1

Preliminary key to the genus Lejeunea in Brazil

(draft version 30 April 2007)

M. Elena Reiner-Drehwald
University of Göttingen, Systematic Botany, [email protected]

Introduction
A first key to 28 selected species of Lejeunea in Brazil was published by Reiner-
Drehwald in Gradstein & Costa 2003.
In the present preliminary key 41 species of Lejeunea already recorded for Brazil or
expected to be found there are treated. It is a partial result of the revision of the
genus Lejeunea for Flora Neotropica (Reiner-Drehwald in prep.).
The key was prepared for
“An Overview of the genus Lejeunea in Tropical America – Workshop on Lejeunea” ,
a two-week workshop held in Recife (Universidade Federal de Pernambuco, Feb-Mar
2007). Comments or suggestions on the present key are welcome and will be used
for its improvement.

Acknowledgments: financial support for the workshop held in Recife was obtained
from the “Conselho Nacional de Desenvolvimento Científico e Tecnológico” (CNPq)
and for the revision of the genus Lejeunea from the “Deutsche
Forschungsgemeinschaft” (DFG RE 1348/4-1). I would like to thank specially Dr.
Kátia Cavalcanti Pôrto (Universidade Federal de Pernambuco, Recife) for inviting me
to this workshop, and the colleagues and students that I met in Recife not only for
their interest and helpful comments on “Lejeunea” but also for the hospitality and
friendship I received there.

Observations on the key:

The key presented here includes two species more than the key tested in Recife.
Most of the species have already been described and/or illustrated (references to
these are included). A few species are not published yet, the references on these will
be included as soon as the publications appear.
Small and sterile collections of Lejeunea are difficult to identify. Fertile specimens
with mature perianths are often needed to recognize the species.
Almost all species of Lejeunea often present undeveloped lobules; shoots with
reduced lobules are not always easy to identify.
Many species appear more than once in the key, in order to take into account the
different “shapes” they possess (e.g. Lejeunea controversa has perianth keels with
teeth, cilia and lacinia, but forms with nearly “entire” perianths are also often found,
as in the course in Recife …). But not all species can be included in a key with all the
possible variation found. Therefore, it is advisable to begin learning to identify
Lejeuneas with large and fertile collections.
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1. Underleaves entire ……………………...………….…………………. L. reflexistipula

1. Underleaves bifid ……………………………………………………….……………….. 2

2. Lobule tooth (2-)3-18 cells long, 1-4 cells wide ……………………..…..…………… 3

2. Lobule tooth 1(-2) cells long, 1 cell wide, or tooth undifferentiated ……….……..… 6

3. Plants with caducous leaves; tooth 5-7 cells long, 1-4 cells wide; underleaves 50-
75 % bifid, lobes ± divergent ……………………..…...………..….. L. ptosimophylla
3. Plants without caducous leaves; tooth (2-)3-18 cells long, 1-4 cells wide;
underleaves 40-60 % bifid, lobes straight ………….……………..…………………. 4

4. Lobule tooth (2-)3-4(-6) cells long, 1(2-) cells wide; perianth not compressed
…………………………………………………………………………..……... L. setiloba
4. Lobule tooth (3-)6-18 cells long, 1-4 cells wide; perianth dorsiventrally compressed
……………………………………………………………………..……………………… 5

5. Lobule tooth 2-4 cells wide, (7-)10-18 cells long, tooth usually extending more than
½ across the lobe …………………..…….………….…………………… L. spiniloba
5. Lobule tooth 1(-2) cells wide, (3-)6-11 cells long, tooth usually extending less than
½ across the lobe ………………………...…………….………………… L. trinitensis

6. Dorsal surface of lobe entirely or at least the distal half roughened (due to
mammillose cells, i.e. with the cell lumen extending into protuberances; ex
Echinocolea, see Ilkiu-Borges 2005) ……………………………………...………….. 7
6. Dorsal surface of lobe smooth ……………………………………..……………….…. 8

7. Plants 0.3-0.5 mm wide; entirely dorsal surface of leaves strongly roughened,

margin and keel of leaves strongly crenate to denticulate ………….. L. asperrima
7. Plants 0.4-0.8 mm wide; distal half of dorsal surface of leaves strongly roughened,
margin of leaves strongly crenate to denticulate, keel and dorsal base of leaves
smooth ……………………………………….…….……………….... L. subspathulata

8. Lobe apex acute to acuminate, (1-)2-3(-6) cells in a row …….……..… L. ramulosa

8. Lobe apex widely rounded to acuminate (then 1-2 cells in a row) ………..…..…… 9

9. Underleaf with a pronounced tooth 1-2(-3) cells long on each margin; cuticle
strongly papillose; perianth with teeth, cilia and lacinia ……….….…….. L. boryana
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9. Underleaf without marginal teeth or only occasionally with 1 small marginal tooth
(when underleaf margin often with teeth, then perianth without ornamentation, see
L. laetevirens); cuticle smooth or papillose; perianth with or without ornamentation
…………………………………………………………………………………………… 10

10. Lobule usually well developed, 2/3 the lobe length, strongly inflated throughout or
only along the keel and flattened near the lateral margin ………….... L. inflexiloba
10. Lobule when well developed smaller ……….……………………..……………….. 11

11. Plants with caducous leaves; dioicous ..… Key to Lejeunea with caducous leaves
11. Plants without caducous leaves; dioicous or autoicous ……..…………………… 12

12. Plants 0.3-0.8 mm wide, cuticle strongly papillose, plants absorb water slowly;
mostly sterile, shoots fragile, vegetative reproduction by fragmentation of the
plants; one of the most abundant and variable species in the Neotropics
……………………………………………………….…………………….. L. laetevirens
12. Plants 0.3-2.2 mm wide, cuticle papillose or smooth; fertile or sterile, vegetative
reproduction absent or when present via cladia, caducous leaves or regenerants,
not by fragmentation ………………………………………………..………………… 13

13. Perianth terete, without keels …………………………………………… L. capensis

13. Perianth 5-keeled, dorsal keel equally developed or shorter and less pronounced
than the others …………………………………………...…………..……………….. 14

14. Perianth keels variously ornamented with teeth, cilia, and lacinia ………...……. 15
14. Perianth keels entire, at most ± strongly crenate …………………….……….….. 28

15. Underleaves with lobes triangular to subulate, apex ciliate, 1-3(-5) cells in a row
…………………………………………….……………………………… L. controversa
15. Underleaves with lobes triangular, apex 1-2 cells in a row ………………..…….. 16

16. Leaf apex (obtuse-) acute, apiculate to acuminate ………………………………. 17

16. Leaf apex rounded ……………………………………………………...……………..21

17. Leaf margin strongly crenate due to mammillose marginal cells; cuticle smooth;
underleaves distant, 1.4-2.5 x the stem width; perianth keels 2-winged, wings
irregular, 1(2-4) cells wide, outermost cells elongate, rarely with few cilia (2-3 cells
long) ………..…………………………………………………………..……. L. raddiana
17. Leaf margin entire to slightly crenate; cuticle smooth to slightly to strongly
papillose; underleaves distant to contiguous to imbricate, 1.4-4 x the stem width;
perianth keels with cilia and lacinia or only occasionally with cilia .…………….... 18
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18. Leaf lobes ovate to lanceolate, apex acute to acuminate, (1-)2-3(-6) cells in a
row; underleaves contiguous to imbricate, 2.5-4 x the stem width; dioicous
………………………………………………………. ……………...………. L. ramulosa
18. Leaf lobes ovate to ovate-falcate, apex acute to apiculate, 1-2 cells in a row;
autoicous ……………………………………………………………...… …………….. 19

19. Underleaves 2.7-4 x the stem width; plants 0.9-1.2 mm wide ….. L. controversa
19. Underleaves 1.4-2.4 x the stem width; plants 0.4-1.0 mm wide ……..……….… 20

20. Perianth wings with lacinia (3-6 cells long, 2-3 cells wide) and cilia (1-3 cells
long); dorsal leaf margin without teeth ……………………….……….. L. grossitexta
20. Perianth wings without lacinia, seldom with short cilia (2 cells long); dorsal leaf
margin occasionally with few obtuse teeth near apex …….………. L. bermudiana

21. Perianth beak long, 100-140 m long ………………………………..... L. grossiretis

21. Perianth beak shorter, 25-50 um long (up to 75 um in L. cristulata) ……….…… 22

22. Leaf lobule usually well developed, 2/3 the lobe length; dioicous
…………………………………………………………………….…..……. L. inflexiloba
22. Leaf lobule when well developed smaller, < 1/3 or seldom up to ½ the lobe
length; autoicous ……………………………………….. ……….…………………… 23

23. Plants small, 0.3-0.7 mm wide …………………………………..……………..…… 24

23. Plants medium sized, (0.6-)0.8-1.6 mm wide …………………..……………….… 25

24. Perianths emergent ½ its length beyond the bracts; keels 2-winged, wings 1(-2-3)
cells wide, outermost cells of wing irregular, often elongated …………… L. elliottii
24. Perianths emergent 1/3 its length beyond the bracts; keels with teeth and cilia;
leaves distant; epiphyte (description based only on scanty type material)
…………………………………………………………………………..… L. spinuliflora

25. Underleaves 2.7-4 x the stem width, underleaf apices ciliate, 1-3(-5) cells long;
cuticle strongly papillose; perianth 5(-4)-keeled, dorsal keel absent or shorter and
less pronounced than the others ……………….…………….. …….. L. controversa
25. Underleaves 1.3-2.4 x the stem width, underleaf apices obtuse to acute, 1 cell at
the apex; cuticle smooth to slightly papillose; perianth 5-keeled, all keels ± equally
developed ………………….…………………………………………………………… 26
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26. Leaf lobe apex rounded to subacute, often recurved (also seen in dried condition
under the dissecting scope); leaf cells with small trigones, without intermediate
thickenings; on rocks in or near rivers or waterfalls, or on soil, almost always in
very humid or wet conditions, also partially submersed …………………….. L. laeta
26. Leaf lobe apex rounded and plane; leaf cells with small to medium sized trigones
and 0-2 intermediate thickenings; epiphytes, epiphylls or on rotten logs …..…… 27

27. Lobule inflated throughout, free margin involute, tooth 20-30 µm long; perianth
lateral keels somewhat expanded above, occasionally with short cilia (2 cells long)
……………………………………………………………...…………….….. L. cristulata
27. Lobule inflated along the keel and the lateral margin appressed to the lobe, tooth
50-55 um long; perianth lateral keels not expanded above, with teeth, cilia and
lacinia ………………………………………………….………………. L. cristulaeflora

28. Underleaves distant, strongly appressed on the stem, 75-95 um wide, 1-1.4 x the
stem width, lobes triangular, apex with 1-2 cells in a row, apical cells of the lobes
and occasionally also marginal cells extremely thin-walled and therefore collapsed
…………………………………………………………………..……………….. L. filipes
28. Underleaves distant to imbricate, not strongly appressed on the stem, wider than
the stem, 1.5-4.5(-7) x the stem width, all cells equally thick walled ..………..…. 29

29. Underleaf apices ciliate, 1-3(-5) cells long; cuticle strongly papillose; perianth
without ornamentation or with few teeth when poorly developed, but often densely
covered with cilia and lacinia ……………………….. ……………….. L. controversa
29. Underleaf apices not ciliate, 1(-2) cells in a row; cuticle smooth or papillose;
perianth keels entire, at most crenate ………………………………………………. 30

30. Underleaves (3-)4-7 x the stem width; gynoecia with 0-1(-2) innovations;
perianths 1-7 in a cymose row …………………...………………………...……..…. 31
30. Underleaves smaller, 2-4 x the stem width; gynoecia with 1-2 innovations (0
innovations not observed); perianths 1-3 in a row ………………………….……… 32

31. Underleaves 4-5.5(-7) x the stem width; cuticle papillose; dioicous; gynoecia with
0-1(-2) innovations; gynoecia 1-2(-3) in a row ……………….………….…. L. cerina
31. Underleaves 3-4.5 x the stem width; cuticle smooth; autoicous; gynoecia up to 5(-
7) in a row ………………………………………….…………………… L. obtusangula

32. Lobule usually well developed, 2/3 the lobe length, strongly inflated throughout or
only along the keel and distal half appressed on the lobe; dioicous
……………………………………………………………………………… L. inflexiloba
32. Lobule when well developed smaller, usually 1/3 the lobe length or smaller,
seldom up to ½ the lobe length (see L. oligoclada); autoicous or dioicous …...… 33
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33. Perianth without beak, apex constricted and slightly depressed; plants 0.35-0.55
mm wide ………………………………………………………………….…. L. erostrata
33. Perianth with beak, 1-3 cells long; plants usually > 0.6 mm wide (when smaller
then perianth with beak conspicuous) ………………………………………………. 34

34. Plants dioicous, often sterile; usually with vegetative reproduction via cladia,
caducous leaves, regenerants from leaf margin, fragmentation of the shoot …... 35
34. Plants autoicous, usually fertile; mostly without vegetative reproduction (seldom
with caducous branches in L. caulicalyx and L. angusta) ……………………….... 38

35. Cuticle strongly papillose, the plants absorb water slowly; usually sterile, often
with vegetative reproduction by fragmentation of the plants; underleaves often with
a tooth on each margin; one of the most common and variable species in tropical
America ……………..…………………………………………..………... L. laetevirens
35. Cuticle smooth or slightly papillose; sterile or fertile; underleaves without marginal
teeth …………………………………………………………………………………….. 36

36. Plants mostly sterile with vegetative reproduction via numerous cladia; oil bodies
numerous, 15-25(-50) per cell ……..……………………………..…….. L. cancellata
36. Plants often with perianths and androecia; vegetative reproduction via
regenerants and caducous leaves, seldom via caducous branches; oil bodies 2-4
per cell (Lejeunea sp. B) or oil bodies not known (L. oligoclada) ……………..… 37

37. Plants 0.4-0.8 mm wide; lobule ¼ to ½ the lobe length; underleaves 1.8-2.6 x the
stem width ……………………………………………………….…...…… L. oligoclada
37. Plants 1.0-1.8 mm wide; lobule < 1/5 the lobe length; underleaves (2.3-)3.0-4.2 x
the stem width …………………………………………….………..….. Lejeunea sp. B

38. Underleaves contiguous to imbricate, seldom distant, 2.5-4.5 x the stem width,
base rounded to slightly auriculate, seldom cuneate ……………….…….……….. 39
38. Underleaves distant, 1.3-2.5(-3) x the stem width, base cuneate ………………. 41

39. Oil bodies numerous, 20-40 per cell; underleaves usually wider than long; lobule
usually reduced, when well developed oval, lateral margin involute and not seen in
situ ……………………………………………………………….………. L. puiggariana
39. Oil bodies 3-8(-9-13) per cell ……………………………………..………………… 40

40. Cuticle papillose; lobe apex rounded; gynoecia 1-3 in a row ……………. L. flava
40. Cuticle smooth; lobe apex narrowly rounded, acute to acuminate, up to 2 cells in
a row; gynoecia 1-5(-7) in a row ………….………………………….. L. obtusangula
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41. Lobe apex subacute to apiculate, 1-2 cells in a row ……...……………..……….. 42

41. Lobe apex widely rounded to subacute ……………………..…………………….. 43

42. Leaf lobe on the dorsal distal half roughened due to convex to mammillose cells,
lobe margin also strongly crenate, without teeth; perianth keels 2-winged, wings
irregular, 1(2-4) cells wide, outermost cells elongate, rarely with few cilia (2-3 cells
long) ……………………………………………………….…………..…….. L. raddiana
42. Leaf lobe on the dorsal side smooth, lobe margin crenate and occasionally with
few obtuse teeth near apex; perianth keels 2-winged, wings 1 cell wide, crenate to
denticulate, seldom with short cilia (2 cells long) ……….………….. L. bermudiana

43. Leaf lobe apex almost always recurved when dry (as seen under dissecting
scope), extended or recurved when wet; underleaves at ± right angles to the stem;
cell walls thin, without intermediate thickenings; on rocks or soil, in or near small
rivers, often partially submersed …………………….....……. L. laeta (= L. geophila)
43. Leaf lobe apex plane or occasionally recurved when dry, extended or recurved
when wet; underleaves mostly appressed on the stem; cell walls ± thin, with or
without intermediate thickenings; epiphyll, epiphyte, rotten logs, seldom on soil or
rocks, not submersed …………………………………………………………………. 44

44. Underleaves 50-70 % bifid, lobes lanceolate, apex 1-2 cells in a row, sinus widely
U- to V-shaped, margin without or with a small tooth; leaf cells with large trigones
and intermediate thickenings; lobule when developed strongly inflated, lateral
margin involute; perianth dorsiventrally compressed, obcordate with lateral keels ±
dilated …………………………………………………………..…………. L. magnoliae
44. Underleaves at most 50 % bifid, lobes triangular, apex 1(-2) cells in a row, sinus
mostly V-shaped, margin without teeth; leaf cells with small to medium-sized
trigones, with or without intermediate thickenings; lobule variable; perianth not
dorsiventrally compressed, at most dorsal keel shorter and less developed than
the others ………………………………………………………………………………. 45

45. Perianth keels 2-winged, wings 1-3 cells wide, …………………….…………….. 46

45. Perianth keels entire or when 1-2-winged with wings only 1 cell wide …………. 47

46. Plants 0.3-0.7 mm wide; lobules ½-1/3 the lobe length; gynoecia with 1 sterile
innovation; perianth beak 35-50 µm long ……………………..…………… L. elliottii
46. Plants 0.5-1.2 mm wide; lobules 1/3-1/4 the lobe length; gynoecia with 1 (rarely 2)
sterile or fertile innovation, up to 3 gynoecia in a row; perianth beak 40-75 µm long
……………………………………………………………………………….. L. cristulata

47. Plants 0.4-0.9 mm wide; cuticle finely to strongly papillose; lobule up to 1/3 the
lobe length ………………………………………………………………..….. L. angusta
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47. Plants 0.7-1.8 mm wide; cuticle smooth to finely papillose; lobule ¼-1/7 the lobe
length ……………………………………………………………………..…………..… 48

48. Perianth shorter or as large as the bracts, seldom emergent up to ¼ its length
beyond the bracts; perianth beak 50-85 µm long; lobe margin strongly crenate due
to bulging cells; almost always growing on rotten logs …………...……. L. immersa
48. Perianth emergent ½ - 2/3 its length beyond the bracts; perianth beak 20-50 µm
long (50-85 µm long in L. monimiae); lobe margin slightly crenate to entire; on
rotten logs or epiphyllous ……………………………………………..……...………. 49

49. Perianth beak 50-85 µm long; almost always epiphyllous ………..… L. monimiae
49. Perianth beak 20-50 µm long; almost always on rotten logs or bases of trees
…………………………………………………………………………..……………….. 50

50. Lobule line of union with stem long, subequal to keel; lobule tooth 1-celled;
gynoecia terminal on short branches with one sterile innovation …… L. caulicalyx
50. Lobule line of union with stem not long; lobule variable, mostly with one basal
rectangular to triangular part and an apical tooth 1-4(-6) cells long and 1(-2) cells
wide; gynoecia terminal on branches or on the main shooth, with one often fertile
innovation, up to 3 gynoecia in a row ………………………………….….. L. setiloba

Key to Lejeunea with caducous leaves

1. Leaf lobule with a rectangular base and a long tooth (1-4 cells wide, 5-7 cells
long). …………………………………………….………………….… L. ptosimophylla
1. Leaf lobule tooth 1-celled . ……………………………....…………………………….. 2

2. Leaf lobule rectangular, apical margin 6-8 cells long; plants large, up to 2.5 mm
wide. …………………………………………………………….…….… Lejeunea sp. A
2. Leaf lobule rectangular to triangular, apical margin 2-3 cells long; plants smaller.
…………………………………………………………………….…………………….… 3

3. Leaf cells with triradiate trigones; bases of underleaves cuneate to quadrate; often
with hyaline rhizoids on leaf margins; plants 0.9-1.7 mm wide ...… L. tapajosensis
3. Leaf cells without triradiate trigones; bases of underleaves rounded to cuneate
……………………………………………………………………………..……………… 4

4. Leaves ± suborbicular; underleaves large, 3.3-6 x the stem width, imbricate, sinus
widely U- or V-shaped; plants 0.7-1.5 mm wide …………………… L. rionegrensis
4. Leaves ± ovate; underleaves smaller, distant to contiguous, sinus V-shaped. …... 5
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5. Leaf lobe apex widely rounded, plane; plants 0.6-1.1 mm wide. …… L. phyllobola
5. Leaf lobe apex rounded to subacute, often recurved; plants larger or smaller. ….. 6

6. Plants 0.9-1.8 mm wide; often with microphyllous branches with large collars;
lobules small, < 1/ 5 the lobe length. ………………………..…….… Lejeunea sp. B
6. Plants smaller, 0.3-0.8 mm wide; without microphyllous branches; lobules when
developed up to ½ the lobe length …………………………….…………..………….. 7

7. Underleaves 40-50 % bifid ……………………………….…………...… L. oligoclada

7. Underleaves 50-75 % bifid ……………………………..………..…… Lejeunea sp. C

References

Lejeunea photographs: http://www.drehwald.info/Lejeunea/lejeunea.html

1. L. angusta Reiner-Drehwald 1999

2. L. asperrima Ilkiu-Borges 2005
3. L. bermudiana Reiner-Drehwald & Goda 2000
4. L. boryana Reiner-Drehwald & Goda 2000
5. L. cancellata Reiner-Drehwald 2000
6. L. capensis Giancotti & Vital 1989
7. L. caulicalyx Reiner-Drehwald & Goda 2000
8. L. cerina Reiner-Drehwald 1999
9. L. controversa Reiner-Drehwald & Goda 2000
10. L. cristulaeflora Reiner-Drehwald & Goda 2000
11. L. cristulata Reiner-Drehwald & Goda 2000
12. L. elliottii Reiner-Drehwald & Goda 2000
13. L. erostrata Reiner-Drehwald & Goda 2000
14. L. filipes Reiner-Drehwald 2000
15. L. flava Reiner-Drehwald 2000
16. L. grossiretis Reiner-Drehwald & Goda 2000
17. L. grossitexta Reiner-Drehwald & Goda 2000
18. L. immersa Reiner-Drehwald 1999
19. L. inflexiloba Reiner-Drehwald & Goda 2000
20. L. laeta (= L. geophila) Reiner-Drehwald 2000
21. L. laetevirens Reiner-Drehwald 2000
22. L. magnoliae Schuster 1980 (as L. caespitosa)
23. L. monimiae Reiner-Drehwald 2000
24. L. obtusangula Reiner-Drehwald 2000 (as
Taxilejeunea obtusangula)
25. L. oligoclada Reiner-Drehwald 1999
26. L. phyllobola Reiner-Drehwald 2000
27. L. ptosimophylla Reiner-Drehwald 2000
28. L. puiggariana Reiner-Drehwald 2000
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29. L. raddiana Reiner-Drehwald & Goda 2000

30. L. ramulosa Reiner-Drehwald 1999
31. L. reflexistipula Reiner-Drehwald 2005
32. L. rionegrensis Reiner-Drehwald 1999
33. L. setiloba Reiner-Drehwald 2000
34. L. sp. A
35. L. sp. B
36. L. sp. C
37. L. spiniloba Schuster 1980 (as Rectolejeunea
spiniloba)
38. L. spinuliflora Reiner-Drehwald 1999
39. L. subspahulata Ilkiu-Borges 2005
40. L. tapajosensis Reiner-Drehwald 2000
41. L. trinitensis Reiner-Drehwald 2000

Giancotti, C. & D. M. Vital (1989). Lejeunea capensis Gott. (Hepaticae:

Lejeuneaceae) Disjunct Between South America and Africa. The Bryologist 93: 305-
307.

Ilkiu-Borges, A. L. (2005). A taxonomic revision of Echinocolea (Lejeuneaceae,

Hepaticae). Nova Hedwigia 80: 45-71.

Reiner-Drehwald, M. E. (1999). Catalogue of the genus Lejeunea Lib. (Hepaticae) of

Latin America. Bryophytorum Bibliotheca 54: 1- 101.

Reiner-Drehwald, M. E. (2000). Las Lejeuneaceae (Hepaticae) de Misiones,

Argentina. VI. Lejeunea y Taxilejeunea. Tropical Bryology 19: 81-131.

Reiner-Drehwald, M. E. (2005). On Amphilejeunea and Cryptognolejeunea, two small

genera of Lejeuneaceae (Jungermanniopsida), and two common neotropical
Lejeunea species. Nova Hedwigia 81: 395-411.

Reiner-Drehwald, M. E. & A. Goda (2000). Revision of the genus Crossotolejeunea

(Lejeuneaceae, Hepaticae). J. Hattori Bot. Lab. 89: 1-54.

Schuster, R. M. (1980). The Hepaticae and Anthocerotae of North America. Vol. IV.
1-1334. Columbia Univ. Press, New York.