, and can be observed

throughout the distributional range of this taxon

either together or uncorrelated, reasons that led us

not to consider this expression at any rank.

Zygodon baumgartneri was described to

accommodate small specimens with clavate

propagula lacking longitudinal septa, leaves broadly

lanceolate, more acute and with wider apices than Z.

viridissimus (Malta 1924). All these features fall into

the variability of Z. rupestris, as previously considered

by other authors (e.g., Dull 1985; Lewinsky-

Haapasaari 1998; Smith 2004), justifying the

synonymy.

Specimens examined. PORTUGAL. ALGARVE: Serra

de Monte Figo, pr. Murta, Casas et al. s.n. (BCB 9764);

ALTO ALENTEJO: proximo de Alagoa (a 1.5 km), Ribeira

das Macieiras, Sergio et al. s.n. (BCB 42343, LISU

165292); BAIXO ALENTEJO: Serpa, entre Quintos e

Cortes da Condessa, Sim-Sim & Sergio M60 (LISU

161404); BEIRA ALTA: Serra da Marofa, Bizarril, Sim-

Sim & Sergio M118 (LISU 161443); BEIRA BAIXA:

Pampilhosa da Serra, estrada para Orvalho, Sim-Sim

& Sergio M67 (LISU 161406); BEIRA LITORAL: Vila Nova

de Aurem, Casal Farto, pr. Maxieira, a 5 km de

Fatima, Casas et al. s.n. (BCB 7608, BCB 7769, BCB

7780); DOURO LITORAL: Santo Tirso, Senhora das

Valinhas, Sim-Sim & Sergio M43 (LISU 161407);

ESTREMADURA: Figueiredo, Casas et al. s.n. (BCB 7675);

MINHO: Braga, Vilar de Veiga, en el camping, Munoz

s.n. (MA 15678); RIBATEJO: Torres Novas, Vole de terra,

estrada de Pedrogao, Moitas Vendas, Casas et al. s.n.

(BCB 7803); TRAS-OS-MONTES E ALTO DOURO: Braganca,


saıda para Gimonte, Sim-Sim & Sergio M89 (LISU

161427). SPAIN. ALAVA: Llodio, ermita Santa Lucıa,

Heras 42/89 (VIT 11564); ALICANTE: El Maigmo, subida

antena radio 2, Ros & Moya s.n. (MUB 4370, MUB

4371); ASTURIAS: Covadonga, Brugues s.n. (BCB 6391);

AVILA: Candeleda-Arenas de San Pedro, rıo Muelas,

junto al camino de la garganta Lobrega, Albertos & al

s.n. (MA 13469); BADAJOZ: Cabeza la Vaca, Sierra de

Tentudia, 6 km W del monasterio hacia Cabeza la

Vaca, Munoz s.n. (MA 18054); BARCELONA: Montnegre,

Vallgorguina, Cros s.n. (BCB 9581). BURGOS: Huidobro,

Fuertes s.n. (MACB 29916); CACERES: Villar del Pedroso,

hoya del Guadarranque, Casas et al. s.n. (BCB 49529-

a); CADIZ: Medina-Sidonia, Sierra Blanquilla, arroyo

del Alisoso, Guerra s.n. (MUB 11325); CANTABRIA:

Rionansa, San Sebastian de Garabandal, Infante &

Heras 866/00 (VIT 26676); CASTELLON: Els Ports,

Fredes, Barranc del Salt, Casas et al. s.n. (BCB 13398).

CUENCA: entre Priego y Tragacete, rıo Escabas, Casas

s.n. (BCB 6394); GERONA: Vidra, La Creu de l’Arc,

Casas s.n. (BCB 19198). GUADALAJARA: Poveda de la

Sierra, Mateo & Cezon 213 (MA 31436); GUIPUZCOA:

Ataun, Baiarrate, Infante 655/93 (VIT 16520); HUELVA:

Sierra de Aracena, Alajar, Puerto de Alajar, Cano et

al. s.n. (MUB 7945); HUESCA: Los Corralones, Aragues

del Puerto, Juaristi s.n. (PAMP 7500). ISLAS BALEARES:

Mallorca, Escorca, Puig Masanella, Aedo s.n. (MA

17204); Menorca, Cala Galdana, barranco der Cala

Macarella, F. Koppe s.n. (BCB 44199); LA CORUNA:

Puentedeume, Caaveiro, Reinoso (BCB 7509, MACB

60912); LEON: Cremenes, Aedo s.n. (MA 17205); LERIDA:

Alt Urgell, vall d’Alinya, La Llosa, vessant N, Saez

23.2 (BCB 53198); MADRID: El Escorial, La Herrerıa,

Lopez & Lopez s.n. (MA 9418); MALAGA: Sierra Bermeja,

pinsapar de los Reales de Estepona, Brugues s.n. (BCB

34295); MURCIA: Yecla, Sierra de Salinas, pr. cerro

Laurel, Cano 1173 (MUB 15138); NAVARRA: Lizarrusti,

de Miguel & Ederra s.n. (PAMP 05375); ORENSE: Parque

do Xures, Cela, Casas et al. s.n. (BCB 47093); PALENCIA:

Branosera, rıo Rubagon hacia Barruelo de Santullan,

Infante & Heras 1254/04 (VIT 33473); SALAMANCA:

Valle de las Batuecas, Elıas s.n. (BCB 22615, SALA

1094); SANTA CRUZ DE TENERIFE: El Hierro, Cruz de los

Reyes, Losada et al. s.n. (BCB 26737); TARRAGONA: La

Cenia, El Mangraners, barranc de la Fou, Casas et al.

s.n. (BCB 14182); TERUEL: Rubielos de Mora, canon del

rıo Mijares, Molino de la Hoz, Infante & Heras 167/

02 (VIT 29074); TOLEDO: Almendral de la Canada,

Cezon et al. s.n. (MA 31451); VALLADOLID:

Roturas, rıo Almonte, Viera s.n. (MA 5159);

VIZCAYA: Guernica, Monte Erenosarre, Heras 506/89

(VIT 12034).

5. Zygodon stirtonii Schimp. in Stirt., Trans. Bot.

Soc. Edinb. 11: 75. 1871; Z. viridissimus subsp.

stirtonii (Schimp.) Dixon, Student’s Handb. Brit.

Mosses 236. 1896; Zygodon viridissimus var.

stirtonii (Schimp.) I. Hagen, Kongel. Norske

Vidensk. Selsk. Skr. (Trondheim) 1907(13): 15.

1908. TYPE: (BM?, GL orig.?). Figs. 7, 9A


green above, brown below. Stems erect, simple, rarely

branched. Rhizoids reddish-brown, roughened to

papillose, at the base of stems. Leaves twisted when

dry, erect-spreading when moist, narrowly lanceolate,

strongly keeled, (0.752)1.021.5 3 0.2220.30 mm,

not or slightly decurrent; apex acute, ending in a

long, strong mucro of ca. 0.15–0.16 mm long;

margins plane, entire, papillose above and smooth

towards leaf base; lamina unistratose, ending

asymmetrically at the apex. Upper leaf cells

isodiametric, rounded-hexagonal, thick-walled, 7.5–

9.0 mm in diam., papillose; papillae 3–4 per cell, low,

blunt, simple. Basal cells 6 isodiametric to short-

rectangular, longer toward the costa and shorter

toward the margins, thick-walled, 12225 3

7213(215) mm, smooth. Costa excurrent, 40–45 mm

wide in lower 1/3, dorsally smooth except at the tip,

where short cells with few papillae covering the

elongate cells, ventrally smooth throughout; in cross-

section with two small ventral guide cells and dorsal

stereids. Propagula clavate, usually hyaline,

sometimes yellowish-brown or brown walls,

(552)702114 3 30240 mm, with transverse and

longitudinal, vertical septa (usually in the central part

of the propagulum). Dioicous. Sporophytes not seen,

similar to those of Z. viridissimus according to Smith

(2004).

Ecology and distribution. Zygodon stirtonii was

collected and correctly identified for the Iberian

Peninsula for the first time by P. Heras and M. Infante,

although they did not publish the record. It is only

known from a single collection in Cantabria province (N

Spain), on a large Quercus ilex close to the sea shore.


Differentiation. Zygodon stirtonii is

unmistakable by its costa widened at the tip and

excurrent as a stout mucro. Moreover, it can be

separated from Z. viridissimus, its closest relative and

the only other species having propagula with many

longitudinal septa, by having smaller, highly

incrassate upper leaf cells (compare Figs. 7 D,E and

8 D,E).

Figure 7. Zygodon stirtonii. A. Habit when dry. B. Habit when moist. C. Leaves. D. Leaf apex. E. Cells in the middle of leaf. F. Leaf

cross-sections. G. Basal leaf cells. H. Propagula. A–G from Heras 815/93 (VIT 16680).


Figure 8. Zygodon viridissimus. A. Habit when dry. B. Habit when moist. C. Leaves. D. Leaf apex. E. Cells in the middle of leaf. F.

Leaf cross-section. G. Basal leaf cells. H. Propagula. A, B from Munoz s.n. (MA 15684); ci, D, E, G, hi from Infante 1037/92 (VIT

15689); cii, F, hiii from Ron & Soria s.n. (MACB 33006); ciii, hiv from Munoz s.n. (MA 15682), civ, hii from Munoz s.n. (MA 15681).


Specimen examined. SPAIN. CANTABRIA: Val de

San Vicente, Munorrodero, rıa de Nansa, Heras

815/93 (VIT 16680).

6. Zygodon viridissimus (Dicks.) Brid., Bryol. Univ.

1: 592. 1826; Bryum viridissimum Dicks., Pl.

Cryptog. Brit. 4: 9, tab. 10, fig. 18. 1801. TYPE:

‘‘In pascuis Scotiae: Rosshire; in arburom

truncis, et in muris, prope Kilcullen-bridge, in

Hibernia, D. Brown’’ (BM 918153!). Figs. 8, 9B

Description. Plants 0.4–0.9 cm tall, in loose

turfs, light green above, reddish-brown below. Stems

erect, simple, rarely branched. Rhizoids reddish-

brown, smooth to roughened, at the base of stem.

Leaves twisted and homomallous when dry, erect-

spreading when moist, lanceolate, weakly keeled,

(0.72)1.021.8 3 0.220.4 mm, slightly decurrent;

apex acute; margins plane, entire, papillose above

and smooth towards leaf base; lamina unistratose.

Upper leaf cells isodiametric, rounded-hexagonal,

thick-walled, 10–13 mm in diam., papillose; papillae

(2–)3–4(–5) per cell, low, blunt, simple. Basal cells

hexagonal, subquadrate or rectangular, shorter

towards the margins, uniformly thick-walled,

(102)17230 3 (72)10215 mm, smooth. Costa

ending below the apex, 37–48 mm wide in lower 1/3,

dorsally smooth except at the upper 1/3, where short

cells with few papillae covering the elongate cells,

ventrally smooth throughout; in cross-section with

two small ventral guide cells and dorsal stereids.

Propagula usually clavate, sometimes ovoid,

yellowish-brown or hyaline walls, (502)802120 3

(272)35248 mm, with transverse and longitudinal

(vertical and oblique) septa. Dioicous. Perichaetial

leaves undifferentiated, slightly larger than vegetative

leaves. Vaginula pilose. Seta ca. 2.5 mm long, yellowish-

brown, dextrorse above, sinistrorse below. Capsule erect,

ellipsoidal, 1.2–1.4 mm long, with 8 longitudinal ribs;

exothecial cells slightly differentiated in longitudinal

bands 3–4 cell rows wide, with 3–4 rows of smaller cells

around capsule mouth; operculum not seen; peristome

lacking. Spores papillose, 12–15 mm in diam. Calyptra

not seen.

Ecology and distribution. Zygodon viridissimus

was reported from scattered localities in northern

Spain (Galicia, Asturias and Pirineos) by Malta

(1926). We extend its distribution to all the

Cantabrian coast, and confirm its presence in a

highly disjunct area in southern Spain (Cadiz

province), already recorded by Casares Gil (1915).

Figure 9. Distribution maps of Iberian Zygodon. A. Circle 5

Z. catarinoi; star 5 Z. conoideus; square 5 Z. stirtonii. B. Star 5

Z. forsteri; circle 5 Z. viridissimus. C. Z. rupestris.


Differentiation. For separation of Z. viridissimus

from Z. rupestris and Z. stirtonii, see these taxa. Plants

of Z. viridissimus usually lack sporophytes in the

Iberian Peninsula; we have only studied one fertile

specimen.

Specimens examined. PORTUGAL. BEIRA LITORAL:

Cantanhede, proximo da saıda da Pena, Sergio & Sim-

Sim s.n. (LISU 157270); ESTREMADURA: Alcobaca, Covao

do Milho, Sergio et al. s.n. (BCB 15509); MINHO:

Esposende, Aoulia, Fonte Boa, Sim-Sim & Sergio M73

(LISU 161435); RIBATEJO: Alcanede, Vale da Trave, Sergio

et al. s.n. (BCB 11032); SANTAREM: Lagoa da Serea, pr.

Vila Nova de Ourem, Sergio et al. s.n. (BCB 11143); TRAS

OS MONTES-ALTO DOURO: Cintra, Casas s.n. (BCB 22616).

SPAIN. ALAVA: Vitoria (ciudad), parque de La Florida,

Ron & Soria s.n. (MACB 33006); ALICANTE: Alcoi, Font

Roja, Casas & Sergio s.n. (BCB 8466); ASTURIAS: Tapia de

Casariego, playa de la Paloma, pr. Tapia, Munoz s.n.

(MA 15684); CADIZ: Sierra del Aljibe, barranco de Puerto

Oscuro, Guerra s.n. (MUB 9899); CANTABRIA: Meruelo,

puente de la Venera, Munoz s.n. (MA 15681); LA CORUNA:

Rianjo, Casares Gil s.n. (MA 8113, MA 8700). LUGO:

Vivero, Casares Gil s.n. (MA 8110, MA 11520); VIZCAYA:

Trucıos, rıo Aguera, Infante 11/93 (VIT 15876a).

ACKNOWLEDGMENTS

We thank the curators of BCB, BM, BP, LISU, M, MA, MACB, MUB, PAMP, S,

SALA, VIT and Esther Fuertes for the loan of specimens. Jesus

Munoz was partially supported by the ‘‘Convenio Marco CSIC

– Junta de Comunidades de Castilla La Mancha’’ to study the

bryophyte diversity in this Autonomous Community. Katia

Cezon kindly allow us to report here her findings of Zygodon

from Castilla-La Mancha.

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zygodon (orthotrichaceae) in the iberian peninsula€¦ · co´ rdoba: sierra morena, camino de...

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