Phycologia (1987) Volume 26 (I), 9-16
gen. nov. and a reappraisal of the
Phyllariaceae Tilden 1935 (Laminariales, Phaeophyceae)
Phyllariopsis
ERIC C. HENRY' AND G. ROBIN SOUTH2*
, Department of Biology, Memorial University of Newfoundland,
St John's, Newfoundland AlB 3X9, Canada
2
Huntsman Marine Laboratory, St Andrews, New Brunswick EOG 2XO , Canada
E.e. HENRY AND G.R . SOUTH. 1987. Phy/lariopsis gen. nov. and a reappraisal
Tilden 1935 (Laminariales, Phaeophyceae). Phyc% gia 26: 9-16.
of the Phyllariaceae
Phy/lariopsis gen. nov . and Saccorhiza de la Pylaie make up the family Phyllariaceae Tilden 1935,
distinguished from other families in the Laminariales by the occurrence �f thlck-walled c�nductIng
.
elements (solenocysts) in the medulla. Several other characters of the famIly mdlcate pnmltlveness:
zoospores possess eyespots, holdfasts lack branched haptera, sorus paraphyses lack thick�ned tips,
the sexual pheromone is not lamoxirene, gametophytes of so�e species are not sexually dimorphic,
and hair tufts occur on young laminae. Phy/lariopsis gen. nov. IS proposed, With the new combmatlOns
Phy/lariopsis brevipes = Phy/laria reniformis (Lamouroux) Rostafinski ex Bornet and Phy/lariopsis
purpurascens = Phy/laria purpurascens (C. Agardh) Rostafinski ex Bornet.
INTRODUCTION
The family Phyllariaceae was established by Til­
den (1935) to include two genera of Laminari­
ales, Saccorhiza de la Pylaie and Phyllaria (Le
Jolis) Rostafinski. Although previous workers
(Rostafinski 1877; Gobi 1878; Setchell & Gard­
ner 1925; Feldmann 1934) had recognized that
these genera constitute a distinct group within
the Laminariales, most subsequent authors have
not recognized the Phyllariaceae and have treat­
ed Saccorhiza and Phyllaria as members of the
Laminariaceae.
A recent ultrastructural study found that an
eyespot was lacking in zoospores of all 17 species
of 14 genera of Lessoniaceae, Alariaceae and
Laminariaceae that were examined (Henry & Cole
1982). The light microscopical literature on this
topic has been equivocal (Fritsch 1945), but an
incidental report of an electron microscopical
observation of a zoospore eyespot in Saccorhiza
polyschides (Lightfoot) Batters (Evans 1966) sug­
gested that this and related species might possess
other distinctive characters, prompting us to re­
examine these species as well as earlier literature
describing them. This paper presents a review of
the literature and recent findings supporting the
* To whom reprint requests should be addressed.
recognition of a distinct family Phyllariaceae; no­
menclatural revision of one genus is proposed.
TAXA INCLUDED IN THE
PHYLLARIACEAE
The Phyllariaceae contains four species (Table
1), whose distributions are confined to the North
Atlantic. Saccorhiza polyschides (Lightfoot) Bat­
ters occurs from southern Norway (Svendsen
1962) to the Tropic of Cancer in west Africa
(Lawson & John 1982), and in the Mediterranean
as far east as the Strait of Messina (Huv€: 1958;
Giaccone 1969). There is also one record from
the Aegean island of Chios (Bornet 1892). Sac­
eorhiza dermatodea (de la Pylaie) J. Agardh is
known in northern Europe from Novaya Zemlya
to Norway (Kjellman 1883), westward to Elles­
mere I. and south to northern Massachusetts in
North America (Taylor 1957). Phyllaria purpur­
ascens (c. Agardh) Rostafinski ex Bornet is known
from northwest Spain to Morocco and in the
Mediterranean to the Strait of Messina (Feld­
mann 1934; Giaccone 1969). Phyllaria renifor­
mis (Lamouroux) Rostafinski ex Bornet has a
similar distribution but also occurs northward
into France in the Atlantic. Both species are rare.
Saccorhiza polyschides was included in Lam9
10
Phycologia, Vol. 26 (1), 1987
inaria (as L. bulbosa) by Lamouroux (1813) when
he established this genus, and S. dermatodea, P.
purpurascens and P. reniformis were all origi­
nally described as species of Laminaria. A sep­
arate genus Saccorhiza was established for S.
polyschides by de la Pylaie (1829, as S. bulbosa).
Although he remarked upon the similarities be­
tween this species and his new species L. der­
matodea, he left the latter in Laminaria. De­
caisne (1842) proposed the genus Haligenia for
S. polyschides. Le Jolis (1855) divided Haligenia
into two sections, Saccorhiza, containing S. poly­
schides (as S. bulbosa) and several doubtful
species, and Phyllaria, including P. dermatodea
and the two species of the Mediterranean and
adjacent Atlantic, P. purpurascens and P. reni­
formis. Phyllaria was established as a separate
genus by Rostafinski in 1877 (Farr et al 1979).
Phyllaria reniformis has a confused history.
Since the time of Bornet (1892) it has been re­
ferred to as Phyllaria reniformis (Lamouroux)
Rostafinski ex Bornet. Laminaria reniformis was
established by Lamouroux's (1813) publication
of only the name and a figure, with the locality
of Cape of Good Hope. C. Agardh (1820) pro­
vided a description of L. reniformis based on
Lamouroux's figure, and he noted the distinct
differences between L. reniformis (long, branched
haptera and regularly dissected lamina) and his
L. brevipes (short, unbranched haptera and ir­
regularly dissected lamina). Lamouroux's figure
appears to be idealized, and it does not corre­
spond to any of the specimens in his collections
at Caen or Paris labelled "L. reniformis Lamx."
(none appears to be labelled in the hand of La­
mouroux). The figure does resemble the South
African Laminaria pallida Greville in J. Agardh
(1848) in regard to the form of the holdfast and
regular splitting of the lamina. Laminaria pallida
is usually a much larger species, but no scale was
given for Lamouroux's figure so the size of his
specimen is unknown. The extant Lamouroux
specimens labelled L. reniformis are all of the
Mediterranean species, and later authors have
used Lamouroux's epithet, assuming that the
Cape of Good Hope locality of L. reniformis was
the result of a labelling error (Feldmann 1934;
Hamel 1938). However, because of the lack of
concordance between the characters illustrated
in Lamouroux's figure and those of the Medi­
terranean species, the possibility must be con­
sidered that Lamouroux intended L. reniformis
to refer to a S. African species. C. Agardh's (1820)
description of L. brevipes is clear and the type,
his No. I I (Lund No. 1952; Fig. I) from Cadiz,
is a complete, fertile specimen. Therefore it seems
best to accept J. Agardh's (1848) evaluation of
L. reniformis as "species dubia", and to consider
the Mediterranean species to be typified by L.
brevipes C. Agardh. Rostafinski's "in herb."
transfer of L. reniformis Lamouroux to Phyllaria
[published by Bornet (1892)] was based on
Shousboe's collections from Morocco.
The two species of Saccorhiza are distin­
guished by significant differences: most striking
are the adult holdfasts (cf. Norton 1972) and the
gametophytes (Henry 1986). However, these
two species resemble each other more closely
than either resembles any other species. We
therefore retain S. dermatodea in Saccorhiza.
Because Saccorhiza dermatodea is the type of
Phyllaria (Le Jolis) Rostafinski 1877, Phyllaria
is superfluous, and we therefore propose a new
genus, Phyllariopsis, to accommodate Laminar­
ia brevipes C. Agardh and Phyllaria purpurascens
(c. Agardh) Rostafinski ex Bornet (Fig. 2).
DIAGNOSIS
Phyllariopsis gen. nov.
Hapteron, discus simplex, aut e brevibus hapteris
non ramosis constans quod vel incremento vel ex­
pansione disci adligationis, primi, simplicis, factum.
Laminae annuae, breviter stipitatae, simplices, a1i­
quando laceratae aetate. Caespites pili, in laminis
iuvenibus, manifesti. Ducti mucilagini non reperti,
medulla laminae solenocystas et allelocystas conti­
nens. Sori sporangiorum in laminae partibus inferi­
oribus facti. Paraphyses sine appendicibus hyalinis.
Zoosporae stigma rubrum habent.
Phyllariopsis gen. nov.
Holdfast a simple disc or with short unbranched
haptera, formed by growth from or expansion of
an initial simple attachment disc. Laminae an­
nual, shortly stipitate, simple, sometimes be­
coming lacerated with age. Hair-tufts evident in
young laminae. Mucilage ducts lacking, medulla
of lamina containing solenocysts and allelocysts.
Sporangial sori formed on lower parts of the lam­
ina. Paraphyses without hyaline appendage. Zoo­
spores with a red eyespot.
TYP
E SPECIES: Phyllariopsis brevipes (c. Agardh)
comb. nov.
Phyllariopsis is distinguished from Saccorhiza
primarily by its simpler holdfast, lacking a sec­
ondary rhizogen (see below). The stipe is always
very short, only a maximum of 5 cm in P. brev­
ipes and 9 cm in P. purpurascens (Hamel 1938),
Henry and South: Phyllariaceae and Phyllariopsis gen. nov.
II
2
1
1
�
Smm
/
Figs 1-5. Phyllariaceae Tilden 1 935 (Laminariales, Phaeophyceae).
Fig. 1. Laminaria brevipes C. Agardh. Holotype, Herb. Agardh No. 1 952. Reproductive sorus (arrow), the dark
region at the base of the lamina, extends to the margin.
Fig. 2. Larninaria purpurascens C. Agardh. Herb. Agardh No. 1 955: 3, one of three specimens on holotype
sheet. Reproductive sorus (arrow) does not extend to margin of the lamina.
Fig. 3. Phyllariella ochotensis Petrov & Vozzhinskaja, holotype. Squash preparation of medulla of lamina
showing presence of trumpet hyphae (arrows). Phase-contrast optics.
Fig. 4. Saccorhiza derrnatodea. Squash preparation of medulla of lamina showing solenocysts (S) and allelocysts
(arrows). Phase-contrast optics.
Fig. 5. Saccorhiza derrnatodea. Upper stipe partially cut, then tom. Solenocysts (arrow) in the medulla are
easily peeled away from the cortex.
12
Phyeologia, Vol. 26 (1), 1987
whereas in S. dermatodea it may reach 60 cm
(Taylor 1957) and in S. polysehides 2 m (Norton
& Burrows 1969) in length.
HISTORICAL BACKGROUND OF
THE PHYLLARIACEAE
The distinctiveness of the two genera in the Phyl­
lariaceae was first explicitly recognized by Ros­
tafinski (1877). This little-known publication (in
Polish) appears to be the first attempt to sub­
divide the "Ordo Laminariaceae (Bory) Rostaf."
on the basis of the internal structure of the lamina
rather than gross external morphology. Rostaf­
inski recognized "Subordo I Simplices", con­
taining Chorda; "Subordo II Fibrosae", com­
posed of Phyllaria and Saeeorhiza; and "Subordo
III Vasculares", composed of all the other Lam­
inariales usually placed in the Laminariaceae,
Alariaceae and Lessoniaceae (Setchell & Gardner
1925). These differences in internal anatomy
among Chorda, Saeeorhiza and most other Lam­
inariales were recognized by Setchell (1891) when
he discussed his observations on S. dermatodea:
the medulla of Chorda contains elongated 'hy­
phae' (cf. South & Burrows 1967) and Saecorhiza
contains thick-walled 'sclcrenchyma' (Setchell
1891), also called 'solenocysts' and 'allelocysts'
(Sauvageau 1918), which form a conducting sys­
tem (Emerson et aI1982), whereas other families
of Laminariales including the Laminariaceae
possess sieve elements, which often appear as
characteristically shaped 'trumpet hyphae'
(Fritsch 1945).
Rostafinski's (1877) partitioning of the Lam­
inariales was accepted by Gobi (1878) but not
by subsequent authors. Gobi thanked Rostafin­
ski for communicating this scheme to him, but
he did not cite Rostafinski's publication, prob­
ably because it had not yet appeared when Gobi
wrote his paper. It appears that later authors were
unaware of Rostafinski's ideas. Setchell & Gard­
ner (1925) clearly regarded Saeeorhiza and Phyl­
laria as belonging in a separate family (Haligeni­
aceae), but they did not explain their reasons for
holding this view nor formally propose the fam­
ily, not did they cite other authors' treatments
of families in the Laminariales. Feldmann (1934)
accepted the distinctiveness of the family Hali­
geniaceae but pointed out that Phyllariaceae was
the earlier name. Tilden (1935) formally pro­
posed "Phyllariaceae familia nova", using only
the lack of hyaline appendages on sorus para-
physes to separate Phyllaria and Saecorhiza from
other lamina-forming Laminariales. Tilden cited
no earlier authors to support her action. Hamel
(1938) and several other subsequent (mostly
French) authors (Huve 1958; Gayral 1958, 1966;
Ardre 1970; Giaccone 1969) have recognized the
family Phyllariaceae, often as "Phyllariacees",
but none cited any authority for the family. Pet­
rov (1974) proposed a division of Laminariales
into two suborders, Phyllarineae, composed of
Phyllariaceae [which he erroneously attributed
to (Kjellm.) Hamel], and Laminarineae, com­
posed of all other Laminariales, including Chor­
da. However, since 1935 most floristic (Taylor
1957; Lund 1959; Rodrigues 1963; Jaasund 1965;
Parke & Dixon 1976; South & Hooper 1980) and
systematic (Fritsch 1945; Papenfuss 1951; Chap­
man 1962; Dawson 1966; Christensen 1980;
Wynne 1982) publications that have treated the
families of the Laminariales have included Sae­
eorhiza and Phyllaria in the Laminariaceae.
DISTINGUISHING CHA
RA
CTERISTICS
OF THE PHYLLARIACEAE
The delimitation of the family as conceived by
Tilden (1935) and others still seems to be correct.
The four species in the Phyllariaceae share sev­
eral characters that serve to separate the family
from the Laminariaceae:
(1) The presence of the unique system of 'so­
lenocysts' and 'allelocysts' (Sauvageau 1918) (Fig.
4) which serve as a conducting system (Emerson
et al 1982) analogous to the sieve tubes of the
Laminariaceae (Fig. 3). These structures have
been described in S. dermatodea (Setchell 189I;
Emerson et a11982) (Figs 4, 5) and S. polysehides
(Sauvageau 1918), and are also found in Phyl­
lariopsis purpuraseens and P. brevipes (Sauva­
geau 1918).
(2) Sorus paraphyses lack the hyaline thick­
ening of the wall at the tip (Setchell 1891; Sau­
vageau 1918; Feldmann 1934; Henry 1986).
Mucilaginous paraphysis tips are characteristic
of species of the Laminariaceae (Feldmann 1934;
Fritsch 1945).
(3) Zoospores possess a red eyespot in the plas­
tid. Although zoospore eyespots have been re­
ported in species in the Laminariaceae, Alari­
aceae and Lessoniaceae, electron microscopy
has shown them to be absent (Henry & Cole
1982). However electron microscopy has con­
firmed the presence of eyespots in zoospores of
Henry and South: Phyllariaceae and PhyllariopSiS gen. nov.
Chorda tomentosa (Toth 1974) in the Chorda­
ceae and Saeeorhiza polysehides (Evans 1966)
and S. dermatodea (Henry 1986). Recent light
microscopical observations confirm the presence
of an eyespot in Phyllariopsis brevipes (Henry
1987). P. purpuraseens remains to be investigat­
ed. Zoospores of Pseudoehorda are also reported
by light microscopy to possess eyespots (Kawai
& Kurogi 1985).
(4) The sexual pheromones of S. polysehides
and S. dermatodea are not lamoxirene or struc­
turally related compounds (Muller et al 1985).
In all the species in the three advanced families
of Laminariales examined to date the sexual
pheromone is lamoxirene (Muller et al 1985).
Neither species of Phyllariopsis has yet been in­
vestigated.
Three additional characters are common to all
species of Phyllariaceae and are present in a few
other Laminariales, or are present in only some
Phyllariaceae and Chorda:
(I) Gametophytes of at least two species of
Phyllariaceae are not sexually dimorphic. Those
of S. dermatodea are monoecious (Henry 1986)
as in Chorda tomentosa (Maier 1984). Game­
tophytes of P. brevipes, although dioecious, are
not distinguishable as to sex until gametangia are
formed (Henry 1987). Gametophytes of S. poly­
sehides are sexually dimorphic as in other Lam­
inariales (Sauvageau 1918); those of P. purpu­
raseens remain unknown.
(2) Hair tufts or pits (cryptostomata) are formed
on young laminae (Setchell 1891; Feldmann
1934). Cryptostomata are also known in a few
other Laminariales: Costaria (Yendo 1911) and
Phyllariella oehotensis Petrov & Vozzhinskaja
(Petrov & Vozzhinskaja 1966) in the Laminar­
iaceae, and Alaria and Undaria (Yendo 1911) in
the Alariaceae. Although tentatively reported in
Nereoeystis luetkeana (Mertens) Postels & Ru­
precht in the Lessoniaceae, the figure illustrating
this finding (MacMillan 1899, pI. 362, fig. 15) is
unconvincing.
(3) Holdfasts lack branched haptera. The hold­
fast of P. purpuraseens is a simple disc no more
than 7 mm in diam. (Feldmann 1934). That of
P. brevipes is composed of short (I cm) un­
branched haptera, pointed or flattening at the tip
upon contact with the substrate (Feldmann 1934;
Henry 1987). These haptera form as outgrowths
of the initial basal disc. The holdfasts of Sae­
eorhiza spp. are formed in a manner unique to
this genus (Setchell 1891; Sauvageau 1918).
Young laminae are attached to the substrate by
13
Table 1. Nomenclature of the four species in the Phyl­
lariaceae
Saccorhiza polyschides (Lightfoot) Batters
Basionym: Fucus polyschides Lightfoot 1777: 936
Synonymy: (for complete synonymy, see Norton &
Burrows 1969)
Laminaria bulbosa (Hudson) Lamouroux 1813: 22
Laminaria elliptica C. Agardh 1820: 119
Saccorhiza bulbosa (Hudson) de la Pylaie 1829: 23
Haligenia bulbosa (Hudson) Decaisne 1842: 345
Saccorhiza dermatodea (de la Pylaie) J. Agardh 1868:
31
Basionym: Laminaria dermatodea de la Pylaie 1824:
180
Synonymy:
Laminaria lorea Bory ex J. Agardh 1848: 130
Phyllitis dermatodea (de la Pylaie) Kiitzing 1849: 567
Haligenia (sect. Phyllaria) dermatodea Le Jolis 1855:
473
Phyllaria dermatodea (de la Pylaie) Gobi 1878: 75
Phyllaria lorea (Bory) Kjellman 1883: 226
Phyllariopsis purpurascens gen. nov., comb. nov.
Basionym: Laminaria purpurascens C. Agardh 1820:
117
Synonymy:
Laminaria brevipes (c. Agardh) J. Agardh 1842: 46
(pro parte)
Phyllitis brevipes (c. Agardh) Kiitzing 1849: 567 (pro
parte)
Laminaria elliptica Montagne 1846: 24 (non L. ellip­
tica C. Agardh 1820: 119, Saccorhiza polyschides)
Haligenia (sect. Phyllaria) brevipes (c. Agardh) Le Jolis
1855: 473 (pro parte)
Phyllaria purpurascens (c. Agardh) Rostafinski ex Bornet 1892: 251
Phyllariopsis brevipes gen. nov., comb. nov.
Basionym: Laminaria brevipes C. Agardh 1820: 116
Synonymy:
Phyllitis brevipes (C. Agardh) Kiitzing 1849: 567 (pro
parte)
Haligenia brevipes (C. Agardh) Lenormand ex Kiitzing
1849: 567 (pro syn.)
Haligenia (sect. Phyllaria) brevipes (c. Agardh) Le Jolis
1855: 473 (pro parte)
Phyllaria reniformis (Lamouroux) Rostafinski ex Bor­
net 1892: 250
=
a small discoid swelling of the base of the stipe,
similar to that of young Phyllariopsis laminae.
In S. polysehides this disc may produce a few
haptera that are essentially unbranched, al­
though they may become lobed at the tips as they
spread against the substrate. A secondary hold­
fast is then produced, beginning as a discoid cir­
cumferential swelling of the stipe circa 0.2 cm
(in S. dermatodea) or 1 cm (in S. polysehides)
above the primary holdfast. In S. dermatodea
this 'rhizogen' (Setchell 1891) forms protuber-
14
Phycologia, Vol. 26 (1), 1987
ances from its margin which grow downward,
forming unbranched haptera surrounding the
original discoid holdfast. A second whorl of hap­
tera completes formation of the holdfast. In S.
polyschides successive whorls of haptera are pro­
duced from the rhizogen which continues to ex­
pand until it forms a bell ('cloche' of Sauvageau
1918). The margin of the bell extends downward
.as it expands, completely surrounding the orig­
inal holdfast and eventually forming an inflated
bulb covered with stubby unbranched haptera
(superbly illustrated by Sauvageau 1918).
At least four of the above characters of the
Phyllariaceae are considered to be primitive in
the Laminariales. Unicellular sorus paraphyses
without thickened tips are otherwise known only
in Chorda; zoospores with eyespots are otherwise
known only in Chorda and Pseudochorda, which
are considered to be the most primitive genera
of Laminariales (Kawai & Kurogi 1985). Like­
wise, monoecious gametophytes are known only
in Chorda tomentosa (Maier 1984) and Pseu­
dochorda (Kawai & Kurogi 1985). Only Lami­
nariaceae, Alariaceae and Lessoniaceae are known
to produce lamoxirene; ectocarpene is produced
by Saccorhiza (I. Maier pers. comm.), Chorda
tomentosa and many other more primitive brown
algae (Muller et al 1985).
Small, simple holdfasts without branched hap­
tera are also probably primitive in the Laminar­
iales. In Chorda the holdfast is a simple disc, but
it is merely an aggregation of filamentous rhi­
zoids (Kanda 1938; Maier 1984). In other Lam­
inariales the holdfast is a parenchymatous tissue
formed by activity of the meristoderm. In a few
Laminariaceae [e.g. Laminaria solidungula J.
Agardh, L. ephemera Setchell, L. yezoensis Mi­
yabe, Cymathere triplicata (Postels & Ruprecht)
J. Agardh] the holdfast is a simple disc. Whether
this character is primitive or derived in these
species, there is no evidence that a Saccorhiza­
like secondary rhizogen is formed.
The many distinctive characters exhibited by
members of the Phyllariaceae indicate that this
family is not only distinct from the Laminari­
aceae, but is in fact the most clearly demarcated
lamina-forming family in the Laminariales. The
solenocysts and allelocysts of the medulla con­
stitute a conducting system comparable to the
sieve tubes of other Laminariales (Emerson et al
1982). They may be interpreted as the parallel
evolutionary development of a mechanism to
meet the long-distance translocation require­
ments of a large thallus. The presence of this
unique and highly differentiated conducting net­
work as well as several primitive characters sug­
gests that the Phyllariaceae diverged from the
other Laminariales early in the evolution of the
order, and that Laminariaceae have given rise to
Alariaceae and Lessoniaceae.
It is generally thought that the Laminariales
had their origin in the North Pacific. This con­
cept is supported by the recent findings concern­
ing Pseudochorda (Kawai & Kurogi 1985), de­
spite the fact that the Pacific Ocean has only one
species of Chorda, whereas the Atlantic has two.
It appears that there are no Phyllariaceae in the
Pacific; rather they are most diverse in the Med­
iterranean and adjacent Atlantic (three species)
where other families of Laminariales are absent
(Pseudochordaceae, Chordaceae, Lessoniaceae)
or sparse compared to the Pacific (Alariaceae,
Laminariaceae). This geographical distribution
supports the concept that the Phyllariaceae are
an offshoot of the main line of evolution in the
Laminariales, which has achieved the greatest
diversity in the colder waters of the North At­
lantic and North Pacific.
ACKNOWLEDGMENTS
We thank P. Silva and M. Wynne for assistance
with nomenclatural problems, F. Ardre and P.
Gayral for information and fragments from La­
mouroux's collections, P. Lassen for access to
collections of the Agardh Herbarium, and Ju. E.
Petrov for access to specimens of Phyllariella.
The Latin diagnosis was provided by Angela
Shipman. This research was supported by Nat­
ural Sciences and Engineering Research Council
of Canada Operating Grant No. 4648 to G.R.
South and by the Newfoundland Institute for
Cold Ocean Science. Final preparation of the
manuscript was expedited by the facilities of Dr
C. Yarish of the University of Connecticut.
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