(1015) Proposal to Conserve the Type of Gracilaria, nom. cons., as G. compressa and Its
Lectotypification (Rhodophyta: Gracilariaceae)
Author(s): M. Steentoft, L. M. Irvine, C. J. Bird
Source: Taxon, Vol. 40, No. 4 (Nov., 1991), pp. 663-666
Published by: International Association for Plant Taxonomy (IAPT)
Stable URL: http://www.jstor.org/stable/1222789
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NOVEMBER 1991
663
(1015) Proposal to conserve the type of Gracilaria, nom. cons., as G. compressa and
its lectotypification (Rhodophyta: Gracilariaceae)
Gracilaria Greville, Alg. Brit. liv, 121. 1830.
T.: G. compressa (C. Agardh) Greville (Sphaerococcus compressus C. Agardh) (typ.
cons. prop.).
In the most recent edition of the International Code of Botanical Nomenclature
(Greuter & al., 1988: 115), Gracilaria confervoides (Stackh.) Greville (1830: liv, 121)
(basionym: Flagellaria confervoides Stackhouse, 1809: 92) is listed as the type of the
conserved name Gracilaria Grev. (see also Farr & al., 1979: 741). This, the earliest
selection of the generic type, was originally made by Schmitz (1889:443; see also: Briquet & al., 1912: 77; Kylin, 1932: 58; Silva, 1952: 265, 293). Fucus confervoides L.
(1763: 1629), although the source of the concept, is an illegitimate epithet-bringing
synonym, a later homonym of E confervoides Hudson (1762: 474) (see Papenfuss,
1950: 195). By default, Stackhouse was the first to create a legitimate species name.
A lectotype has not previously been selected for Fucus confervoides L. In the protologue, Linnaeus cited a synonym from Ray (1724: 52, "Fucus teres rubens
ramosissimus", which in fact refers to Cystoclonium purpureum), but he used a completely new phrase name, "Fucus subdichotomus teres filiformis ramosissimus; ramis
inaequalibus setaceis", suggesting that he based the name E confervoides on a
specimen in his own herbarium.
Of the four sheets in the Linnean Herbarium, LINN (herbarium abbreviations
follow Holmgren & al., 1990) labelled Fucus confervoides, only two, pinned together
(Nos. 1274.111and 1274.112),bear Gracilaria-likealgae. On No. 1274.111,bearing two
specimens side by side, the annotations "confervoides"deleted and reinstated, and
"muscoides" deleted, made by Linnaeus himself, show that he considered this
material carefully, while No. 1274.112bears no annotation other than "Fucus confervoides". We select the larger, right hand specimen on No. 1274.111as the lectotype
specimen of F confervoides L.
Both specimens mounted on No. 1274.111are cystocarpic, with obvious protruding
lateral cystocarps, a fact referredto in the epithet "confervoides",that is to say, in the
air-dried and at most lightly pressed condition, such specimens resemble a string of
beads, not unlike "some of the larger Confervae when shrivelled" (Stackhouse, 1802:
97).
Microscopical examination of sections of a cystocarp of the lectotype through both
base and apex, and including both gonimoblast and pericarp, revealed a lack of
tubular cells between the gonimoblast and pericarp. The gonimoblast is also domelike, and composed of small, apparently densely protoplasmic cells, with chains of
carposporangia radiating from it. These are all generic characteristics of
GracilariopsisDawson (1949: 40; see also Fredericq& Hommersand, 1989b). The Linnean specimen of F confervoides thus belongs to Gracilariopsis Dawson, not to
Gracilaria sensu Fredericq & Hommersand (1989a), and synonymy with Gracilaria
verrucosa (Hudson) Papenfuss (1950: 195) sensu Fredericq& Hommersand (1989a) is
incorrect.
A case can be made for selecting the lectotype of Fucus confervoides L. as the lectotype of Flagellaria confervoides Stackhouse (1809: 92), since Fucus confervoides L.
is given as a synonym. Stackhouse specimens are known to exist in BM, K (now united
664
TAXON VOLUME 40
in BM) and LINN, and in Lamouroux's herbarium in CN (Dixon, 1962), but no
specimen of Flagellaria confervoides Stackh., annotated in his own hand, is available.
Stackhouse's (1809: 92) description of his E confervoides is brief ("Fr[onde] ...
ramulis longis, subsecundis") but he referred to one of his own illustrations of F
longissimus (1802: pl. 16). The illustration of Fucus longissimus is accompanied by a
fuller description: "fronde filiforme, irregulariter et sparsim ramosa - ramis inaequalibus distichis; extremis prolongis; fructu minuto laterali orbiculari depresso",and
referencesto F longissimus S. Gmelin (1768: 134, pl. 13) are added (Stackhouse, 1802:
99-100). Stackhouse commented that "the fructification is in some sorts different",
that is from the "fructu minuto laterali orbiculari depresso"of his description, and the
Stackhouse (1802: pl. 16) drawing of the cystocarp is that of a Cystoclonium,
described by him as "consisting of inflations; these are like imbedded vesicles"
(Stackhouse, 1802: 100).
The specimen of "Fucus longissimus Gmel'' in the sole surviving volume of
Stackhouse's herbarium (BM: British Fuci named, 1801: 8) is a Cystoclonium.
Stackhouse's illustration (1802: pl. 16) does not correspond exactly with the specimen
(see Dixon, 1962). Thus Stackhouse's description included two species, one of which
is, from the synonymy and Latin description cited above (Stackhouse, 1802: 99), a
Gracilariopsis.The other, from the English description, comments in "Observations"
(Stackhouse, 1802: 100) and the specimen found by one of us (BM), is a Cystoclonium.
There is thus nothing to indicate that Stackhouse was describing a Gracilariasensu
Dawson (1949) and Fredericq& Hommersand (1989a).
The retention of Gracilariaconfervoides (Stackh.) Grev., based on Flagellaria confervoides Stackh., as the type of GracilariaGrev., would involve renaming the more
than 100 species of Gracilaria sensu Fredericq & Hommersand (1989a) at present
placed in the genus, and transferring the fewer species at present in Gracilariopsisto
Gracilaria.However,the name Gracilariais well known and accepted not only in algal
taxonomy, but also in world commerce. A number of its species are important sources
of agar and support industries in both tropical and temperate regions.
In the interests therefore of nomenclatural stability (Greuter & al., 1988, Preamble
1), we propose that a new type be conserved for GracilariaGrev.The only species name
included by Grevillethat taxonomically belongs to Gracilariasensu Fredericq& Hommersand (1989a) is the fourth species, G. compressa (Agardh) Greville (1830: 125),
based on Sphaerococcuscompressus Agardh (1822: 308). According to the protologue
of the basionym, the ideal lectotype specimen would be at LD, have a holdfast ("radix
scutata"), be non-carposporangial ("fructus ignotus"), and be either from Spain
(Cadiz (Gades), leg. Cabrera), or Italy (Oneglia, leg. Brongniartjun.) (cf. Agardh,
1822: 308; Bird, unpubl. obs.).
The Brongniart syntype (LD No. 28988) proved to be a member of the Dictyotales
(Phaeophyta) (Steentoft, pers. obs.), but from the remaining syntypes, Bird selected
and annotated as the lectotype specimen LD No. 28984 (Cabrera,sine loco), the only
non-cystocarpic specimen with a holdfast, which also possesses the faint transverse
striations of the cuticle referred to by Agardh (1822: 308, "transversim striatae").
Cabrera collected both in Cadiz and Malaga (Cremades Uguarte, 1990), but the lectotype compares very well with LD No. 28987 (Cadiz), LD No. 28991 ("misit Cabrera"
in Agardh's hand) and LD No. 28992 (sine coll., sine loco), which, in addition, is
tetrasporangial, as is the lectotype. All have the unusually thick cuticle characteristic
of European Atlantic Gracilariacompressa (Steentoft, pers. obs.), and the striations
noted by Agardh are apparently caused by the contraction of this cuticle in drying.
NOVEMBER 1991
665
Gracilaria compressa is nowadays known by its taxonomic synonym, G. bursapastoris (S. Gmelin) Silva (1952:265). This latter name is the type of Ceramianthemum
Donati ex Leman, against which Gracilariahas been conserved. This was proposed by
Bdrgesen & al. (1909: 323, 325) and was first listed as conserved in the Brussels Rules
(Briquet & al., 1912;see also Silva, 1952).
The proposed conservation of the type of Gracilariawith a synonym of G. bursapastoris conflicts with the infrageneric nomenclature of Yamamoto (1975), wherein G.
subg. Gracilaria(the autonym) encompassed the then-accepted generic type, G. verrucosa (Hudson) Papenfuss, in accordance with Art. 22 of the International Code of
Botanical Nomenclature (Greuter& al., 1988). If this proposal is passed, G. subg. Textoriella Yamamoto, including G. bursa-pastoris and its synonym G. compressa, will
yield to the autonym in the system of subgenera, and the subgenus in which G. verrucosa falls will requirea different name.
Acknowledgments
We wish to express our thanks to the Library of the University of Lund for access to the Nord-
stedt archiveand to Dr. TygeChristensen,Copenhagen,for searchingthat archivefor us; to
Professors Ove Almborn and Sven Snogerup, former and current directors of the Botanical
Museum of the University of Lund for hospitality and access to the Agardh herbarium, and to
Dr. Per Lassen for curatorial help; to the Linnean Society of London and Dr. N. Robson for
access to the Linnean herbarium, and to the Appleyard Fund for a grant to the first author; to
Dr. C. E. Jarvis, Mr. J. H. Price and the Natural History Museum, London, for discussions on
the typification of Linnean species, for bringing to our attention information on Clemente's
herbarium and for access to specimens and literaturerespectively;to Dr. C. Billard, University
of Caen, for her searches in the Lamouroux herbarium, and to Professor Paul Silva, University
of California at Berkeley,for sharing insights into the typification of Gracilaria.
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Proposed by: M. Steentoft, Portsmouth Polytechnic Marine Laboratory, Hayling Island,
POll ODG, U.K.; L. M. Irvine, Natural History Museum, London, SW7 5BD, U.K.; and C. J.
Bird, Institute for Marine Biosciences, National Research Council, Halifax, N.S. B3H 3Z1,
Canada.