Phycologia (1996) Volume 35 (5),470-483

The typification and status of Leptophytum

(Corallinaceae, Rhodophyta)
L. DOWEL

AND

S. WEGEBERG1

Department of Phycology, Botanical Institute, University of Copenhagen,

(Z)ster Farimagsgade 2D, DK-/353 Copenhagen K, Denmark

L. DOWEL AND S. WEGEBERG. 1996. The typification and status of Leptophytum (Corallinaceae, Rhodophyta). Phycologia
35: 470-483.

The status of the genus Leptophytum Adey has been uncertain, because the location of the nomenclatural type, Lithophyllum
laeve Stromfelt nom. illeg., was unknown for years. In order to solve the problem of the missing type, new material was
collected at the type locality of Lithophyllum laeve, Eyrarbakki, South Iceland. Part of the holotype has recently been found
but does not show any diagnostic characters. The material collected at the type locality agrees well with Stromfelt's de
scription of Lithophyllum laeve, and an epitype is designated. The epitype is found to be conspecific with Phymatolithon
lenormandii (Areschoug in 1. Agardh) Adey. Leptophytum laeve Adey, the type species of Leptophytum, therefore becomes
a nomenclatural synonym of P. lenormandii and Leptophytum a nomenclatural synonym of Phymatolithon Foslie. The status
of the genus Leptophytum and the features distinguishing the genus from Phymatolithon are discussed.
Leptophytum laeve must be renamed however, because Adey's concept of it is taxonomically different from P. lenormandii.
The lectotype of Lithothamnion tenue Rosenvinge is therefore designated and redescribed. Adey's concept of Leptophytum
laeve is in full accordance with the lectotype of Lithothamnion tenue, which belongs to the genus Phymatolilhon Foslie,
and the new combination Phymatolithon tenue (Rosenvinge) Diiwel et Wegeberg is proposed. The holotype of Lithothamnion
foecundum Kjellman has been examined, and the new combination Phymatolithonfoecundum (Kjellman) Diiwel et Wegeberg
is also proposed.

INTRODUCTION
In 1966 Adey erected the genus Leptophytum using Litho
phyllum laeve Stromfelt nom. illeg. (1886) as the nomencla
tural type. The binomial L. laeve StromfeIt (1886) is a later
homonym of Lithophyllum laeve Kiitzing (1847) and therefore
illegitimate according to Art. 53.1 of the International Code
of Botanical Nomenclature (ICBN, Greuter 1994). Adey
(1966) did not examine the type material of L. laeve StromfeIt,
the location of which was unknown for years (Foslie 1895;
Woelkerling 1988; Chamberlain 1990). The taxonomic agree
ment between Leptophytum laeve Adey (1966) and L. laeve
Stromfelt is thus uncertain, rendering the status of the genus
Leptophytum Adey uncertain as well.
In the summer of 1992 we collected material from the type
locality of L. laeve Stromfelt in an attempt to designate a
neotype. However, part of the holotype, a single slide, was
recently rediscovered by W.J. Woelkerling in the Swedish Mu
seum of Natural History, Stockholm, during completion of
this study. The slide is supplied with complete collection data
by H. Stromfelt, and we have compared it with the newly
collected material from the type locality.
As Stromfelt (1886) indicated that his new species was sim
ilar to Lithophyllum lenormandii (Areschoug in J. Agardh)
Rosanoff (1866) ( = Phymatolithon lenormandii (Areschoug
in J. Agardh) Adey (1966, we also have examined the type
material of this species. In addition, the type material of Litho
thamnion foecundum Kjellman (1883) ( = Leptophytum foe
cundum (Kjellman) Adey (1966 and the herein designated
lectotype of Lithothamnion tenue Rosenvinge (1893) were exI

Author for correspondence.

470

amined as the two species in many respects correspond to

Adey's (1966) concept of Leptophytum laeve.

MATERIALS AND METHODS

Field collections were made at Eyrarbakki, Iceland, the type
locality of Lithophyllum laeve Stromfelt, on 4 June 1992. The
shore consisted of lava. The tidal amplitude was 4 m and at
low tide large, connected pools appeared. Coralline algae ad
herent to small lava stones were collected from the mjd- to
low littoral zone (about 200 m from the upper shore) as de
scribed by Stromfelt (1886). Collections were preserved in 5%
formaldehyde in sea water for 24 h. Before storing in 7 : 2: I
solution of ethanol, distilled water, and glycerol, the speci
mens were rinsed in cold tap water. Specimens are deposited
at C2 and ICEU.
We examined the following herbarium material:
(I) The holotype (S2) of Lithophyllum laeve Stromfelt (Fig.
I).
(2) The lectotype (LD2) of Melobesia lenormandii Areschoug
in J. Agardh, herbarium no. 50674 (I) A, B, & C (Fig. 2).
(3) A dried specimen (C) identified by L.K. Rosenvinge as
Lithothamnion tenue Rosenvinge, herbarium no. 879 (Fig.
3).
2 C, Botanical Museum, Copenhagen; rCEL, Icelandic Museum of
Natural History (Department of Botany), Reykjavik; LD, Botanical
Museum, Lund; S, Swedish Museum of Natural History (Department
of Cryptogamic Botany), Stockholm; UPS, Botanical Museum, Uni
versity of Uppsala. Herbarium abbreviations according to Holmgren
& Keuken (1974).

Diiwel and Wegeberg: Typification and status of Leptophytum

1b

1a

471

1c

'S"H X ID ca- 6/l'

(,M!q041U1)OU01SUO
'qreJ,.{3!lh.reJodS
'ds'U"IW9lIS"H

elo8f!'WIlJl,{qOOqll1

TYPUS

2cm

Figs la, b,

100 fJm

e. Holotype (S) of Lithophyllum laeve Stromfelt.

Fig. la. The holotype slide with H.F.G. Stromfelt's original label. The slide has been provided with an English translation. The note in the

parentheses: 'In the bay?' refers to H.F.G. Stromfelt's note: ' . . . i fjrere', which, however, means 'in the littoral zone'.
Fig. lb. Bisporangium.
Fig. Ie. Pore plate of a multiporate asexual conceptacle.

(4) The type collection (UPS2) of Lithothamnion foecundum

Kjellman. We have sectioned the specimen marked 'orig.
ex.' (Fig. 4).
Material was prepared for microscopy by methods adapted
from Woelkerling (1988). For light microscopy, small frag
ments of the selected specimens were cut off with a razor blade
and decalcified in 0.3 M HCP for approximately 24 h. The
material was rinsed once or twice in distilled water, stained in
2% KMn04 for 15-20 min, and rinsed again in distilled water
prior to the dehydration process. The material was dehydrated
consecutively in 30, 60, 90, and 99.99% ethanol at 20-min in
tervals. Finally, the material was embedded in LR-white resin
(London Resin Company, UK) in three steps: LR-white resin
overnight in stoppered vials, fresh LR-white resin for 1-2 h in
stoppered vials, and fresh LR-white resin in PVC Multi Well
Research Trays (Acrylic Industries Pty Ltd, Australia) covered
to exclude oxygen during resin hardening. The trays were
placed in a cold oven and subsequently heated to 58C, for
20-24 h. Sections of 5 [.Lm thickness were cut with a glass
knife on a Reichert Jung 2050 Supercut microtome, placed on
slides, stretched with Histoclear (BN, Denmark), and mounted
in Eukitt (Bie & Berntsen, Denmark).
For scanning electron microscopy, small fragments of the
specimens were cut off, air dried, and mounted on aluminium
stubs with Scotch doublestick tape (3M). The material was
coated once with gold and examined in a Phillips Scanning
Electron Microscope 515 at 25 kY.
RESULTS
Dimensions and other characteristics of the taxa studied
are summarized and compared in Tables 1-4.

Lithophyllum laeve Stromfelt

Historical background (Table 1)
Stromfelt (1886) described a thin calcified crust, which he
3 It should be noted that HCl must not be used as a decalcifier on
material that has been preserved in formaldehyde. The combination
of HCI and formaldehyde makes chlorine compounds that are carci
nogenic.

named Lithophyllum laeve, on small lava stones in the littoral

zone at Eyrarbakki, south Iceland. The binomial was, how
ever, a later homonym of Lithophyllum laeve Kiitzing (1847)
and is therefore illegitimate. Stromfelt described his new spe
cies as being similar to Lithophyllum lenormandii (Areschoug
in J. Agardh) Rosanoff (1866) ( = Phymatolithon lenormandii
(Areschoug in J. Agardh) Adey (1966, but found, however,
that the species were different in two ways. First, he observed
only bisporangia in L. laeve, whereas L. lenormandii had both
bi- and tetrasporangia. Second, he found L. laeve's sporangia
to be twice the size of those of L. lenormandii.
Foslie (1890) found specimens, which he identified as L.
laeve Stromfelt. Foslie, however, like Stromfelt (1886), con
sidered it difficult to distinguish the taxon from L. lenorman
dii. He later (Foslie 1891) reduced L. laeve Stromfelt to a form
of L. lenormandii, viz. L. lenormandii f. laeve (Stromfelt) Fos
lie. However, as Stromfelt's taxon is illegitimate it cannot
serve as basionym, and L. lenormandii f. laeve (Stromfelt)
Foslie is therefore also illegitimate. The specimens Foslie
(1891) referred to were in agreement with Stromfelt's descrip
tion (1886) except that the sporangia were larger and tetra
sporic. However, Foslie considered the bisporangia found by
Stromfelt to be immature tetrasporangia.
Foslie (1895) established the species Lithothamnion strom
feltii (with two forms, f. macrospora and f. tenuissima) based
on Lithothamnion tenue Rosenvinge (1893). The latter species
was described upon material from Greenland and was char
acterized by very large conceptacles and sporangia. Further
more both bi- and tetrasporangia were observed. Foslie (1895)
listed L. laeve Stromfelt and L. lenormandii f. laeve (Strom
felt) Foslie as synonyms to L. stromfeltii, and in contrast to
his 1891 paper, now regarded L. laeve Stromfelt to be a spe
cies separate from L. lenormandii. He considered L. laeve
Strornfelt to be identical to L. stromfeltii but admitted that he
was not able to prove the conspecificity because he could not
locate Stromfelt's type material. For the same reason he would
not adopt the epithet laeve.
By establishing the new name Lithothamnion stromfeltii,
Foslie (1895) created a superfluous name because Litho
thamnion tenue Rosenvinge (1893) was legitimate. Foslie's

472

Phycologia, Vol. 35 (5), 1996

'/I. M
n;f.l';'iv

"tV, K1:..
LU'..f:'11

1cm

4a
.,.m,

J("ut.'f. Jor".

f("lotu:ffw,o.manda.. ,

4cm

3cm

Fig. 2. Lectotype (LD) material of Melobesia lenormandii Areschoug in

1. Agardh.

Fig. 3. Lectotype (C) of Lithothamnion tenue Rosenvinge.

Figs 4a, b. Holotype (UPS) of Lithothamnion foecundum KjeJlman.
Fig. 4a. The note: 'New Siberian Islands Arctic USSR' is written and enclosed by Y.M. Chamberlain (Y.M. Chamberlain, personal commu
nication).
Fig. 4b. Holotype.

(1895) binomial is illegitimate according to Art. 52.1 of the

ICBN (Greuter 1994). However, Foslie (1895) considered L.
tenue Rosenvinge ( \ 893) to be a homonym of Lithothamnion
tenue (Kjellman) Foslie (1895), although he had created the
homonym himself by moving Lithophyllum tenue Kjellman
(1889) to the genus Lithothamnion.
Even though Foslie (in Rosenvinge 1898) still had not seen
Stromfelt's type material, he now felt sufficiently convinced
of the conspecificity of L. laeve Stromfelt and L. tenue Ro
senvinge to reapply the epithet laeve to the taxon. However,
Lithothamnion laeve Foslie in Rosenvinge (1898) (correctly
cited, as Stromfelt's binomial is illegitimate) is also a super-

fluous name for L. tenue Rosenvinge (\ 893) and hence ille

gitimate.
Adey (1966) removed Lithothamnion laeve Foslie in Ro
senvinge from the genus Lithothamnion Heydrich (1897) be
cause of disagreement in the epithallial cells and the meristem
cells with the branching forms of Lithothamnion. The species
was placed in a new genus Leptophytum based on Stromfelt's
Lithophyllum laeve with the binomial Leptophytum laeve Adey
as a result. As synonyms, Adey (1966) listed Lithothamnion
laeve Foslie in Rosenvinge, Lithophyullum laeve Stromfelt,
Lithophyllum lenormandii f. laeve Foslie, and Lithothamnion
stromfeltii f. tenuissima Foslie, none of which is legitimate.

Diiwel and Wegeberg: Typification and status of Leptophytum

473

Table 1. The history of Stromfelt's taxon. The second column lists the nomenclaturally correct citation of the binomial. Note how the concept
of the species changes during time; the number of spores per sporangium changes and the length of the sporangia increases

Stromfelt (1886)
Foslie (1890)
Foslie (1891)

Rosenvinge (1893)
Foslie (1895)
Foslie in Rosenvinge (1898)

Foslie (1905)
Rosenvinge (1910)
Rosenvinge (1917)
Adey (1966)
Chamberlain (1990)

Lithophyllum laeve
Stromfelt nom. illeg.
as above
Lithophyllum lenormandii
(Areschoug in J. Agardh) Rosanoff f.
laeve Foslie nom. illeg.
Lithothamnion tenue
Rosenvinge
Lithothamnion stromfeltii Foslie f.
tenuissima Foslie nom. illeg.
Lithothamnion laeve
Foslie in Rosenvinge
nom. illeg.
as above
as above
as above
Leptophytum laeve Adey
as above

Leptophytum laeve Adey is based on an illegitimate bino

mial, which therefore cannot serve as basionym. The new
combination established in 1966 by Adey is therefore to be
treated as a nomen novum, and the date of the species is thus
1966 (P.c. Silva, personal communication). The type of the
species and genus is, however, still Stromfelt's type material
of Lithophyllum laeve.

Epitypification
The holotype material of L laeve Stromfelt is a slide (Fig. l a)
annotated with the collection data (Eyrarbakki) and H.F.G.
Stromfelt's initials (H.S.) and located at the Swedish Museum
of Natural History, Stockholm (S). Unfortunately, all that can
be seen are a few bisporangia (Fig. Ib) and the pore plate of
a multiporate asexual conceptacle (Fig. Ic). The size of the
sporangia is 140-150 f..lm, which is in accordance with Strom
felt's (1886) description (125-160 f..lm).
Because this slide is the only original material and as it
does not show any taxonomic characteristics except sporangia

Length of sporangia
(j,Lm)

Spores per
sporangium

Binomial

Reference

125-160

4
4

150-210 (450)

2 or 4

200-660

150-200

200-600
240-360
126-129
220 (Fig. 36)
140-208

2 or 4
2
2
2 or 4
2

size, it is possible to select an epitype according to Art. 9.7.

in the ICBN (Greuter 1994).
We collected material at Eyrarbakki which agreed with
Stromfelt's (1886) description of morphology, i.e. , thin crusts
strongly adherent to a substrate of small lava stones. The
length of the sporangia of our specimens was 50-130 f..lm ( I I I
sporangia measured), rather smaller than the length recorded
by Stromfelt. In our resin-embedded material we observed
only one sporangium of similar size (130 f..lm). Accordingly,
we examined several sporangia of the Eyrarb akkimaterial
during the decalcification, staining, and dehydration processes.
After squashing a bisporangial conceptacle in hydrochloric
acid on a slide, we measured the length of the sporangia. We
then completed the whole procedure of decalcification, stain
ing, and dehydration on the slide and measured the size of the
sporangia during each process. The sizes of the sporangia var
ied greatly during the procedure, being larger at the outset
than after embedding in resin (respectively, 100-190 f..lm and
50-130 f..lm). If we compare the sporangia observed initially

Table 2. Measured characters in the examined type material. Numbers of measured conceptacles/sporangia are given in parentheses

Lithophyllum
laeve
holotype

Thallus thickness (j,Lm)

Asexual conceptacle diameter (j,Lm)
Diameter of pore plates (j,Lm)
Asexual conceptacle chamber diameter (j,Lm)
Number of cells in asexual conceptacle
roof filaments
Length of sporangia (j,Lm)
Carpogonial/carposporophytic conceptacle
diameter (j,Lm)
Carpogonial/carposporophytic conceptacle
chamber diameter (j,Lm)
Male conceptacle diameter (j,Lm)
Male conceptacle chamber diameter (j,Lm)

Lithophyllum
laeve epitype
(not resinembedded)

100-265
275-380 (8)
175-265 (7)
215-290 (8)
5-6
140-150 (4)

5 0-130 (I l l )
[100-190 (25)]
395-495 (4)
180-250 (5)
380-450 (2)
c. 140 (2)

Lithothamnion
tenue
lectotype

110-200
400-500 (3)
6-7
225-270 (3)

Lithothamnion
foecundum
holotype

:51100
300 (4)
5-7

Melobesia
lenormandii
lectotype

75-200
200-285 (2)
135-175 (3)
135-200 (4)
3-5
80-95 (4)
300-350 (3)
185-225 (3)

474

Phycoiogia, Vol. 35 (5), 1996

Figs 5-10. Epitype (C) of Lithophyllum laeve Strbmfelt nom. illeg. from Eyrarbakki, Iceland.
Fig. 5. Habit. (EY920102).
Fig. 6. Section through dorsal region showing domed epithallial cells and the meristem cells beneath. Note the increasing size of the cells

cut off the meristem cells. (EY920106).

Fig. 7. Monomerous vegetative thallus. (EY920106).
Fig. 8. Thallus surface. (EY920119).
Fig. 9. Asexual conceptacle with bisporangia. Note the large, pale apical cells in the conceptacle roof (arrowhead). (EY920102).
Fig. 10. Immature asexual conceptacle containing sporangia initials (arrowhead). Note the large, pale apical cells at the surface of the

conceptacle roof. (EY920110).

during this experiment with those drawn by Stromfelt (1886,

tab. 1, figs. 11, 12), they are much more alike. The resin
embedded material showed somewhat shrunken sporangia, not
filling the sporangial cavity completely.
In summary, our material corresponds to Stromfelt's de
scription in (1) morphology and habitat, (2) size and form of
the sporangia, and (3) presence of bisporangia only. These
similarities confirm that the alga we have collected is the same
as the one Stromfelt found and described and can thus serve
as epitype.

Observations on morphology and anatomy

The following description of the epitype is based on a single
collection at Eyrarb akki
on 4 June 1992 (collection no. EY9201,
specimen nos EY920101-04, EY920106-1O, EY920112-14,
and EY920118-19) (C).
The external appearance is a thin (100-265 J,Lm) crust with
out protuberances, strongly adherent to the substrate (Fig. 5).
Conceptacles are seen as small white spots.
The thallus is pseudoparenchymatous, each filament ter-

Duwel and Wegeberg: Typification and status of Leptophytum

11

475

12

";;::.,:...
, -,;0:'
."1.

,>5\

....

","

Figs 11-16. Epitype (C) of Lilhophyllum laeve Stromfelt nom. illeg. from Eyrarbakki, Iceland. Uniporate female conceptacle.
Fig. 11. Section through conceptacle after fertilization. Note the deeply stained, flask-shaped terminal cells of the filaments lining the pore
canal (arrowhead). The crowded small cells in the conceptacle chamber are the apical cells of filaments constructing the conceptacle (confirmed
by serial sections). (EY9 20107).
Fig. 12. Early stages of carpogonial branches. (EY920102).
Fig. 13. Carpogonium with a trichogyne. (EY92010 2).
Fig. 14. Young carposporophyte with fusion cell (arrowhead). (EY920 l l4).
Fig. 15. Young carposporophyte with laterally extended fusion cell (arrowheads). (EY9201l 4).
Fig. 16. Carposporangium. (EY92010 2).

minated by a single rounded to domed epithallial cell (Fig. 6)

above a meristem cell that cuts off cells ventrally without prior
elongation. After division, the daughter cell becomes larger.
This phenomenon results in cell size increasing inwardly in
the thallus (Fig. 6). The thallus consists of few ventral fila
ments subparallel to the substrate, curving upward and becom
ing perpendicular to the substrate (monomerous) (Fig. 7). The
thallus surface appears as cells with relatively thick calcareous
walls surrounding concavities and can be more or less covered
by a veil (Fig. 8). Cell fusions are present but uncommon.
Asexual conceptacles are multiporate and 275-380 f.Lm in
diameter. Pore plates are 175-265 f.Lm in diameter and the
conceptacle chambers are 215-290 f.Lm. Bisporangia (Fig. 9)
have apical pore plugs and measure 50-130 f.Lm (excluding
the pore plugs) when embedded in resin. The conceptacles are

raised and the conceptacle roofs consist of vertically oriented

5- to 6-celled filaments with a characteristic large, pale apical
cell (Fig. 9). These cells can be sloughed off, leaving cup
shaped remnants on the conceptacle roofs. In immature asex
ual conceptacles (Fig. 10), sporangial initials appear as long,
deeply stained cells in cavities with a pore, and the filaments
of the conceptacle roof have characteristic apical cells.
Carpogonial conceptacles are uniporate with an external
basal diameter of 395-495 f.Lm and an inner diameter of 180245 f.Lm. The terminal cells of the filaments that terminate
along the pore canal are flask-shaped and deeply stained (Fig.
11). Carpogonial branches arise from the conceptacle floor
(Fig. 12), each consisting of 4 cells: a basal cell, a supporting
cell, a hypogynous cell, and the carpogonium (Fig. 13). From
the carpogonium, a long trichogyne (:'S 60 f.Lm) protrudes (Fig.

476

Phycologia, Vol. 35 (5), 1996

Figs 17-21. Epitype (C) of Lithophyllum laeve Strbmfelt nom. illeg. from Eyrarbakki, Iceland. Uniporate male conceptacles.
Fig. 17. Section through conceptacle with dendroid spermatangial systems. Note the deeply stained, flask-shaped terminal cells of the filaments
lining the pore canal (arrowhead). (EY920103).
Fig. 18. Early stages of spermatangial filaments covering the entire conceptacle floor. (EY920103).
Fig. 19. Spermatangial filaments with elongated apical cell (arrowhead). (EY920113).
Fig. 20. Spermatangial filaments showing the early development of the dendroid system (arrowheads). (EY920113).
Fig. 21. Different stages of spermatangial development within the same conceptacle. (EY920103).

13). Upon fertilization, the trichogynes decay and a fusion cell

can be observed (Fig. 14). Subsequently the fusion(s?) ex
tends laterally (Fig. 15), but it could not be determined wheth
er gonimoblast filaments develop peripherally or all across the
floor. The difficulties in obtaining this evidence indicate that
the carposporophyte is very fragile. Carposporangia are ulti
mately formed (Fig. 16).
Male conceptacles are uniporate with an external basal di
ameter of 380-450 /-Lm and an inner chamber diameter of
approximately 140 /-Lm. As in the female conceptacle, the ter
minal cells of filaments forming the conceptacle roof are flask
shaped and deeply stained along the pore canal (Fig. 17).
Spermatangial filaments develop on the conceptacle floor only
(Figs 17, 18). The early stages resemble the carpogonial
branches in consisting of four cells, the apical one being elon
gated (Fig. 19). Cells of the spermatangial filaments divide
(Fig. 20) and develop dendroid systems (Fig. 21). Different
stages of spermatangial development are present within one
conceptacle (Fig. 21).

When Foslie (1895) revised his classification of the corallines,

he regarded Lithophyllum as a subgenus of Lithothamnion,
thereby creating the combination Lithothamnion lenormandii
(Areschoug in J. Agardh) Foslie. In the same paper, Foslie
erected a new species, Lithothamnion squamulosum, but later
(Foslie 1905) regarded it as a form of Lithothamnion lenor
mandii and stated that this form, L. lenormandii f. squamulosa
(Foslie) Foslie, might represent senescent specimens of L. len
ormandii f. typica. Heydrich (1911) established the genus
Squamolithon, characterized principally by the development
of the gonimoblasts, with Lithothamnion lenormandii as the
type. However, this combination was not recognized by con
temporary or later corallinologists. Adey (1966) transferred
Lithothamnion lenormandii to Phymatolithon, thus creating
the presently used combination Phymatolithon lenormandii
(Areschoug in J. Agardh) Adey.
Typification

Melobesia lenormandii Areschoug in J. Agardh

LECTOTYPE (LD): Melobesia lenormandii, Arromanches,

Calvados, France. Specimen 50674 (I) designated by Woelk
erling (1988, in herb.) (Fig. 2).

Historical background

Observations on morphology and anatomy

Melobesia lenormandii was described by Areschoug in J.

Agardh (1852). Rosanoff (1866) made the new combination
Lithophyllum lenormandii (Areschoug in J. Agardh) Rosanoff.

The following description is based on specimens B and C

from the lectotype collection.
The external appearance is a thin crust (75-200 /-Lm) with-

Diiwel and Wegeberg: Typification and status of Leptophytum

477

22

1cm

23

24

100 \-1m

25

28

100 f.Jm

Figs 22-28. Lectotype (LD) of Melobesia lenormandii (Areschoug in J. Agardh) Rosanoff.

Fig. 22. Habit.
Fig. 23. Section through dorsal region showing epithallial cells and the meristem cells.
Fig. 24. Monomerous vegetative thallus.
Fig. 25. Asexual conceptacle with bisporangia. The large size of the conceptacJe is caused by fusion of two conceptacJes.
Fig. 26. Section through a carpogonial conceptacle. Note carpogonial branches terminated by trichogynes and the elongated cells terminating
the roof filaments in the pore region.
Figs 27, 28. Carposporophytic conceptacle with gonimoblasts issued from fusion cells across the floor of the conceptacJe chamber.

out protuberances (Fig. 22) possessing raised uni- and multi

porate conceptacles. Filaments of the pseudoparenchymatous
thallus are terminated by epithallial cells that are flattened,
rounded, or domed (Fig. 23), and beneath the epithallial cells
are meristem cells. These cut off cells which elongate succes
sively (Fig. 23). The thallus is monomerous (Fig. 24) and cell
fusions are observed.
Asexual conceptac\es (Fig. 25) are multiporate and 200285 /-Lm in diameter. Pore plates are 135-175 (370) /-Lm in
diameter and the conceptacle chambers 135-200 (350) /-Lm.
The extreme values of conceptac\e size are caused by fusion
of two conceptacles (Fig. 25). Bisporangia (Fig. 25) have api
cal pore plugs and measure 80-95 /-Lm (excluding pore plugs)
when embedded in resin. The filaments of the conceptac\e

roofs consist of 3-5 cells. Immature asexual conceptacles

were not observed.
Carpogonial conceptac\es are uniporate (Fig. 26). One con
ceptacle measured 350 /-Lm in diameter, the conceptac\e cham
ber being 185 /-Lm; carposporophytic conceptac\es measured
300 /-Lm externally and 200-225 /-Lm internally. The filaments
of the conceptac\e roofs terminate by elongated cells in the
pore canal region. From the conceptac\e floor arise carpogo
nial branches terminated by trichogynes (Fig. 26). Several fu
sion cells with various degrees of extension (Figs 27, 28) are
observed on the floor and walls of the chamber. From the
fusion cells, gonimoblasts arise (Fig. 27) and spread across
the chamber floor (Fig. 28). Male conceptac\es were not ob
served.

478

Phycologia, Vol. 35 (5), 1996

Lithothamnion tenue Rosenvinge

Historical background
The history of L. tenue is treated in the discussion of Litho
phyllum laeve (see also Table I).
Lectotypification
L.K. Rosenvinge did not designate a specimen or collection
as type specimen for his taxon Lithothamnion tenue, but based
his description on collections from Greenland made by him
self and several other persons. Two numbered specimens of
L. tenue Rosenvinge are located at C. One specimen (herbar
ium no. 879) corresponds to two slides in Rosenvinge's slide
collection (Fig. 3). The slides are annotated Lithothamnion
tenue by Rosenvinge and fully supplied with collection data
to match the data in his description. The other specimen of L.
tenue (no. 331) is labelled, and supplied with collection data.
However, this crust does not possess any conceptacles, only
scars of shed ones. Specimen no. 879 is therefore selected as
lectotype.
LECTOTYPE (now designated) (C): Lithothamnion tenue, Hol
steinsborg, Greenland, 12 fathoms. 20 June 1890, collected by
N. Hartz, no. 879 (Fig. 3).
Observations on morphology and anatomy
The external appearance of the lectotype specimen is a thin
crust (1 lO-200 fLm) without protuberances. Conceptacles ap
pear as small white spots (Fig. 29). The epithallial cells ter
minating the pseudoparenchymatous filaments are flattened to
rounded (Fig. 30). Subepithallial meristem cells cut off cells
without prior elongation. After division, the daughter cells be
come successively larger (Fig. 30). The thallus consists of few
ventral filaments subparallel to the substrate, which curve up
wards and become perpendicular to the substrate (monomer
ous) (Fig. 31). Cell fusions are present (Fig. 31).
Asexual conceptacles are multiporate (Fig. 32). The con
ceptacles are raised above the thallus surface. The pore plates
measure 400-500 fLm in diameter. The roof consists of fila
ments of 6-7 cells. The filaments next to the pores consist of
distinctive cells, being larger and more deeply stained than
the other cells in the conceptac\e roof (Fig. 32). Bisporangia
(Fig. 33) have apical pore plugs and measure 225-270 fLm
excl. pore plugs. Gametophytic conceptacles are not present
in the specimen.

Lithothamnion foecundum Kj ellman

Historical background
In 1883 Kjellman separated Lithothamnion foecundum from
the heterogenous taxon Lithothamnion polymorphum (Linnae
us) Areschoug in J. Agardh (1852) [see Woelkerling & Irvine
(1986) for disentanglement of nomenclatural and taxonomical
problems of Lithothamnion polymorphum]. Adey (1966)
transferred Lithothamnion foecundum Kjellman (1883) into
Leptophytum, creating the new combination Leptophytumfoe
cundum (Kjellman) Adey.

Typification
The original collection (UPS) of L. foecundum consists of two
specimens and five slides marked Lithothamnion foecundum
Kjellman. One of the specimens is marked 'orig. ex.' (Fig.
4a) and must be regarded as the holotype. This is supported
by the appearance of the specimen (Fig. 4b) that corresponds
with the drawing of L. foecundum in the original description
(Kjellman 1883, pI. 5, fig. II). The holotype specimen is not
supplied with any collection data. However, the other speci
men is fully supplied with collection data: 'Sibiriska Ishafvet
nara Cap Taimur, 12 august 1878, F.R. Kjellman'. The five
slides are annotated 'Kariska Hafvet' (Kara Sea).
HOLOTYPE (UPS): Lithothamnion foecundum. Labelled 'orig.
ex.'.
Observations on morphology and anatomy
The external appearance of the holotype specimen is a crust
without protuberances covered with multiporate conceptacles.
Filaments of the pseudoparenchymatous thallus are terminated
by flattened to rounded epithallial cells (Fig. 34). Meristem
cells are located beneath the epithallial cells and cut off cells
without prior elongation. Newly formed cells subsequently
elongate and become squarish (Fig. 34). The thallus is mon
omerous (Fig. 35) and relatively thick (up to l lO0 fLm). Cell
fusions are present (Fig. 34).
Asexual conceptacles are multiporate (Fig. 36) and raised
above the thallus surface. The pore plates measure 300 fLm in
diameter. The roof consists of vertical filaments of 5-7 cells,
with the pores surrounded by characteristic larger and deeply
stained cells (Fig. 37) that may be kidney-shaped. Old con
ceptacles become buried in the thallus and filled with second
arily formed cells (Fig. 38). Gametophytic conceptacles are
not present in the holotype specimen.

TAXONOMIC CONCLUSIONS
Status and disposition of the genus Leptophytum
As the genus Leptophytum is based on Lithophyllum laeve
Stromfelt and as type material of this species was unknown
until very recently, the status of the genus has been uncertain
(Woelkerling 1988; Chamberlain 1990). Therefore, the delim
itation between Leptophytum and Phymatolithon has always
been problematic for modern corallinologists (Adey 1966;
Adey 1970; Lebednik 1978; Woelkerling & Irvine 1986;
Woelkerling 1988; Chamberlain 1990; Chamberlain & Irvine
1994; Chamberlain & Keats 1994). The characters considered
to separate Leptophytum and Phymatolithon will be discussed
below and are summarized in Table 3 as represented by se
lected species.
Elaboration of the thallus surface has been proposed by
Chamberlain (1990) as useful in delimiting the two genera.
She distinguished between a Phymatolithon-type and a Lep
tophytum-type, respectively consisting of thickened calcareous
ridges surrounding central concavities and flat cell surfaces,
each with a minute central hole and surrounded by a thin
calcareous wall. She stated that the genus Phymatolithon has
a thallus surface of the Phymatolithon-type only while Lep
tophytum has both the Leptophytum- and Phymatolithon-type.

Diiwel and Wegeberg: Typification and status of Leptophytum

479

32

50l-lm

Figs 29-33. Lectotype (C) of Lithothamnion tenue Rosenvinge. Morphology, vegetative anatomy, and multiporate asexual conceptacle.
Fig. 29. Habit. Asexual conceptacles are seen as small white spots.
Fig. 30. Section through dorsal region showing flattened to rounded epithaUial cells and the underlying meristem cells. Note the inwardly

increasing size of the cells cut off the meristem cells.

Fig. 31. Monomerous vegetative thallus. Note cell fusions (arrowheads).
Fig. 32. Asexual conceptacle roof with deeply stained, large cells (arrowhead) surrounding the pores with pore plugs.
Fig. 33. Asexual conceptacle containing bisporangia with apical pore plugs (arrowheads).

However, Wilks & Woelkerling (1994) have shown that the

type species of Phymatolithon (P. calcareum) possesses both
thallus surface types and they explain the differences as ar
tefacts due to drying. We therefore do not recognize this char
acter as a diagnostic feature.
Adey (1966) observed deeply stained cells lining the pore
canals of the multiporate conceptacles of Leptophytum and
included this observation (as 'pore cells') in the diagnosis of
the genus. When erected, the genus accommodated two spe
cies, L. laeve and L. foecundum, both possessing the pore
cells. However, this character is now regarded as variable
within Leptophytum according to species (Chamberlain 1990;
Chamberlain & Irvine 1994). A similar situation exists in the
genus Mesophyllum, wherein differences in the cells lining the
pores are used in delimiting species (Woelkerling & Harvey
1992).
The location of conceptacle primordia has also been con
sidered as diagnostic for the two genera (Adey 1970; Cham
berlain 1990, table 2). Wilks & Woelkerling (1994, p. 200)
have carefully considered this character and did not regard it
as useful in delimiting genera. They found that the location
of conceptacle initiation depends on a subjective interpretation
of Adey's (1966) description of the conceptacle primordia as
developing a few cells down in the vegetative thallus of Lep
tophytum or, as Adey (1964) described for Phymatolithon, de
veloping from cells sunken in the vegetative thallus.

Lebednik (1978) discussed the development of male con

ceptacles in several melobesioid genera and referred to Adey
(1966), who observed elongated cells in the spermatangial
conceptacles of Leptophytum laeve. Lebednik (1978) regarded
these cells as corresponding to the protective cells he observed
in two species of Mesophyllum. He ascribed this character to
the genus Leptophytum and stated that no such cells appear
to be formed in Phymatolithon. However, other authors have
not confirmed the presence of these cells in Leptophytum, and
at present it is not possible to demonstrate the importance of
this character at the generic level.
Lebednik (1978), Chamberlain (1990), Chamberlain & Ir
vine (1994), and Chamberlain & Keats (1994) further sug
gested that the structure of spermatangial conceptacle contents
distinguishes Leptophytum from Phymatolithon. They found
that the spermatangial systems of Leptophytum are simple in
the periphery of the conceptacle and somewhat elaborate in
the centre, whereas Phymatolithon has dendroid spermatangial
systems across the entire floor of the conceptacle. In our ma
terial of the epitype of Lithophyllum laeve, we observed both
types of elaboration of the spermatangial systems (respective
ly, Figs 18, 21, and 17), which invalidates the character. We
believe that the different elaborations of spermatangial sys
tems represent different stages in development, in that Fig. 17
shows a mature male conceptacle and Figs 18 and 21 show
earlier stages of development, Fig. 18 being the youngest.

480

Phycoiogia, Vol. 35 (5), 1996

34

200iJm

40 JJm

Figs 34-38. Holotype (UPS) of Lithothamnion foecundum Kjellman.

Fig. 34. Section through dorsal region showing flattened epithallial cells and the meristem cells. Note the cell fusions (arrowhead).
Fig. 35. Monomerous vegetative thallus.
Fig. 36. Roof of multiporate asexual conceptacle.
Fig. 37. Asexual conceptacle roof with pores lined by characteristic, deeply stained, larger cells (arrowhead).
Fig. 38. Buried, old conceptacles become filled with secondarily formed cells.

Table 3. Occurrence of proposed Phymatolithon and Leptophytum generic characters.

+, present; 7, absent; ND, no data

Phymatolithon
calcareum neo
type (Woelker
Phymatolithon Leptophytum
Litho
Litho
ling & Irvine
laeve
repandum
1986; Wilks & Phymatolithon
thamnion thamnion
(Adey 1966;
(Wilks &
tenue
Chamberlain Lithophyllum foecundum
Woelkerling
Woelkerling
lenormandii
laeve epitype holotype lectotype
1990)
1994)
lectotype
1994)
Leptophytum-type surface cells present
Phymatolithon-type surface cells present
Tetra-Ibisporangial conceptacles initiated
adventitiously
Pores of asexual conceptacles lined with
distinctive cells
Gonimoblast filaments peripheral in conceptacle
Gonimoblast filaments all across the floor
Protective cells present in immature male
conceptacles
Dendroid spermatangial systems all over
conceptacle floor
Sperrnatangial systems dendroid in center of
conceptacle, while simple in periphery

'Wilks & Woelkerling 1994, fig. 3C.

Chamberlain 1990.

+
+

ND
ND
ND

+,
+'
+

+
+
+

+
+
ND

+
+

NO
ND
ND

+
ND

ND

ND

ND

ND

ND
ND
ND

+/7

ND

ND

+/7
ND

ND
ND

ND
ND

ND

ND

ND

ND

Dendroid and
simple systems
intermixed

+2
ND
ND

Diiwel and Wegeberg: Typification and status of Leptophytum

Adey (1970) stated that in Phymatolithon, gonimoblasts are

'vertical from fusion cell' in contrast to Leptophytum, which
has gonimoblasts 'lateral from fusion cell' , i.e., gonimoblasts
developed on the entire conceptacle floor and gonimoblasts
confined to the periphery of the conceptacle chamber, respec
tively. Wilks & Woelkerling (1994) observed gonimoblast fil
aments and carposporangia developing both across the floor
and confined to the periphery of the conceptacle chamber
within Phymatolithon repandum (Foslie) Wilks et Woelker
ling. This character must therefore also be considered invalid.
On the basis of the above discussion of characters used in
distinguishing Leptophytum from Phymatolithon, we can only
conclude that the two genera cannot be considered distinct.
The same conclusion was reached by Wilks & Woelkerling
(1994) in their account of the status of Leptophytum and is
here confirmed by our study of epitype material of Litho
phyllum laeve Stromfelt. The material corresponds to Woelk
erling's (1988) description of the genus Phymatolithon and to
his list of diagnostic features in the following ways: (I) it is
nontaeniform; (2) haustoria are not present; (3) the thallus
possesses a ventral core of noncoaxial filaments; (4) the epi
thallial cells are rounded (or domed) but not flared; and (5)
the meristem cells are as short as or shorter than their im
mediate inward derivatives. Woelkerling's (1988) sixth char
acter diagnostic for Phymatolithon concerns the location of
the asexual conceptacle primordia, which arise adventitiously
within the thallus. We do not have evidence of this character
in the epitype of L. laeve. It delimits Phymatolithon from
Clathromorphum as the latter genus develops asexual concep
tacles from groups of subepithallial initials (Woelkerling
1988). However, these genera also differ by the meristem
cells, which in Clathromorphum are as long as or longer than
their immediate inward derivatives (Woelkerling 1988). This
is indeed not the case in the epitype of Lithophyllum laeve,
and consequently we conclude that the epitype of Lithophyl
lum laeve belongs to Phymatolithon.
Stromfelt (1886) delimited Lithophyllum laeve from Litho
phyllum lenormandii (Areschoug in J. Agardh) Rosanoff by
the presence of only bisporangia and by larger sporangial size.
However, bisporangia are observed in the type material of
Phymatolithon lenormandii and Stromfelt (1886), Suneson
(1943), Adey (1966), and Chamberlain & Irvine (1994) have
also recorded bisporangia in other specimens of P. lenorman
dii. Furthermore, the length of the bisporangia of the L. laeve
epitype (50-130 f.Lm) is commensurate with that measured in
the type material of P. lenormandii (80-95 f.Lm) and reported
for other specimens of P. lenormandii (Foslie 1895; Rosen
vinge 1917; Suneson 1943; Chamberlain & Irvine 1994).
In the material from Eyrarbakki we have observed char
acteristic, large, pale apical cells in the conceptacle roof fila
ments. The same phenomenon may be observed in illustra
tions of immature asexual conceptacles of Phymatolithon len
ormandii in Chamberlain & Irvine (1994, fig. 114D) and in
Suneson (1943, fig. I as Lithothamnion lenormandii). The
cells have not been observed in the type material of P. len
ormandii, but their absence may be due to different methods
of preservation or age of the conceptacles. This seems prob
able as the cells are readily sloughed off.
Phymatolithon lenormandii is described with gonimoblasts
developed over the entire floor of the conceptacle (Rosenvinge
1917; Suneson 1943; Adey 1966; Chamberlain & Irvine

481

1994), which we also observed in the type material. However,

we were unable to determine whether gonimoblasts were pe
ripheral or widespread in the epitype of Lithophyllum laeve.
Since both elaborations of the carposporophyte are observed
within Phymatolithon repandum (Wilks & Woelkerling 1994),
this character is invalid at even the species level.
The above comparison of the epitype of Lithophyllum laeve
with the type material of Phymatolithon lenormandii shows
that Stromfelt (1886) misidentified a specimen of P. lenor
mandii. We did not observe any specimens at the type locality
or later in our examination of our extensive Eyrarbakki col
lection that can be referred to Adey's (1966) <?oncept of Lep
tophytum laeve, which we do recognize as a different taxon
from P. lenormandii (see below).
As the type of Leptophytum laeve Adey is now shown to
be Phymatolithon lenormandii, L. laeve Adey is synonymous
with P. lenormandii (Areschoug in J. Agardh) Adey and Lep
tophytum Adey is synonymous with Phymatolithon Foslie.
Disposition of Adey's (1966) concept of

Leptophytum laeve
As the type of Leptophytum laeve is actually Phymatolithon
lenormandii, Adey (1966) has erroneously used Stromfelt's
type as nomenclatural type for his taxon. Adey's (1966) con

cept of L. laeve is taxonomically different from P. lenorman

dii by distinctive cells lining the pore canals of asexual con
ceptacles.

In order to identify Adey's taxon we examined the closely

related species Lithothamnion joecundum Kjellman and Lith
othamnion tenue Rosenvinge, which has been synonymized
with Lithothamnion laeve Foslie in Rosenvinge. Adey (1966)
listed the latter species as one of the synonyms of Leptophy
tum laeve.
The holotype of Lithothamnion joecundum corresponds to
Adey's concept of Leptophytum laeve by presence of the char
acteristic pore cells and the elaboration of epithallial and mer
istem cells. However, L. joecundum differs by having a rela
tively thick thallus, generally smaller pore plates, and buried
old conceptacles. The last character has never been reported
for L. laeve.
In contrast, the lectotype of Lithothamnion tenue Rosen
vinge fully agrees with Adey's (1966) description of Lepto
phytum laeve. The lectotype possesses the characteristic large,
deeply stained cells surrounding the pores in the asexual con
ceptacle roofs as described by Adey (1966, figs 36, 62). Bi
sporangia in Adey's material (Adey 1966, figs 36, 62) are
225-275 f.Lm long, in agreement with the length of bisporangia
of the L. tenue lectotype (225-270 f.Lm). The diameter of the
pore plates given by Adey (1966) is 270-480 f.Lm, and the
pore plates of the lectotype are 400-500 f.Lm. In vegetative
anatomy, Adey's (1966, figs 35, 53) illustrations of Leptophy
tum laeve are identical with the L. tenue lectotype (Figs 31,
30). In both cases the elongation of cells cut off by meristem
cells is postdivisional, the thallus is monomerous, thin (Adey
1966; 50-320 f.Lm, the lectotype: 110-200 f.Lm), and cell fu
sions are present. Consequently, Adey's (1966) concept of
Leptophytum laeve Adey is in full accordance with the lec
totype of Lithothamnion tenue Rosenvinge and is thereby
identified.

482

Phycologia, Vol. 35 (5), 1 996

Table 4. Characters diagnostic of Phymatolithon foecundum, P. len

ormandii, and P. tenue based on the type material

P.
P.
foecundum lenormandii

Thallus thickness (/-Lm)

Diameter of pore plates (/-Lm)
Large, darkly stained cells lining the pore canals in asexual conceptacles
Buried old conceptacles

:5 1100
300

Present
Present

P.
tenue

75- 200
135-175

110- 200
400-500

Absent
Absent

Present
Absent

Pedersen, for guidance. Many thanks are due to the late Tyge
Christensen and to Paul C. Silva for their support and advice
in disentangling the nomenclature. Our thanks are also due to
an anonymous referee and to Yvonne M. Chamberlain who
provided constructive and valuable comments which im
proved the manuscript. We also wish to express our thanks to
the following funds for financial support: Dansk-Islandsk
Fond, Ing. Svend G. Fiedler og hustrus legat, Knud H0jgaards
Fond, Dansk Botanisk Forening, Mag. art. Marcus Lorenzens
Legat, Nordisk Kollegium for Marinbiologi and Japetus
Steenstrups Legat.

Status and disposition of

Lithothamnion tenue and L. foecundum

Lithothamnion tenue Rosenvinge does not belong to the genus
Lithothamnion, because we have not observed flared epithal
lial cells and elongated meristematic cells which are charac
teristic for the genus according to Woelkerling ( 1 988). Instead
we have observed vegetative characters as described for Phy
matolithon (Woelkerling 1 988): flattened to rounded epithal
lial cells and postdivisional elongation of cells cut off by mer
istematic cells (Fig. 30). We therefore propose the following
new combination.

Phymatolithon tenue (Rosenvinge) comb. nov.

BASIONYM: Lithothamnion tenue Rosenvinge ( 1 893), p. 778,

figs 4-7.
NOMENCLATURAL SYNONYMS: Lithophyllum lenormandii
(Areschoug in J. Agardh) Rosanoff f. laeve Foslie nom. iIIeg.
( 1 89 1 ). Lithothamnion stromfeltii Foslie nom. iIIeg. ( 1 895).
Lithothamnion laeve Foslie in Rosenvinge nom. iIIeg. ( 1 898).
LECTOTYPE (C): No. 879, Holsteinsborg, Greenland, 20 June
1 890, collected by N. Hartz, two slides by L.K. Rosenvinge
and 1 8 slides by L. Diiwel & S. Wegeberg.

The holotype specimen of Lithothamnion foecundum Kjell

man possesses features characteristic of the genus Phymato
lithon Woelkerling ( 1 988) as described for P. tenue. Therefore
we hereby propose the following new combination.

Phymatolithon foecundum (Kj ellman) comb. nov.

BASIONYM: Lithothamnion foecundum Kjellman (1883), p.
99, pI. 5, figs 1 1 - 1 9.
NOMENCLATURAL SYNONYM: Leptophytum foecundum (Kjell
man) Adey (1 966).

The characters diagnostic of and separating the three spe

cies in question, viz. Phymatolithon foecundum, P. lenorman
dii, and P. tenue, are summarized in Table 4. A study of the
remaining species in the former Leptophytum Adey will reveal
whether they all belong to the genus Phymatolithon Foslie or
whether a separate genus has to be established.

ACKNOWLEDGEMENTS
We wish to thank William J. Woelkerling for inspiration and
invaluable help during the whole project, Karl Gunnarson for
assistance during our stay in Iceland, and our supervisors from
University of Copenhagen, Aase Kristiansen and Poul M011er

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Accepted 3 June 1 996