C Nova Hedwigia Vol.

100 (2015) Issue 3–4, 589–601

published online January 14, 2015; published in print May 2015
Article

A revision of Archilejeunea s.str. (Lejeuneaceae,

Marchantiophyta)

Xue-Qin Shi1,2 and Rui-Liang Zhu1,3*

1
School of Life Sciences, East China Normal University, 3663 Zhong Shan North Road,
Shanghai 200062, China
2
School of Life Sciences, Anhui Normal University, 1 Bei Jing East Road, Wuhu,
Anhui 241000, China
3
Shanghai Key Lab for Urban Ecological Processes and Eco-Restoration, East China
Normal University, 500 Dongchuan Road, Shanghai 200241, China

With 5 figures

Abstract: The genus Archilejeunea s.str. was taxonomically reviewed. Six species are recognized,
including five species known in tropical America and one new species, A. gradsteinii known only
from Africa. The new species differs from the closely related species A. fuscescens in the usually
apiculate leaf apex, mostly double subfloral innovations, and female bract lobule 1/4–1/3 as long as
the bract lobe. A key to the species of Archilejeunea is provided.
Key words: Africa, Archilejeunea gradsteinii, Hepaticae, new species, Ptychanthoideae, Sierra Leone.

Introduction

Archilejeunea (Spruce) Steph. was first described as subgenus Archi-Lejeunea Spruce in

the broadly defined genus Lejeunea Lib. Subsequently, Stephani (1888) reported a new
species, Archilejeunea erronea Steph., and Archi-Lejeunea was raised to the generic
level. However, Archilejeunea erronea was considered a synonym of Leucolejeunea
clypeata (Schwein.) A.Evans (Gradstein & Geissler 1997) [≡ Cheilolejeunea clypeata
(Schwein.) W.Ye & R.L.Zhu]. Although many authors usually cited the genus as
"(Spruce) Schiffn. 1893" and proposed to conserve "(Spruce) Schiffn. 1893" (Grolle
1995 p. 18, Gradstein & Geissler 1997), Söderström et al. (2014) have shown "(Spruce)
Steph. 1888" to be a valid genus name free of conservation and the type of Archilejeunea
sensu Spruce can be used for Archilejeunea Steph. 1888.

*Author for correspondence: rlzhu@bio.ecnu.edu.cn

© 2015 J. Cramer in Gebr. Borntraeger Verlagsbuchhandlung, Stuttgart, www.borntraeger-cramer.de
Germany. DOI: 10.1127/nova_hedwigia/2015/0246 0029-5035/2015/0246 $ 3.25
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Archilejeunea s.l. is a pantropical genus with about 20 species (Shi et al. 2014,
Frey & Stech 2009). It was divided into two heterogeneous subgenera, i.e., A. subg.
Archilejeunea (pycnolejeuneoid innovations of gynoecia and never reduced leaf lobules)
and subg. Dibrachiella (lejeuneoid innovations of gynoecia and often reduced leaf
lobules) (Gradstein & Buskes 1985, Gradstein 1994, Wilson et al. 2007). Traditionally,
Archilejeunea is characterized by the stems without an enlarged epidermis, isodiametric
leaf cells with unpigmented walls, simple-triangular to radiate trigones, usually
segmented oil bodies, presence of gynoecial innovations, and perianths with 4-5
smooth to weakly toothed keels (Gradstein 1994).
Recent molecular-phylogenetic studies reveal that Archilejeunea is not monophyletic
(Wilson et al. 2007), and that A. subg. Dibrachiella should be raised to the generic level
(Shi et al. in prep.). Morphologically, Archilejeunea s.str. (A. subg. Archilejeunea) is
separated from Dibrachiella by the pycnolejeuneoid innovations of gynoecia and never
reduced leaf lobules. Morphological-anatomical characters and molecular-phylogenetic
analyses reveal that Archilejeunea s.str. is sister to Verdoornianthus Gradst., a small
genus from Amazonia (Gradstein 1994, Shi et al. in prep.) and Dibrachiella is closely
related to the paleotropical Ptychanthus Nees and Spruceanthus Verd, and the East
Asian endemic Tuzibeanthus S.Hatt. Both genera have tropical Afro-American
distributions, but the range of Archilejeunea s.str. is much more restricted than that
of Dibrachiella.
Here we present a taxonomic revision of Archilejeunea s.str. Six species of Archile-
jeunea were recognized, including a new species, A. gradsteinii, discovered from West
Africa during our studies on the Archilejeunea specimens collected by E.W.Jones from
Sierra Leone in 1971.

Material and methods

About 300 herbarium specimens from BR, E, EGR, GOET, HSNU, PC, STU, and U were examined
in the present study. Specimens were observed, measured, and illustrated with Olympus BX43
microscope and Nikon Y-IDT drawing tube instructions. Measurements were made as described in Zhu
& So (2001) and Zhu & Gradstein (2005) and always represent the complete observed morphological
variation of the plants. Cells and cross sections of the stem were examined and illustrated using
cover slips, while shoots, leaves, underleaves, bracts, bracteoles, etc., were observed and illustrated
without cover slips. Habitat data on the species were obtained from herbarium labels and literatures.

Taxonomic treatment

Archilejeunea (Spruce) Steph., Hedwigia 27: 113. 1888.

Lejeunea subg. Archilejeunea Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 88. 1884. Lejeunea
subg. Archilejeunea Spruce sect. Monotropella Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 89.
1884. Archilejeunea subg. Monotropella (Spruce) Schiffn., in Engler & Prantl, Nat. Pflanzenfam.
1, 3: 130. 1893. Type: Archilejeunea ludoviciana (Lehm.) P.Geissler & Gradst. subsp. porelloides
(Spruce) Gradst. (≡ Lejeunea porelloides Spruce).
Plants glossy yellowish-brown or dark brown in herbarium specimens, 1-3 mm
wide. Stems slender, ventral merophyte 4-6(-8) cells wide; epidermis cells similar

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to medullary cells; subepidermis absent. Vegetative branching of the Lejeunea-type.
Microphyllous branches absent. Leaf margins entire; midleaf cells isodiametrical or
longer than width, trigones rather thick, radiate and sometimes confluent; oil bodies
segmented. Lobules never reduced, short rectangular with truncate apex and 0-1 teeth.
Underleaves imbricate, rarely distant, 3-6 × stem width. Dioicous, rarely paroicous (A.
badia). Androecia terminal or intercalary on elongated branches, bracts in (1-)4-11
pairs, bracteoles borne throughout the androecium. Gynoecia usually with a single,
repeatedly fertile pycnolejeuneoid innovation (mostly double gynoecial innovations
in A. gradsteinii), bracts and bracteoles entire; bracts as large as vegetative leaves or
larger, lobe apex rounded to apiculate, lobules usually lanceolate-acuminate and 2/3
of lobe length (1/4–1/3 of lobe length in A. gradsteinii), never reduced; bracteoles
ovate-oblong, apex entire to shortly bifid. Perianths (2-)4-5-keeled, keels smooth or
slightly toothed, the ventral keels sometimes reduced in A. fuscescens.
Archilejeunea s.str. consists of six species: Archilejeunea badia (Spruce) Steph.,
A. crispistipula (Spruce) Steph., A. fuscescens (Lehm. & Lindenb.) Fulford,
A. ludoviciana subsp. ludoviciana, A. ludoviciana subsp. porelloides, and A. nebeliana
Gradst. & Schäf.-Verw. from tropical America, ranging from Costa Rica to southern
Ecuador and eastern Brazil (Gradstein 1994); and A. gradsteinii from west Africa. The
species mainly inhabits tree trunks or branches, rarely rotted logs (e.g. A. fuscescens)
in lowland rainforests, common at altitudes of 0–550 m, sometimes up to 1075 m
(Schäfer-Verwimp et al. 2013).

Key to the species of Archilejeunea s.str.

1 Leaf apex narrowly and obtusely pointed; cells of leaf lobule thin-walled and almost colorless
..................................................................................................................................A. nebeliana
1' Leaf apex broadly rounded or apiculate; cells of leaf lobule moderately thick-walled and not
colorless ......................................................................................................................................2
2 Leaf apex apiculate or rounded; lobules of female bract 1/4–1/3 × lobe length; mostly double
gynoecial innovations; known from tropical Africa ............................................... A. gradsteinii
2' Leaf apex rounded; lobules of female bract 2/3 × lobe length; always single gynoecial innova-
tions (paired innovations also present on the base of shoot in A. ludoviciana subsp. porelloides);
known from tropical America .....................................................................................................3
3 Underleaf margins undulate; ventral leaf margin usually auriculate at the junction with the keel
............................................................................................................................. A. crispistipula
3' Underleaf margins not undulate; ventral leaf margin never auriculate.......................................4
4 Leaves suborbicular, lobe apex often incurved, the angle between ventral leaf margin and the
lobule keel usually less than 150°; ventral keels of perianth often reduced ............A. fuscescens
4' Leaves ovate-oblong, lobe apex plane, the angle between ventral leaf margin and the lobule keel
more than 150°; ventral keels of perianth well-developed .........................................................5
5 Paroicous; leaf lobules strongly swollen, lobular tooth 1–4 cells long and 2–4 cells wide at base
........................................................................................................................................ A. badia
5' Dioicous; leaf lobules rather flat, lobular tooth lacking or present.............................................6
6 Leaf lobules 1/3–1/2 × lobe length, lobular tooth indistinct ....A. ludoviciana subsp. porelloides
6' Leaf lobules 1/5–1/3 × lobe length, lobular tooth uniseriate, 1–2 cells long................................
................................................................................................A. ludoviciana subsp. ludoviciana

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Archilejeunea gradsteinii X.Q.Shi & R.L.Zhu sp. nov. Fig. 1
Type: Sierra Leone. Kenema District: Kambui Hills. Bambawo. 27 Mar. 1971, E.W.Jones 1528
(holotype, E! Barcode: 00018881, female).
Archilejeunea gradsteinii differs from A. fuscescens in its rounded or apiculate leaf apex,
often double subfloral innovations, and the lobule of female bract 1/4–1/3 × lobe length.
Dioicous. plants 1–1.5 cm long × 1.2–1.5 mm wide, yellowish-brown in herbarium
specimens. stems in cross section orbicular in shape, 125–150 µm in diam., composed of
12–14 thick-walled epidermal cells surrounding 21–23 similar medullary cells; ventral
merophyte four cell rows wide. Branching of the Lejeunea-type. leaves imbricate,
when dry widely spreading and flat, dorsal lobe ovate-oblong, ca. 0.75 × 0.5 mm,
apex rounded or usually apiculate, dorsal margin plane, arched, ventral margin plane
or slightly incurved, forming an angle of ca. 120–150° with the keel. Lobe cell thin to
moderately thick-walled, marginal cells quadrate to rectangular, 10–20 × 10–15 µm,
median cells suborbicular to slightly elongate, 15–33 × 14–25 µm, basal cells hexagonal,
longer than wide, 20–45 × 20–23 µm, trigones small to medium, simple-triangular,
intermediate thickenings 0–1 per wall; oil bodies not seen. Lobules invariably well-
developed, short-rectangular, ca. 1/3 × lobe length, apex truncate, with one short,
outwardly pointing tooth consisting of 1 cell, tooth sometimes lacking. unDerleaves
contiguous to slightly imbricate, flat, suborbicular or reniform, 0.25–0.32 × 0.36–
0.5 mm, 3–4 × stem width, apex truncate, margins plane, insertion line shallowly
curved. anDroecia terminal or intercalary on lateral branches or stem, bracts in 4–5
series, hypostatic, 0.5–0.6 × 0.3–0.4 mm, apex rounded, margins entire, antheridia
2 per bract; male bracteoles slightly larger than underleaves, borne throughout the
androecium. Gynoecia with 2(–1) pycnolejeuneoid innovations, bracts suberect, lobe
obovate to oblong, 0.8–1.0 × 0.5–0.65 mm, apex apiculate or obtuse, margins entire,
lobules rectangular to ligulate, sometimes with the apex extending into a long, weird-
looking extension, 1/4–1/3 × lobe length; bracteole oval, 0.6–0.8 × 0.3–0.57 mm,
apex obtuse to broadly rounded. perianth immature. sporophytes and veGetative
reproDuctive orGans not seen.

EtymoloGy: The species is dedicated to Dr. S.Robbert Gradstein, who has made an
outstanding contribution to Archilejeunea.
Archilejeunea s.str. was previously known only from tropical America. A. gradsteinii
is the first record of Archilejeunea s.str. from Africa. A. gradsteinii stands out by its
rounded or usually apiculate leaf apex, mostly double innovations, and smaller lobule
of the female bract, ca. 1/4–1/3 × lobe length. The species is similar to A. fuscescens
which is extremely common in the lowland rain forests of Amazonia and the Guianas
(Gradstein 1994). However, the latter differs in the constant rounded leaf apex, single
gynoecial innovation, and larger lobules of the female bract, ca. 2/3 × lobe length.
Moreover, the shoots of A. fuscescens are more robust (ca. 1.5–2.5 mm wide) and the
margins of the underleaves are often recurved.
The leaves and leaf lobules of Archilejeunea gradsteinii also resemble A. nebeliana,
which is the only other species in the Archilejeunea s.str. with pointed leaves. However,
in A. nebeliana, the leaves become long pointed acuminate and cells of leaf lobule are
colorless and thin-walled. Moreover, the female bracteoles in that species are bifid and
the gynoecial innovations are single.

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Fig. 1. Archilejeunea gradsteinii. A–B. Portion of shoots, ventral view. C–D. Leaves. E. Underleaf.
F. Cross section of stem. G. Leaf lobule. H. Female bracteole. I–J. Female bracts. K–L. Apical
marginal cells of leaf lobe. M. Median cells of leaf lobe. N. Basal cells of leaf lobe. (All from
holotype, E.W.Jones 1528.)

Specimens examineD: sierra leone. Kenema District: Kambui Hills. Bambawo. 29 Mar. 1971,
E.W.Jones 1547A (E).
Distribution: Known only from the type locality in Sierra Leone, on tree trunks.

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Archilejeunea badia (Spruce) Steph., Sp. Hepat. 4: 711. 1911. Fig. 2. A–F
Lejeunea badia Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 92. 1884. Type: Brazil. Rio Uaupés,
IX. 1852. Spruce L104 (n.v.).
For further synonyms see Gradstein (1994).
Illustration: Gradstein & Buskes (1985, p. 93, fig. 1b, e–f. as Archilejeunea juliformis (Nees)
Gradst.); Stephani (1985, icones no. 793, no. 815 as Archilejeunea recurvans (Spruce) Steph.).
Description: Spruce (1884, as Lejeunea badia), Stephani (1911), Gradstein (1994).
Archilejeunea badia is recognized by the 1) paroicous sexuality, 2) strongly inflated
leaf lobule with a remarkable tooth 3–5 cells long and (1–)2–4 cells wide at the base,
and 3) lobule apex of the female bract usually with a long uniseriate tip (Fig. 2F).
Archilejeunea ludoviciana subsp. porelloides is similar to A. badia, but differs in its
larger size (shoots 2.7–3.5 mm wide, only 1.5–2.5 mm wide in A. badia), dioicous
sexuality, and lack of a distinct lobular tooth.
Specimens examineD: brazil. Amazonas: Rio Negro, in arborum ramulis. Spruce s.n. isotype
of Lejeunea recurvans Spruce (BR, E); Panuré fl. Uaupés, in cortice. Spruce s.n. (E).
Distribution: Brazil and Guyana (Gradstein 1994). Often on small twigs, sometimes
on leaf surface.

Archilejeunea crispistipula (Spruce) Steph., Sp. Hepat. 4: 712. 1911. Fig. 2. G–K
Lejeunea crispistipula Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 93. 1884. Type: Brazil.
Amazonas: Rio Negro, Uanauaca, XII. 1851–1852, Spruce L60 (lectotype, designated by Gradstein
& Buskes 1985, MANCH, n.v.).
Illustration: Gradstein & Buskes (1985, p. 97, fig. 2. g–j); Stephani (1985, icones no. 797–798).
Description: Spruce (1884, as Lejeunea crispistipula), Stephani (1911), Gradstein &
Buskes (1985), Gradstein (1994).
Archilejeunea crispistipula is separated from other Archilejeunea species by its glossy
yellow plant, undulate underleaf margin, an angle of 90° between the ventral leaf margin
with the lobule keel, and moreover, the ventral leaf margin usually "auriculate" at the
junction with the lobule keel.
Archilejeunea crispistipula and A. gradsteinii are the smallest species in Archilejeunea,
both with shoots 1.2–1.5 mm wide and ventral merophytes four cells wide. A. gradsteinii
stands out by its usually apiculate leaf apex, plane underleaf margin, and mostly double
innovations.
Specimens examineD: brazil. Amazonas: Sao Gabriel fl. Negro, Spruce s.n. isosyntype of Lejeunea
crispistipula (BR, E), in Campina forest, Km 60 along Manaus-Caracarai road, Griffin et al. 971 (HSNU).
Distribution: Northern South America (Gradstein 1994). On tree trunks.

Archilejeunea fuscescens (Lehm. & Lindenb.) Fulford, Bryologist 45: 174. 1942. Fig. 3
Lejeunea fuscescens Lehm. & Lindenb., Nov. Stirp. Pug. 7: 16. 1838. Marchesinia fuscescens
(Lehm. & Lindenb.) Kuntze, Revis. Gen. Pl. 2: 837. 1891. Type: Peru (?). "in cortic. Chinae reg."
ex hb. Hampe (n.v.).

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Fig. 2. A–F. Archilejeunea badia. A. Portion of shoot, ventral view. B–C. Leaf lobule apex. D. Median
cells of leaf lobe. E. Leaf. F. Female bract. G–K. Archilejeunea crispistipula. G. Portion of shoot,
ventral view. H. Leaf. I. Underleaf. J. Leaf lobule. K. Female bract. [A–F from Spruce s.n. (BR),
G–K from Spruce s.n. (BR).]

For further synonyms see Gradstein (1994).

Illustration: Gradstein (1994, p. 56, fig. 14); Gradstein & Ilkiu-Borges (2009, p. 54, fig. 28: A–C).
Description: Gradstein (1994), Gradstein & Ilkiu-Borges (2009).
Archilejeunea fuscescens is widely distributed in northern South America (Gradstein
1994). The species is characterized by the glossy brownish color, suborbicular leaf

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Fig. 3. Archilejeunea fuscescens. A. Portion of shoot, ventral view. B. Leaf. C. Underleaf. D. Leaf
lobule. E. Female bracteole. F. Female bract. G. Perianth, ventral view. H. Cross section of perianth.
I. Median cells of leaf lobe. J. Cross section of stem. (A from H.J. Cornelissen, H. ter Steege & S.R.
Gradstein C070; B–J from S.R.Gradstein 4708.)

lobes, and often recurved underleaf apex. In addition, the ventral keels of the perianth
are sometimes developed weakly, and tend to be reduced.
The differences between Archilejeunea fuscescens and A. gradsteinii are shown under
the latter.
Specimens examineD: brazil. Amazonas State: road Manaus-Itacoatiara km 27, Reserva Florestal
Ducke. G.T.Prance et al. 11307 (BR, E). colombia. Amazon: Caquetá. L.V.Campos J5-C (HSNU).
costa rica. Heredia: "La Selva" Biological Station. A.Bernecker-Lücking 97–53. (E, EGR);
Pantarenas: Osa Peninsula Rincón. S.R.Gradstein 9348, 9345 (HSNU). Guyana. Mabura Hill: 180
km S of Georgetown. H.J.Cornelissen, H.ter Steege & S.R.Gradstein C070 (E, EGR); East Demerara
district: Timehri, Dakara creek, Thompson farm. S.R.Gradstein 4708 (BR, E).
Distribution: Bolivia, Brazil, Colombia, Costa Rica, French Guyana, Guyana, Peru,
Suriname, Trinidad, and Venezuela (Gradstein 1994). On tree trunks and branches,
occasionally on rotted logs.

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Archilejeunea ludoviciana (Lehm.) P.Geissler & Gradst., J. Hattori Bot. Lab. 75:
202. 1994.
Phragmicoma ludoviciana Lehm., Nov. Stirp. Pug. 10: 11. 1857. Type: Tropical America, "in cortice
Cinchonae," L.Dufour s.n. (n.v.).

Archilejeunea ludoviciana subsp. ludoviciana, Fig. 4. A–H

For synonyms see Gradstein (1994).
Dioicous. plants 5 cm long, 2.5–3 mm wide, dull green to yellowish-brown in
herbarium specimens. stems rather fragile, in cross section suborbicular in shape,
200–300 µm in diam., composed of 22–24 thick-walled epidermal cells surrounding
55–65 thick-walled medullary cells, epidermal cells not larger than medullary cells;
ventral merophyte 6–7 cell rows wide. Branching of the Lejeunea type, flagelliform
branches lacking. leaves obliquely spreading at an angel of 60–80°, subimbricate;
dorsal lobe ovate-oblong, 1.1–1.6 × 0.7–0.8 mm, apex rounded, dorsal margin plane
and arched, ventral margin slightly curved, forming a broad angle with the keel; margin
cells suborbicular to hexagonal, isodiametrical, 20–30 × 15–25 µm, median cells longer
than width, 30–40 × 20–25 µm, basal cells similar to median cells, 30–60 × 20–30 µm,
trigones small to medium, simply to radiate, intermediate thickenings 0–2 per
wall; oil bodies not seen. Lobules short rectangular, ca. 1/5–1/3 × lobe length, apex
truncate, with one sharp, 1–2(–3) cells long tooth. unDerleaves imbricate, rounded or
longer than width, 0.8 × 0.5–0.7 mm, 3–4 × stem width, apex truncate or emarginate,
margins plane or weakly recurved, entire, insertion line arched. anDroecia terminal
on lateral branches, bracts in 5–7 series, hypostatic, bracts lobe 0.80–1.00 × 0.50–
0.56 mm, apex rounded , margins entire, bract lobules 0.55–0.65 × 0.30–0.35 mm, 2/3
× lobe length, antheridia 2 per bract; male bracteoles smaller than underleaves, borne
throughout the androecium. Gynoecia with 1 pycnolejeuneoid innovation, usually
repeatedly fertile, the bracts and bracteoles in 2 series, bracts as long as vegetative
leaves but much narrower, suberect, inner bract lobe oblong-lanceolate, 1.75–1.90 ×
0.58–0.72 mm, apex obtusely pointed or acuminate, margins entire, lobules lanceolate,
1.45–1.50 × 0.37–0.40 mm, 2/3 × lobe length, apex apiculate; inner bracteole ovate-
oblong, 1.75–2.10 × 0.85–1.10 mm, apex bifid to 1/5 its length and the lobes narrowly
acute; outer female bracteole about 1/2-2/3 as long as inner bracteole, ovate, apex
shortly bifid and the lobes obtuse. perianths emergent, cylindrical-obovate, 2.2 ×
1.1 mm, with 5 almost smooth to irregularly subdentate keels. sporophytes not
observed.
Archilejeunea ludoviciana subsp. ludoviciana is closely related to A. ludoviciana subsp.
porelloides. The former, however, can be distinguished by the more slender plants and
smaller leaf lobules (ca. 1/5–1/3 × lobe length) with an obvious tooth. In Archilejeunea
ludoviciana subsp. porelloides, the plants are 2.7–3.5 mm wide, and the leaf lobules
are ca. 1/3–1/2 × lobe length, without a distinct tooth.
Gradstein (1994) mentioned an unusual trait in the species that the distal end of the
lobular free margin is truncate and connate with the leaf lobe across 1-2 cells. This
character was not seen in the specimens cited below.
Specimens examineD: colombia. Chocó: Nuqui, El Amargal. S.R.Gradstein 8870 (HSNU). Ecuador.

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Fig. 4. A–H. Archilejeunea ludoviciana subsp. ludoviciana. A. Portion of shoot, ventral view. B, F.
Leaves. C, E. Underleaves. D. Leaf lobule. G. Cross section of stem. H. Median cells of leaf lobe.
I–K. Archilejeunea ludoviciana subsp. porelloides. I. Portion of shoot, ventral view. J. Leaf. K.
Cross section of stem. [A–D from A. Schafer-Verwimp 31733, E–H from S.R.Gradstein 8870, I–K
from Spruce s.n. (BR).]

Napo: Tena, Estación Biológica Jatunsascha. A.Schäfer-Verwimp 31733 (PC, HSNU). venezula.
Juan de Dios 28/B (BR).
Distribution: Colombia, Ecuador, Panama, Venezula (Gradstein 1994). On tree trunks.

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Archilejeunea ludoviciana subsp. porelloides (Spruce) Gradst., Fl. Neotrop. Monogr.
62: 58. 1994. Fig. 4. I–K
Lejeunea porelloides Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 90. 1884. Archilejeunea
porelloides (Spruce) Schiffn., Hedwigia 33: 181. 1894. Type: Venezuela. Amazonas: San Carlos,
Spruce L44 (lectotype, designated by Gradstein & Buskes 1985, MANCH, n.v.).
For further synonyms see Gradstein (1994).
Illustration: Gradstein & Buskes (1985, p. 97 fig. 2a–f as Archilejeunea porelloides); Stephani
(1985, icones no. 812–813 as Archilejeunea porelloides).
Description: Spruce (1884, as Lejeunea porelloides), Stephani (1911, as Archilejeunea
porelloides), Gradstein & Buskes (1985, as Archilejeunea porelloides), Gradstein
(1994).
Archilejeunea ludoviciana subsp. porelloides is the largest species of the genus. It can
be easily recognized by the 1) robust size (2.7–3.5 mm wide, ventral merophyte 6–8
cells wide), 2) leaf lobule almost without a distinct tooth, and 3) cylindrical hyaline
papilla at the proximal margin of the leaf lobule. The species is similar to Archilejeunea
ludoviciana subsp. ludoviciana. For their differences see the latter.
An important morphological trait of A. ludoviciana subsp. porelloides is the absence
of a distinct tooth of the leaf lobule. This character, however, is not stable. In several
specimens (e.g. Venezuela. Amazonas: San Carlos. Spruce s.n. E00280573, E0028575)
a lobular tooth 1–2 cells long is usually developed. Gradstein (1994, p. 60) reported
leaf lobules with or without a tooth in a specimen of A. ludoviciana subsp. ludoviciana,
and considered the two subspecies are closely related. Our studied confirmed his
speculation.
The differences between Archilejeunea ludoviciana subsp. porelloides and A. badia
are given under the latter.
Specimens examineD: brazil. Amazonas: Km 130 along Manaus-Caracarai road, Rio lages, in Campina
forest. Griffin et al. 418 (HSNU). columbia. Amazonas: Rio Caquetá, near Araracoara. I.Wolf 1615
(HSNU). venezuela. Amazonas: San Carlos. Spruce s.n. (BR, E).
Distribution: Brazil, Columbia, Ecuador, Peru, Venezuela (Gradstein 1994). On tree
trunks and twigs.

Archilejeunea nebeliana Gradst. & Schäf.-Verw., Cryptog. Bryol. 33(2): 108. 2012.
Fig. 5
Type: Ecuador. Zamora-Chinchipe: ca 5 km S of Zamora, Parque Nacional Podocarpus, entrance Rio
Bombuscara, sendero Mirador, 4°06,831'S, 78°58,017'W, on thin trunks in submontane rain forest,
1075 m alt., 25 Jan. 2011, A.Schäfer-Verwimp & M.Nebel 31924 (n.v.).
Illustration: Gradstein & Schäfer-Verwimp (2012, p. 109, figs. 1–17).
Description: Gradstein & Schäfer-Verwimp (2012).
Archilejeunea nebeliana is easily recognized by the 1) narrowly and obtusely pointed
leaves, 2) thin-walled leaf lobule cells, and 3) narrowly elongate female bracts.
The differences between Archilejeunea nebeliana and A. gradsteinii are given under
the latter.

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Fig. 5. Archilejeunea nebeliana. A. Portion of shoot, ventral view. B–C. Leaves. D. Underleaf.
E. Cross section of perianth. F–G. Female bracts. H. Female bracteole. I. perianth, ventral view.
J. Leaf lobule. K. Median cells of leaf lobe. L. Cross section of stem. (All from M.Nebel & A.Schäfer-
Verwimp 111599.)

Specimens examineD: ecuaDor. Zamora-Chinchipe: Parque Nacional Podocarpus, Zugang Rio

Bombuscara. M.Nebel & A.Schäfer-Verwimp 111599 (STU).
Distribution: Ecuador (Gradstein & Schäfer-Verwimp 2012). On tree trunks or
branches.

Acknowledgments

We thank the curators and staff of BR, E, EGR, GOET, HSNU, PC, STR, and U for lending specimens
including types for our study. Special thanks also go to the curators of E for permission to extract
DNA from herbarium specimens, and to L.Campos (Bogotá), S.R.Gradstein (PC), D. Long (E), and
T.Pócs (Eger) for sending material for our study. This research was sponsored by the National Natural
Science Foundation of China (nos. 31370238 and 31170190), Special Program for National Basic
Work of the Ministry of Science and Technology of China (No. 2012FY110600).

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Manuscript submitted September 6, 2014; accepted October 26, 2014

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