Marine benthic macroalgae collected by Vieillard from New Caledonia and described as new species by Ku tzing

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Phycologia (2005) Volume 44 (5), 536–549 Published 15 September 2005

Marine benthic macroalgae collected by Vieillard from New Caledonia and
described as new species by Kützing

ALAN J.K. MILLAR1* AND WILLEM F. PRUD’HOMME VAN REINE2
1
Royal Botanic Gardens Sydney, Mrs Macquaries Road, Sydney, NSW 2000, Australia
2
Nationaal Herbarium Nederland, University of Leiden, PO Box 9514, 2300 RA, Leiden, The Netherlands

A.J.K. MILLAR AND W.F. PRUD’HOMME VAN REINE. 2005. Marine benthic macroalgae collected by Vieillard from New
Caledonia and described as new species by Kützing. Phycologia 44: 536–549.

During the 19th century, the German phycologist Friedrich Kützing described 66 new species of marine benthic macroalgae
that had been collected by the French naturalist Eugène Vieillard from New Caledonia and the nearby Ile des Pins in the
southwestern Pacific Ocean. Many of these species have not since appeared in the literature, nor have they been critically
re-examined in light of modern classifications and nomenclatural status. As a direct result of the present research, many
have now been found to be conspecific with either previously described species, or are species for which more recent names
have been unwittingly applied. This work has resulted in seven new combinations and three new names being proposed:
Cladophora mamillata Leliaert nom. nov. (replaced name: Valonia cladophora Kützing); Meristotheca polychotoma (Kütz-
ing) Millar comb. nov. (basionym: Euhymenia polychotoma Kützing); Polyopes emarginatus (Kützing) Millar comb. nov.
(basionym: Grateloupia emarginata Kützing); Halichrysis irregularis (Kützing) Millar comb. nov. (basionym: Iridaea ir-
regularis Kützing); Halymenia kuetzingii Millar nom. nov. (replaced name: Iridaea fimbriata Kützing); Laurencia kuetzingii
Millar nom. nov. (replaced name: Laurencia flagellifera Kützing); Sarcodia marginalis (Kützing) Millar comb. nov. (bas-
ionym: Mastocarpus marginalis Kützing); Schottera angustifolia (Kützing) Millar comb. nov. (basionym: Sphaerococcus
angustifolius Kützing); Rhodymenia caulescens (Kützing) Millar comb. nov. (basionym: Sphaerococcus caulescens Kützing);
Gracilaria chondracantha (Kützing) Millar comb. nov. (basionym: Sphaerococcus chondracanthus Kützing).

INTRODUCTION city in 1845. Kützing then made many drawings and descrip-
tions of Sonder and Binder material, and he often managed to
At present count, some 72 species of marine benthic macroal- get small fragments from these herbaria. He also annotated
gae have been described based on type material collected from many of these specimens in his own hand. Much of Sonder’s
the French Pacific islands of New Caledonia and the Ile des herbarium now resides at MEL because it was purchased by
Pins. The majority (66) were collected by Eugène Vieillard Ferdinand von Mueller (the then Director of the Botanic Gar-
(1819–1896) and described by Friedrich Kützing during the dens in MEL) in the 1800s.
mid-1800s (1863a, 1863b–1869). Many have not appeared in Kützing was well aware of the value of original or type
the literature since, nor have they been critically re-examined material and occasionally he annotated specimens that he had
taking into account updated taxonomy and nomenclature. described or illustrated with the word ‘original’ or ‘unicum’.
A.J.K. Millar & C.E. Payri (unpublished observations) are He made all his figures personally and did that from material
presently working on numerous deep-water collections from at hand or from sketches he had made beforehand. These fig-
New Caledonia and, as a consequence, have discovered many ures were engraved in stone, and thus each figure in his Ta-
new records and species for the region. In an effort to clean bulae Phycologicae is a mirror image of the actual specimen.
up the existing records of marine macroalgae from New Ca- After 1845, Kützing had contacts with Sonder (and maybe
ledonia, the present authors have examined all of Vieillard’s also with von Mueller), but it is assumed that he retained the
specimens that were described by Kützing. Much of the New ‘type’ specimens, especially of those he made figures from.
Caledonian algal literature has been reviewed by Garrigue & Thus Kützing’s specimens in L should be considered the ho-
Tsuda (1988) and is not repeated here. lotypes and lectotypes, whereas those in MEL should be iso-
Kützing’s herbarium is deposited mostly in the Nationaal types or isolectotypes. As for designating specimens as lec-
Herbarium Nederland in Leiden (L), although many of Kütz- totypes or holotypes, we were reluctant to accept Vieillard
ing’s isotype specimens can be found in the National Herbar- material as a holotype when there is no label in Kützing’s
ium of Victoria in Melbourne (MEL), and it was at these two hand and when there is no figure in the Tabulae Phycologicae
institutions, and at the Royal Botanic Gardens in Sydney, that that corresponds to the pressed plant. Additionally, the correct
much of the present work was carried out. Most of the Kütz- number of the figure in the Tabulae Phycologicae must also
ing collections in MEL are samples that were in the Sonder be recorded by Kützing himself directly on the sheet. In such
(or Binder) collections in Hamburg when Kützing visited that instances where there is only one Vieillard specimen filed un-
der Kützing’s name, it must, by default, be considered the
* Corresponding author (alan.millar@rbgsyd.nsw.gov.au). holotype.
536

Millar & Prud’homme van Reine: Vieillard marine algae from New Caledonia 537

Eugène Vieillard’s collections from New Caledonia were Vieillard from Ile des Pins. This is the only specimen and thus
made during the periods 1855–1860 and 1862–1867. Eugène is the holotype. The sheet is labelled Caulerpa freycinetii var.
Vieillard was himself a naval surgeon and along with his col- typica forma serrulata (J. Agardh) Weber-van Bosse. That
league, Emile Deplanche, was posted to the four garrisons on species is considered to be conspecific with C. serrulata
New Caledonia (Baláde, Canala, Gatope and Wagap) where (Forsskål) J. Agardh (1837) by Børgesen (1932) and we
both also serviced the passing vessels (Vieillard & Deplanche would agree with that decision. Caulerpa vieillardii is thus a
1863). Apart from a year’s leave (1861), when he visited New heterotypic synonym of C. serrulata.
Zealand to collect ferns (his principal passion), before moving Chauvinia microphysa Kützing 1863a: 11.
on to Sydney and back to Noumea, the amateur botanist spent Type is L 937,337-749 (barcode L 0194018) collected by
11 years (1855–1867) collecting both terrestrial and marine Vieillard # 1958 from Baláde, New Caledonia. Specimens of
plants. His collections were prodigious, and figures of 400 the two sheets represent the same species. Weber-van Bosse
species represented by 12,000–15,000 specimens have been (1898: 361), who presumably thought Kützing’s name was
estimated as his cache (M.E. McKee & H.S. McKee 1981). unpublished, used his epithet ‘microphysa’ when she de-
All these specimens were sent back to France by military post scribed the forma of Caulerpa racemosa v. clavifera f. mi-
to his friend, Sébastien-René Lenormand, who was a lawyer crophysa Weber-van Bosse. That forma, however, is based on
by profession but keen amateur botanist who also lived in a type specimen from Macassar (Celebes, Indonesia). This
Vieillard’s home region of Vire (Calvados, Normandy, Indonesian species was raised to specific rank by Feldmann
France). Lenormand then distributed the specimens to bota- (1955) and is generally known as Caulerpa microphysa (We-
nists and museums throughout Europe, and at least for the ber-van Bosse) Feldmann (see Silva et al. 1996). Thus the two
algal collections, we know they are housed in L, Paris (PC) lineages are separate. Kützing’s New Caledonian species, if
and Caen (CN; Valet 1967), with one small collection existing transferred to the genus Caulerpa, cannot retain the epithet as
in Michigan (M. Wynne, personal communication). These col- it would become a later homonym. On morphological char-
lections are numbered as species, not as specimens, which acters, however, Chauvinia microphysa appears to be conspe-
means that a single number is used for a species when col- cific with the earlier described Caulerpa lentillifera J. Agardh
lected on different localities or on different dates (or both). (1837).
This may also mean that under one number similar specimens Cladophora physarthra Kützing 1863a: 11.
of different species can occur (M.E. McKee & H.S. McKee Type is L 937,253-471 (barcode L 0054853) collected by
1981). So respected was Vieillard as an experienced botanist Vieillard # 1977 from New Caledonia. This species was not
that ultimately he became Professor of Botany at CN (France), illustrated by Kützing in his Tabulae Phycologicae, and there
suggesting that his surgical skills presumably became a bit are two specimens on this sheet. We herein designate the spec-
rusty. Anna Weber-van Bosse bought parts of the Lenormand imen marked ‘1977’ as the lectotype. It is a profusely
and Suringar herbaria, and this explains why they are now branched specimen that we would identify as the earlier Bood-
housed in L. We do not know if Kützing returned specimens lea composita (Harvey) Brand (1904).
to Lenormand after he had identified them (and thus had his Cladophora radians Kützing 1863a: 10.
handwriting on them). Type is L 937,253-433 (barcode L 0194019) collected by
The species in this paper are arranged firstly by Phylum Vieillard # 2006 from Ile des Pins, New Caledonia. We herein
and then alphabetically by basionym genus and species. All designate this specimen as the lectotype because Kützing did
of these specimens can be seen either on the L Collections not illustrate this species in his Tabulae Phycologicae. We
Database website [www.nationaalherbarium.nl/virtual] or in believe this species is conspecific with Acrosiphonia arcta
Kützing’s Tabulae Phycologicae and we direct the reader to (Dillwyn) Gain (1912).
that site and Kützing’s work rather than re-illustrate all the Dictyosphaeria ulvacea Kützing 1863a: 12.
specimens here. We have illustrated only those specimens for Type is L 937,183-35 (barcode L 0055026) collected by
which a name change has resulted. Vieillard # 1978 from Baláde, New Caledonia. There are two
envelopes on the sheet with each containing a specimen. Valet
(1966, 1967) examined these and stated that the one on the
MATERIAL AND METHODS left was the correct type specimen and was conspecific with
the more recently described D. intermedia Weber-van Bosse,
All specimens were hand-sectioned, using single-edged razor but the specimen on the right was a juvenile form of D. cav-
blades and were placed firstly in an aniline blue solution of ernosa (Forsskål) Børgesen. We have no reason to dispute
93% distilled water, 1 g aniline blue and 7% acetic acid. They this, but note that the specimens are small, not well pressed,
were then mounted in a aniline blue solution made according and thus difficult to identify with certainty.
to the methods of Millar & Wynne 1992, and observed using Polyphysa spicata Kützing 1863a: 12. 1866: 1, table 1c–f.
an Olympus CH-2 compound microscope. Herbarium abbre- Type is L 937,183-066 (barcode L 0054969) collected by
viations follow Holmgren et al. 1990. Vieillard from Baláde, New Caledonia. This specimen was
used by Kützing to prepare fig. 1d and is thus the holotype.
This is the basionym of Halicoryne spicata (Kützing) Solms-
RESULTS AND OBSERVATIONS Laubach (1895: 31, pl. 4, figs 3, 7, 9, 11), a currently accepted
name and species (Berger et al. 2003). A homotypic synonym
Chlorophyta is Pleiophysa spicata (Kützing) Sonder ex De Toni.
Caulerpa vieillardii Kützing 1863a: 11. Struvea delicatula Kützing 1866: 1, table 2e–g.
Type is L 937,336-456 (barcode L 0194017) collected by Type is L 937,183-109 (barcode L 0237969) collected by

538 Phycologia, Vol. 44 (5), 2005

Vieillard # 2111 from Wagap, New Caledonia. This is the Clerck (2003) the specimens in question represent the Vaugh-
holotype, according to F. Leliaert (personal communication). aniella stage of an unidentified species of Padina.
This species is generally considered to be a synonym of Stru- Dictyota vieillardii var. B. filiformis Kützing 1863a: 14.
vea anastomosans Harvey, which is now considered to rep- Type is L 936,291-4 (barcode L 0194023) collected by
resent Phyllodictyon anastomosans (Harvey) Kraft & Wynne Vieillard # 2035a from Ile des Pins, New Caledonia. The sin-
(1996: 139) and we would agree with this decision. F. Leliaert gle specimen annotated accordingly in Kützing’s hand writing
(personal communication) maintained the taxon as distinct is thus the holotype. De Clerck & Coppejans (1997) and De
from anastomosans, only in that he refers to both as pheno- Clerck (2003) have determined that these specimens represent
demes of what he calls the Cladophoropsis composita com- the Vaughaniella stage of an unidentified species of Padina.
plex. We further point out that there seems to be no difference be-
Struvea scoparia Kützing 1863a: 12. 1866: 1, table 2a–d. tween these specimens and those of the type variety Dictyota
Type is L 937,183-105 (barcode L 0062221) collected by vieillardii as above.
Vieillard from Ile des Pins, New Caledonia. The type sheet
has two specimens. The one on the left was used by Kützing Rhodophyta
to prepare fig. 2a and is thus the holotype. This is the basion-
ym of Apjohnia scoparia (Kützing) Valet 1976. Murray & Acrocarpus capitatus Kützing 1868: 12, table 35a–c.
Boodle (1888: 266) and more recently Denizot (1965) consid- Type is L 941,11-104 (barcode L 0194024) collected by
ered the New Caledonian species to be synonymous with the Vieillard # 2048 in 1863 from Wagap, New Caledonia. It is
earlier described and southern Australian species Apjohnia difficult to determine which specimen was used by Kützing
laetivirens Harvey, and the type sheet has been labelled ac- to prepare fig. 35a, but all specimens represent the same spe-
cordingly. Womersley (1984) considered the two species to be cies. Dawson (1954) concluded that Acrocarpus capitatus was
distinct. Although we would agree with the former decision conspecific with Gelidiopsis intricata (C. Agardh) Vickers
(1905). We concur with Dawson’s decision and further point
(based on general habit), freshly collected specimens and mo-
out the striking similarity Price & Scott’s (1992: 51, fig. 13)
lecular analyses will help define these taxa more accurately.
description and illustration bears to Kützing’s illustrations and
Valonia cladophora Kützing 1863a: 12. 1866: 1, table 1a, b.
specimens, especially the swollen (tetrasporic) apices.
Fig. 1
Acrocarpus delicatulus Kützing 1868: 12, table 35g, h.
Type is L 937,183-020 (barcode L 0054999) collected by
Type is L 941,11-103 (barcode L 0194025) from New Ca-
Vieillard # 1975 from Baláde, New Caledonia. Only one spec-
ledonia, but no collector stated. It is not clear which of the
imen exists under this name and it is thus the holotype. How-
several tufts on this sheet have been used by Kützing to pre-
ever, a separate specimen labelled with the manuscript name
pare fig. 35g, but all represent the same species. This is the
Valonia polyclados (L 937, 183-019), corresponds to Kütz-
basionym of the species Gelidium delicatulum (Kützing) P.
ing’s (1866, table 1) illustration of Valonia cladophora. Al-
Crouan & H. Crouan (as listed by Mazé & Schramm 1878)
though both specimens represent the same species, Kützing’s and also Gelidium delicatulum (Kützing) Setchell (1924: 164).
original description (growing in tufts, filiform, entangled, ir- Taylor (1960: 356) listed Gelidium delicatulum as a synonym
regularly branched, cylindrical throughout, diameter of the fil- of Gelidium crinale (Turner) Gaillon (1828), a decision that
aments identical throughout, apical branches frequently re- we agree with. The specimen’s habit and cross-sections, which
curved and obtuse) clearly refers to the type specimen L reveal rhizines in the medulla of what are very thin (160 mm),
937,183-020. Valonia cladophora falls morphologically with- compressed, irregularly branched axes, match well with G.
in the circumscription of the Cladophora section Aegagropila crinale as interpreted by Millar & Freshwater (2005).
as described by van den Hoek (1963, 1984), and closely re- Amansia fasciculata Kützing 1865: 2, table 4a–d.
sembles Cladophora catenata (Linnaeus) Kützing (F. Leliaert, Type is L 938,19-61 (barcode L 0194026) collected by
personal communication). It can be distinguished from that Vieillard # 2042 from New Caledonia. There are three spec-
species by its larger cell diameter and by the mamillose cross- imens in the one envelope (all representing the same species).
walls. In transferring Valonia cladophora to the genus Cla- One of these was used by Kützing to prepare fig. 4a and is
dophora, the species epithet needs to be changed in order to thus the holotype. This species is considered conspecific with
avoid a tautonym (Article 23 of the International Code for the earlier described Amansia glomerata C. Agardh (1822) by
Botanical Nomenclature). We therefore propose the new name Norris (1988) [Heterotypic synonym: Melanamansia glomer-
Cladophora mamillata Leliaert nom. nov. (replaced name: Va- ata (C. Agardh) Norris (1995)] and we would agree with that
lonia cladophora Kützing 1863a: 12) the epithet referring to decision. N’Yeurt (2002) and Masuda & Abe (2002) demon-
the typical mamillose cross-walls. strated sound reasons for the merger of Melanamansia Norris
(1988) with the earlier Amansia Lamouroux (1809).
Heterokontophyta Bangia vieillardii Kützing 1863a: 10.
Type is L 939,271-79 (barcode L 0194027) collected by
Dictyota vieillardii Kützing 1863a: 14. Vieillard # 2010 from New Caledonia. We consider this single
Type is L 936,291-5 (barcode L 0194020) collected by specimen to be the holotype. De Toni (1897: 12) listed it in
Vieillard # 2035 from New Caledonia. There are several spec- his ‘species mihi omnio ignotae aut valde dubiae’. The ma-
imens under this name, and Kützing did not illustrate the spe- terial is not well preserved and thus difficult to identify with
cies in his Tabulae Phycologicae. We herein designate the certainty.
specimen annotated with Kützing’s handwriting as the lecto- Bostrychia vieillardii Kützing 1863a: 18. 1865: 10, table
type. According to De Clerck & Coppejans (1997), and De 26a–e.

Millar & Prud’homme van Reine: Vieillard marine algae from New Caledonia   539

Figs 1–6. Kützing’s type specimens.
  Fig. 1. Valonia cladophora (5 Cladophora mamillata). Scale 5 10 mm.
  Fig. 2. Euhymenia polychotoma (5 Meristotheca polychotoma). Scale 5 30 mm.
  Fig. 3. Grateloupia emarginata (5 Polyopes emarginatus). Scale 5 10 mm.
  Fig. 4. Iridaea irregularis (5 Halichrysis irregularis). Scale 5 30 mm.
  Fig. 5. Iridaea fimbriata (5 Halymenia kuetzingii). Scale 5 10 mm.
  Fig. 6. Laurencia flagellifera (5 Laurencia kuetzingii). Scale 5 20 mm.

540 Phycologia, Vol. 44 (5), 2005

Type is L 935,329-10 (barcode L 0055696) collected by of this name, nor a type of any species or name. An annotation
Vieillard # 2039 from New Caledonia. This specimen corre- label by Max H. Hommersand dated ‘1997’ states it is rep-
sponds best with Kützing’s fig. 26a and is thus the holotype. resentative of Nothogenia fastigiata (Bory) Parkinson.
Possible isotypes L 938,92-259 and MEL 672311. Post (1936) Chondrus edulis Kützing 1867: 19, table 63c–e.
placed this species under Bostrychia binderi Harvey, whereas Type is L 938,92-309 (barcode L 0055877) collected by
King & Puttock (1989) placed it in Bostrychia tenella (La- Vieillard # 2060 from New Caledonia. This is the specimen
mouroux) J. Agardh ssp. tenella and we would agree with the Kützing used to prepare fig. 63c and should thus be the ho-
latter decision. lotype. The sheet is annotated in Kützing’s hand ‘Les indi-
Bostrychia vieillardii var. B. pectinata Kützing 1865: 10, gènes mangent cette algue’ (The inhabitants eat this alga).
table 26f, g. This is the basionym of Eucheuma edule (Kützing) Weber-van
Type is L 935,329-13 (barcode L 0055694) collected by Bosse, a currently accepted name and species (Silva et al.
Vieillard # 2040 from New Caledonia. This specimen corre- 1996).
sponds best with Kützing’s fig. 26f, g and Kützing added an Digenea vieillardii Kützing 1865: 11, table 28f–l.
exclamation mark on its label. Possible isotype: L 939,271- Type is L 940,347-120 (barcode L 0194029) collected by
64. King and Puttock (1989), who placed this variety within Vieillard from New Caledonia. This specimen was used by
Bostrychia tenella (Lamouroux) J. Agardh ssp. tenella, stated Kützing to prepare fig. 28f and is thus the holotype. This is
that the type (which they had not seen) is from Yaté, but no considered to be a heterotypic synonym of Digenea simplex
such annotation is marked on the sheet. We agree with their (Wulfen) C. Agardh by De Toni (1903: 963), and our exam-
taxonomic decision. ination suggests no reason to dispute this.
Callithamnion vieillardii Kützing 1863a: 14. 1863b: 2, ta- Echinocaulon ramelliferum (Kützing) Kützing 1868: 14,
ble 3c, d. table 39d–f.
Type is L 938,92-234 (barcode L 0055703) collected by Type is L941,11-63 (barcode L 0055886) collected by
Vieillard from New Caledonia. Kützing used this specimen to Vieillard # 2061 from Baláde, New Caledonia. See below un-
prepare fig. 3c, d and it is thus the holotype. The specimen is der entry for Gelidium ramelliferum Kützing (1863a: 16).
fragmented and badly preserved and thus difficult to identify Euhymenia polychotoma Kützing 1867: 22, table 73a–d.
with certainty. Judging from Kützing’s illustrations, this spe- Fig. 2
cies does appear to represent a species of the Callithamnion Type is L 941,11-2 (barcode L 0194030) collected by Vieil-
group in that it is has monosiphonous, ecorticate axes with lard # 2025 from Ile des Pins, New Caledonia. There are two
one lateral borne alternately on each axial cell. specimens in this collection, and one has been used by Kütz-
Champia vieillardii Kützing 1866: 14, table 37e, f. ing to prepare fig. 73a and is therefore the holotype. J. Agardh
Type is L 938,303-132 (barcode L 0055786) collected by (1876: 166) placed this species in his inquirendae section and
Vieillard # 2109 in 1863 from Wagap, New Caledonia. This related it to Cryptonemia ligulata J. Agardh. Euhymenia po-
specimen was used by Kützing for his fig. 37e and is thus the lychotoma was listed by De Toni (1905: 1613) under the genus
holotype. This is a valid name and currently accepted species Cryptonemia as ‘species quod genus valde dunia’. Cross sec-
of the genus Champia (Dawson 1954; Millar 1990: 372-373; tions of these plants show an anatomy typical of the genus
Coppejans & Millar 2000: 325). Meristotheca. Within that genus, only Meristotheca papulosa
Chondrococcus spinulosus Kützing 1868: 11, table 32a–c. has a habit that displays superficially similar dichotomous
Type is L 940,284-248 (barcode L 0194028) collected by blades. Euhymenia polychotoma differs from all the Meristo-
Vieillard # 2034 from New Caledonia. There are two speci- theca species in having bluntly rounded apices and very reg-
mens representing the same species on this sheet, and it is ularly dichotomous blades. Meristotheca polychotoma (Kütz-
difficult to determine exactly which one has been used by ing) Millar comb. nov. [Basionym: Euhymenia polychotoma
Kützing to prepare fig. 32a. This is the basionym of Portieria Kützing 1867: 22, table 73a–d].
spinulosa (Kützing) P.C. Silva (Silva et al. 1987). We cannot Gelidium caerulescens Kützing 1868: 19, table 56c, d.
find any differences between this species and the earlier de- Type is L 941,11-91 (barcode L 0056117) collected by
scribed and widely distributed Portieria hornemannii (Lyng- Vieillard # 2103 in 1863 from Wagap, New Caledonia. This
bye) P.C. Silva and suggest that they are conspecific. single specimen was used by Kützing to prepare fig. 56c and
Chondrus coccineus Kützing 1867: 18, table 62a, b. is thus the holotype. Five specimens on another sheet L
Type is L 940,284-245 (barcode L 0055804) collected by 941,11-98 (barcode L0056116) are all isotypes. One of the
Vieillard # 2105 in 1863 from Wagap, New Caledonia. Kütz- specimens is cystocarpic, and sections reveal an internal struc-
ing (1867) specifically cites the specimen # 2105 of Vieillard ture typical of the genus Pterocladiella as illustrated by San-
and used the same plant for fig. 62a (which is a mirror image). telices & Hommersand (1997, fig. 3b). This is the basionym
This is therefore the holotype. De Toni (1897: 183) placed of the currently accepted name and species Pterocladiella cae-
this in his species inquirendae with no comment as to its rulescens (Kützing) Santelices & Hommersand (1997). Ho-
affinities. The habit and internal anatomy of this specimen motypic synonym: Pterocladia caerulescens (Kützing) San-
show it to be the same as Grateloupia emarginata Kützing, telices (1976).
also from Wagap, New Caledonia. Chondrus coccineus Kütz- Gelidium multicorne Kützing 1868: 23, table 66a–d.
ing thus becomes a heterotypic synonym of Polyopes emar- Type is L 941,11-92 (barcode L 0055897) collected by
ginatus (Kützing) Millar (see below). Another specimen in L Vieillard # 2095 from Baláde, New Caledonia. This specimen
is filed under the name Chondrus coccineus Kützing has been used by Kützing to prepare fig. 66a and is thus the
(L940,284-428 5 barcode L 0055805) and has a ‘Type’ label holotype. This is the basionym of Carpopeltis (?) multicornis
attached to it as well as the location ‘Chile’. It is not the type (Kützing) De Toni (1905: 1604) and Cryptonemia multicornis

Millar & Prud’homme van Reine: Vieillard marine algae from New Caledonia             541

(Kützing) Sonder (1871: 63). Examination of the New Cale-          (see Silva et al. 1996). A specimen of Harvey’s Ceylon exsic-
donian specimen shows it to belong to Carpopeltis and also          catae # 50 is filed in NSW (287603), and it is extremely sim-
to be identical to the widely distributed Indo-Pacific species      ilar to the Vieillard specimen (# 2096) from New Caledonia.
Carpopeltis maillardii (Montagne & Millardet) Chiang (1970)         Although the Ceylonese plants are taller; the branch widths,
that was recently recorded and illustrated from nearby Papua        dichotomous branching, lateral proliferations and internal
New Guinea by Coppejans & Millar (2000, fig. 8).                    anatomy are precisely comparable to Vieillard’s specimens
   Gelidium ramelliferum Kützing 1863a: 16.                        from New Caledonia. They represent the same species of the
   Type is L941,11-63 (barcode L 0055886) collected by              genus Polyopes. Thus the correct name for this species is Po-
Vieillard # 2061 from Baláde, New Caledonia. Kützing (1868:       lyopes emarginatus (Kützing) Millar comb. nov. [Basionym:
14) specifically cited the Vieillard sheet # 2061, and this is      Grateloupia emarginata Kützing 1867: 9, table 29a, b]. Chon-
the same specimen as used by him to prepare fig. 39d, e. It         drus coccineus Kützing is a heterotypic synonym (see above).
is thus the holotype. Weber-van Bosse has labelled the spec-           Grateloupia opposita Kützing 1867: 9, table 31a, b.
imen ‘Gelidiopsis rigida (Vahl) W.v.B.’ [5 Gelidiopsis rigida          Type is L 938,92-311 (barcode L 0194032) collected by
(C. Agardh) Weber-van Bosse (1904)], which is now consid-           Vieillard # 1927 from New Caledonia. This specimen was
ered a heterotypic synonym of Gelidiella acerosa (Forsskål)        used by Kützing to prepare fig. 31a and is thus the holotype.
Feldmann & Hamel (1934), and we would identify this spec-           This is a later homonym for Grateloupia opposita Kützing
imen with the latter species. Gelidium ramelliferum and             (1863a: 15) that he described based on a specimen he cited
Echinocaulon ramelliferum (Kützing) Kützing (1868: 14, ta-        as being from the New Hebrides and collected by Ferdinand
ble 39d–f) thus become heterotypic synonyms of Gelidiella           von Müller. We believe that Kützing may have erred in this
acerosa.                                                            instance because as far as we know, nobody has ever seen the
   Gelidium repens Kützing 1868: 21, table 60a, b.                 New Hebrides material, nor does it exist in L. Because the
   Type is L 941,11-84 (barcode L 0055781) collected by             specimen was a Müller collection, Kützing may have returned
Vieillard # 2104 from Wagap, New Caledonia. Kützing used           it to Müller in Melboure, but it was not located there either.
this specimen to prepare fig. 60a so it is the holotype. This is    His 1863 and 1867 descriptions are very similar, and he
the basionym of Gelidiopsis repens (Kützing) Weber-van Bos-        marked the New Caledonian specimen with ‘unicum’, illus-
se (Silva et al. 1996), a currently accepted name and species.      trated it, and was thus certain of its provenance. De Toni
Homotypic synonym: Ceratodictyon repens (Kützing) R.E.             (1905: 1572) suggested that Vieillard’s specimen was more
Norris (1987).
                                                                    like a Hypnea species. Weber-van Bosse (1928: 415) reduced
   Gongroceras subtile Kützing 1863a: 14. 1863b: 1, table 2a, b.
                                                                    this species (the New Caledonian sp.) to a form of Eucheuma
   Type is L 938,303-191 (barcode L 0193965) collected by
                                                                    muricatum (S.G. Gmelin) Weber-van Bosse forma opposita
Vieillard # 1904 from New Caledonia. It is epiphytic on the
                                                                    (Kützing) Weber-van Bosse. Silva et al. (1996) have shown
type specimen of Liagora tomentosa Kützing (see below).
                                                                    that the earliest name for this species complex is Eucheuma
This is the only specimen in the collection and thus the ho-
                                                                    denticulatum (Burman) Collins & Hervey. We are not sure if
lotype. Grunow (1874: 31), realising this was a species of
                                                                    the forma merits maintenance.
Ceramium, proposed the combination Ceramium kuetzingian-
                                                                       Griffithsia heteromorpha Kützing 1863a: 14. 1863b: 2, ta-
um because the name Ceramium subtile J. Agardh (1851) al-
                                                                    ble 3a, b.
ready existed for a Caribbean species. However, we would be
                                                                       Type is L 941,61-42 (barcode L 0055921) collected by
inclined to relate the New Caledonian species to, and suggest
it is conspecific with, the earlier described and very wide-        Vieillard # 1906 (no. 291) from New Caledonia. There are
spread Ceramium flaccidum (Kützing) Ardissone (1871).              four envelopes on the sheet, but Kützing (1863b) specifically
   Grateloupia emarginata Kützing 1867: 9, table 29a, b.           cited specimen # 1906 and we suggest this specimen (upper
   Fig. 3                                                           envelope of second row) was most probably used for fig. 3a
   Type is L 938,92-310 (barcode L 0194031) collected by            and thus is the holotype. This is a currently accepted name
Vieillard # 2096 in 1863 from Wagap, New Caledonia. This            and species (Silva et al. 1996).
specimen was specifically cited by Kützing in the protologue          Gymnophloea gracilis Kützing 1867: 1, table 1a–c.
and has been used to prepare fig. 29a. It is thus the holotype.        Type is L 939,69-914 (barcode L 0194033). Kützing illus-
A possible isotype is in MEL (684993). When describing the          trated this specimen, which he used for fig. 1a, but it is not
species Cryptonemia ligulata, J. Agardh (1876: 162) did so          from New Caledonia as stated in the protologue, but from
based on a specimen of Harvey’s Ceylon exsiccatae (# 50)            Pointe de Galle, Ceylon (Sri Lanka). This species is conspe-
that Harvey had named on the label, but not validly published       cific with Dermonema virens (J. Agardh) Pedroche & Avila
as Gymnogongrus ligulatus. J. Agardh, in this same proto-           Ortiz (Silva et al. 1996: 119 and 915–916).
logue, cited Gymnogongrus ligulatus Kützing (1869: 23, table          Hypnea aspera Kützing 1868: 14, table 40e–h.
63a, b) and Grateloupia emarginata Kützing (1867: 9, table            Type is L941,97-186 (barcode L 0055947) collected by
29a, b) (with a question mark) as synonyms. De Toni (1905:          Vieillard # 1913a from Wagap, New Caledonia. These are the
1596) transferred Gymnogongrus ligulatus to the genus Po-           fragments Kützing used to prepare fig. 40e and thus should
lyopes (with a question mark) but incorrectly assigned the          be the holotype. They appear to be parts of what would have
authority as P. ligulatus (Harvey) J. Agardh, when in fact          been a larger thallus. De Toni (1897: 481) related this species
Kützing validly published the basionym Gymnogongrus ligu-          to the earlier described H. spinella (C. Agardh) Kützing
latus. Furthermore, De Toni (1905: 1573) related Grateloupia        (1847), but due to the many fine proliferous lateral branchlets
emarginata Kützing to Eucheuma gelatinum (Esper) J.                along the main axes, which are clearly visible under the mi-
Agardh, which is now Betaphycus gelatinum (Esper) Doty              croscope, we would suggest that it is conspecific with the later

542    Phycologia, Vol. 44 (5), 2005

described and widely reported Hypnea boergesenii Tanaka             Cribb (1958) placed this species as a synonym of Laurencia
(1941).                                                             concinna Montagne (1842), which in turn has been generally
   Hypnea vaga Kützing 1863a: 16. 1868: 8, table 23a, b.           considered a synonym of the widely distributed Laurencia
   Type is L 941,61-133 (barcode L 0055957) collected by            brongniartii J. Agardh (1841; Yamada 1931; Saito & Wom-
Vieillard # 1941 from Baláde, New Caledonia. Kützing (1868:       ersley 1974). The first author has collected this New Cale-
8) specifically cited specimen # 1941, and this is also the same    donian species many times along the eastern mainland coast
as used by Kützing to prepare fig. 23a. It is thus the holotype.   of Australia and has determined that it is a different species
A probable isotype is filed in MEL (unnumbered). Although           from Laurencia brongniartii based on several diagnostic mor-
seemingly a good species of Hypnea (cross sections reveal a         phological characters. This has also been confirmed by Pro-
central axial filament surrounded by pseudoparenchymatous           fessors Yuzuru Saito and Ki Wan Nam. We thus recognise
medullary cells) its affinities with other species are not yet      this species as distinct and a currently accepted name.
clear, and further study is required.                                  Laurencia decumbens Kützing 1863a: 18. 1865: 18, table
   Hypoglossum vieillardii Kützing 1863a: 19. 1866: 4, table       51a, b.
10a–c.                                                                 Type is L 941,119-50 (barcode L 0055982) collected by
   Type is L 935,329-1 (barcode L 0055716) collected by             Vieillard # 2033 from New Caledonia. This is most likely the
Vieillard # 2026 in 1863 from Wagap, New Caledonia. The             specimen Kützing used to prepare fig. 51a and is thus the
lower specimen labelled ‘Wagap’ appears to have been used           holotype. Possible isotypes are L 941,149-161. This is a cur-
by Kützing to prepare fig. 10a and is thus the holotype. Wynne     rently accepted name and species (Silva et al. 1996).
& Kraft (1985) and King & Puttock (1994) placed this species           Laurencia flagellifera Kützing 1865: 19, table 53c, d.
in synonymy with Caloglossa leprieurii (Montagne) J.                   Fig. 6
Agardh, including the homotypic synonym Delesseria vieil-              Type is L 934,263-8 (barcode L 0055983) collected by
lardii (Kützing) Zanardini. More recently, however, Kamiya         Vieillard # 1934 and # 2030 from New Caledonia. Three spec-
et al. (2003) justified the maintenance of this as a distinct       imens representing the same species make up this collection,
species known as Caloglossa vieillardii (Kützing) Setchell.        two marked # 1934 and one marked # 2030. The former were
   Iridaea irregularis Kützing 1867: 3, table 7d, e.               used by Kützing to prepare fig. 53c. Laurencia flagellifera
   Fig. 4                                                           Kützing is a later homonym of Laurencia flagellifera J.
   Type is L 941,119-12 (barcode L 0055963) collected by            Agardh (1852; type locality – India), which is the basionym
Vieillard # 2051 from Wagap, New Caledonia. The specimen            of Chondrophycus flagelliferus (J. Agardh) Nam (1999) and
appears to have been used by Kützing to prepare fig. 7d and        a very different species. The present taxon has secondary pit-
is thus the holotype. Although sterile, this specimen has a         connections between outer cortical cells and four pericentral
distinctive habit of coalescent blades and internally displays      cells per axial cell and thus is a Laurencia species. In overall
a large celled pseudoparenchymatous medulla. It is what is          habit, it shares similarities with Laurencia majuscula (Harvey)
currently known as Halichrysis coalescens (Farlow) Norris &         Lucas, but differs in being almost completely denuded of lat-
Millar (Norris 1991), which thus becomes a heterotypic syn-         eral branches along its main axes. It also looks similar to
onym of Halichrysis irregularis (Kützing) Millar comb. nov.        Lauencia obtusa var dendroidea J. Agardh, but is closely ad-
[Basionym: Iridaea irregularis Kützing 1867: 3, table 7d, e.]      herent to the paper and thus most likely flacid in texture.
as does Herpophyllon coalescens Farlow (1902: 97).                  Agardh’s variety has a woody texture (see below under L.
   Iridaea fimbriata Kützing 1867: 3, table 8a, b.                 vieillardii). Kützing’s species thus requires a new name: Lau-
   Fig. 5                                                           rencia kuetzingii Millar nom. nov. [Replaced name: Laurencia
   Type is L 941,119-10 (barcode L 0194034) collected by            flagellifera Kützing 1865: 19, table 53c, d.]
Vieillard # 2101a in 1863 from Wagap, New Caledonia. This              Laurencia thuyoides Kützing 1865: 26, table 74a, b.
specimen has been used by Kützing to prepare fig. 8a and is           The only sheet located was L 941,149-178 (ex herb. Len-
thus the holotype. It is a small, flat-bladed cystocarpic plant     ormand; barcode L 0194036) No locality is given, nor does it
of several lobes about 2 cm across. Its ampullate cystocarps        have any Kützing annotations, only those of Lenormand stat-
suggest the Halymeniaceae, and although the medulla is seem-        ing ‘Laurencia thuyoides Kütz’. It may well be from Vieil-
ingly densely packed (a result of the plant being pressed and       lard’s collections, but there is no real certainty. De Toni (1903:
not rehydrating well) and thus reminiscent of the genera Car-       788) and Yamada (1931) determined that this species was con-
popeltis or Polyopes, the occasional cross-linking medullary        specific with the earlier described Laurencia paniculata (C.
filaments joining either side of the blade are characters typical   Agardh) J. Agardh (1852) and this decision has modern ac-
of the genus Halymenia. Because of the existence of Haly-           ceptance (Boisset et al. 2000). Nam (1999) decided that L.
menia fimbriata Zanardini (1874: 503), the type of which,           paniculata was conspecific with the earlier Chondrophycus
from Lord Howe Island, actually represents a specimen of            patentirameus (Montagne) Nam. Based on type specimens,
Sarcodia ciliata Zanardini (1874: 498; G.T. Kraft, personal         Boisset et al. (2000) demonstrated that L. paniculata was in-
communication), this species will require a new name: Haly-         deed a species of Chondrophycus, proposed the new combi-
menia kuetzingii Millar nom. nov. [Replaced name: Iridaea           nation Chondrophycus paniculatus (C. Agardh) Furnari (plac-
fimbriata Kützing 1867: 3, table 8a, b.]                           ing Laurencia thuyoides as a synonym), but determined that
   Laurencia calliptera Kützing 1865: 24, table 69c, d.            C. patentirameus and C. paniculatus are in fact two separate
   Type is L 941,119-53 (barcode L 0194035) collected by            species. Furnari et al. (2001), realising that C. paniculatus was
Vieillard # 1932 from New Caledonia. This is the specimen           illegitimate as its basionym (Chondria obtusa var paniculata)
used by Kützing to prepare fig. 69c and is thus the holotype.      had no priority outside its rank, proposed the new combination
J. Agardh (1876: 661) and later De Toni (1903: 806) and             Chondrophycus thuyoides (Kützing) Furnari. Liao et al.(2004)

Millar & Prud’homme van Reine: Vieillard marine algae from New Caledonia 543

recently reported this species from the Anambas and Natuna specific, Kützing’s name would have priority, but the New
Island in Indonesia. Caledonian species is more likely a Chondria. Sections reveal
Laurencia vaga Kützing 1863a: 18. 1865: 18, table 50a–c. what seems to be a slightly darker staining central axial cell,
Type is L941,119-132 (barcode L 0055999) collected by typical of Chondria species, but until spermatangial plants are
Vieillard # 12b from New Caledonia. The several specimens collected, the generic placement of this species must remain
in the envelope on this sheet were used by Kützing to prepare in doubt.
fig. 50a, c. Laurencia vaga Kützing is a later homonym of Liagora tomentosa Kützing 1863a: 13.
Laurencia vaga Zanardini (1851), which in turn is the bas- Type is L 941,149-236 (barcode L 0056005) collected by
ionym of Alsidium vagum (Zanardini) Zanardini (see De Toni Vieillard from New Caledonia. This species is being critically
1903: 868), and a very different taxon to Kützing’s species. examined separately (J.M. Huisman & A.J.K. Millar, unpub-
De Toni (1903: 807) and Yamada (1931) determined that lished observations). Isotype on sheet L 940,284-299 (barcode
Kützing’s species was conspecific with the earlier described L 0055801) with the holotype of Gongroceras subtile Kütz-
Laurencia perforata (Bory de Saint-Vincent) Montagne ing.
(1840), and this decision has modern acceptance (Cribb 1983; Lophura armata Kützing 1866: 2, table 3a, b.
Price & Scott 1992). Laurencia perforata, the type locality for Type is L 940,284-299 (barcode L 0055801) collected by
which is the Canary Islands, is a very widespread species and Vieillard # 2089 in 1863 from Wagap, New Caledonia. This
has been recorded from the western Indian Ocean (Silva et al. single specimen has been used by Kützing to prepare fig. 3a
1996), eastern Atlantic Ocean, and the western Pacific. Nam and is therefore the holotype. Isotypes on L 040,284-300. This
(1999), without reference to whether he examined type spec- is the basionym of the currently accepted name and species
imens, proposed the new combination Chondrophycus perfor- Chondria armata (Kützing) Okamura (1907).
ata (Bory) Nam, presumably because descriptions of this spe- Mastocarpus marginalis Kützing 1863a: 15. 1867: 11, ta-
cies invariably mention the lack of secondary pit connections ble 39a, b.
between outer cortical cells that are radially elongated and Fig. 7
palisade-like. Examination of Kützing’s specimens revealed Type is L941,149-56 (barcode L 0056014) collected by
plants with two pericentral cells per axial cell and outer cor- Vieillard # 2025 from New Caledonia. This single specimen
tical cells that are palisade-like, radially elongated and lacking was used by Kützing to prepare fig. 39a and is thus the ho-
secondary pit connections between them. Such characters are lotype. The type specimen is cystocarpic, the cystocarps borne
typical of the genus Chondrophycus, and we would agree with only on the margins of the flattened, dichotomously to divar-
the above-mentioned synonymy. The possible conspecificity icately branched, smooth surfaced blades. The anatomy of the
of Laurencia perforata and the New Caledonian Laurencia cystocarps is typical of Sarcodia species with its basal retic-
decumbens Kützing (see above) has often been debated (J. ulum of small cells and massive, thick pericarp. The vegeta-
Agardh 1876; Yamada 1931; Saito 1969), especially as both tive sections display a filamentous medulla and pseudopar-
share a decumbent, tufted habit and occur in similar pan-trop- enchymatous inner cortex, again typical of the genus Sarco-
ical habitats. They are now considered to be in two different dia. There is a very strong resemblance between this species
genera. and the southern Australian Sarcodia marginata J. Agardh
Laurencia vieillardii Kützing 1865: 17, table 45a, b. (1892) as illustrated by Womersley (1994, fig. 101A–D).
Type is L 941,119-51 (barcode L 0194037) collected by Kützing’s name would have priority. The South African en-
Vieillard # 1935 from Baláde, New Caledonia. This is the demic Sarcodia dentata (Suhr) R.E. Norris also shares the
specimen Kützing used to prepare fig. 54a, and is thus the flattened, dichotomously branched habit, but in that species
holotype. J. Agardh (1876: 650) listed this species (with a there are numerous ‘tubercules’ covering the female blade sur-
question mark) as a synonym of his earlier described Lauren- faces (Stegenga et al. 1997: 336, pl. 121, figs 1, 2), a feature
cia dendroidea J. Agardh (1852: 753) from Brazil. He also lacking in the New Caledonian species. Sarcodia marginalis
placed under that species (with no question mark), the Indo- (Kützing) Millar comb. nov. [Basionym: Mastocarpus mar-
Pacific Laurencia obtusa var. majuscula Harvey (1863: synop. ginalis Kützing 1863a: 15].
309b). Harvey’s variety was later recognised at species rank Melanthalia fastigiata Kützing 1869: 16, table 44a, b.
as Laurencia majuscula (Harvey) Lucas (1935), which is a Type is L 941, 149-79 (barcode L0194083) collected by
currently accepted name and species (Womersley 2003: 457) Vieillard from Ile des Pins, New Caledonia. This specimen is
that is distinct from the Atlantic species. Laurencia dendro- clearly the one used by Kützing to prepare fig. 44a and is thus
idea is generally considered to be a variety of Laurencia ob- the holotype. An isotype exists in MEL 697584. This species
tusa (Yamada 1931; Saito 1969). The type specimen of Lau- is conspecific with what is currently known as Melanthalia
rencia dendroidea J. Agardh (LD 36669) has a dark, woody concinna J. Agardh (1876) and thus has priority. Womersley
main axis and laterals that do not adhere well to the paper. It (1996: 37) noted the confusion that has existed between Me-
is very different from the type of Laurencia vieillardii that lanthalia concinna J. Agardh and Fucus concinnus R. Brown
has translucent axes and is closely adherent to the paper. Spec- ex Turner. The latter actually represents Trematocarpus con-
imens of Laurencia majuscula from southern and eastern Aus- cinnus (R. Brown ex Turner) De Toni (1900).
tralia share these translucent/adherent characters but are gen- Melanthalia vieillardii Kützing 1863a: 17. 1869: 16, table
erally much more densely branched than Vieillard’s specimen 42c, d.
is. Furthermore, the New Caledonian specimen has all the Type is L 941,149-80 (barcode L 0056017) collected by
characteristics of a small, delicate Chondria species and, in Vieillard # 2041 from New Caledonia. One of the two spec-
fact, is labeled as such by either Vieillard or Kützing. If Lau- imens has been used by Kützing to prepare fig. 42c and is
rencia vieillardii and Laurencia majuscula prove to be con- thus the holotype. This species is clearly conspecific with the

544    Phycologia, Vol. 44 (5), 2005

Figs 7–10. Kützing’s type specimens.
  Fig. 7. Mastocarpus marginalis (5 Sarcodia marginalis). Scale 5 20 mm.
  Fig. 8. Sphaerococcus angustifolius (5 Schottera angustifolia). Scale 5 20 mm.
  Fig. 9. Sphaerococcus caulescens (5 Rhodymenia caulescens). Scale 5 20 mm.
  Fig. 10. Sphaerococcus chondracanthus (5 Gracilaria chondracantha). Scale 5 20 mm.

earlier described Melanthalia obtusata (Labillardière) J.             Type is L 941,181-434 (barcode L 0061144) collected by
Agardh (1852; Womersley 1996).                                      Vieillard # 2114 in 1863 from Wagap, New Caledonia. This
   Nemalion filicoides Kützing 1866: 23, table 65a, b.             is the specimen used by Kützing to prepare fig. 90c and is
   Type is L 941,181-267 (barcode L 0056029) collected by           thus the holotype. It is a currently accepted name and species
Vieillard # 2094bis in 1863 from Wagap, New Caledonia. This         (Denizot 1968).
is the specimen used by Kützing to prepare fig. 65a and thus          Plocamium botryoides Kützing 1866: 18, table 50a–c.
is the holotype. It bears a striking resemblance to the more           Type is L941,240-49 (barcode L 0056037) collected by
recently described Ganonema samaense (Tseng) Huisman                Vieillard # 1930 from Wagap, New Caledonia. Kützing spe-
(2002), and the specimen is being critically examined sepa-         cifically cited Vieillard # 1930, and two different specimens
rately (Huisman & Millar, unpublished observations).
                                                                    representing the same species have been used to prepare fig.
   Nothogenia livida Kützing 1869: 17, table 46e, f.
                                                                    50a–c. De Toni (1897: 602) related this species to Plocamium
   Type is L 938,303-141 (barcode L 0194038) collected by
                                                                    abnorme Hooker & Harvey (Hooker 1845), which was shown
Vieillard # 2099 in 1863 from Wagap, New Caledonia. The
specimen has been used by Kützing to prepare fig. 46e and is       to be a synonym of Plocamium angustum (J. Agardh) Hooker
thus the holotype. De Toni (1897: 121) listed this species as       & Harvey (Hooker 1847) by South & Adams (1979). This
uncertain, and it appears not to have been mentioned in the         species looks very similar to, if not conspecific with, the later
literature since. The specimens are seemingly identical with        described Plocamium hamatum J. Agardh (1876: 338), which
what Coppejans & Millar (2000, fig. 8) identified as Carpo-         was originally described from nearby Norfolk Island. Al-
peltis maillardii (Montagne and Millardet) Chiang (1970)            though the New Caledonian plant lacks the characteristic
from nearby Papua New Guinea.                                       hooks of that species, specimens of P. hamatum collected by
   Peyssonnelia neocaledonica Kützing 1869: 32, table 90c, d.      the first author from Norfolk Island often lack these.

Millar & Prud’homme van Reine: Vieillard marine algae from New Caledonia 545

Polysiphonia pauperula Kützing 1863a: 17. 1863b: 20, ta- cellular anatomy is typical for species of the genus Rhody-
ble 62a, b. menia. With a distinctive stipe and regularly dichotomously
Type is L 941,253-168 (barcode L 0056093) collected by branched blades, the species would presently be identified
Vieillard # 1948 from Yaté, New Caledonia. Kützing specif- with the eastern and southern Australian species Rhodymenia
ically cited Yaté as the locality in the protologue so this spec- sonderi Silva (see Silva et al. 1996). Silva (in Silva et al.
imen is the holotype. Another isotype is L 941,253-141, but 1996) proposed the new name sonderi because the intended
a specimen in MEL (663746) was collected from Wagap and basionym of Rhodymenia australis Sonder was a superfluous
is thus not a type. King & Puttock (1989: 31) reduced this and illegitimate name, and there were no apparent synonyms
species to a heterotypic synonym of Bostrychia moritziana available. Kützing’s name is available for this taxon, and the
(Sonder ex Kützing) J. Agardh (1863). new combination is here effected Rhodymenia caulescens
Polysiphonia polyphysa Kützing 1863a: 18. 1863b: 20, ta- (Kützing) Millar comb. nov. [Basionym: Sphaerococcus cau-
ble 62c, d. lescens Kützing 1868: 32, table 92c, d.]
Type is L 941,253-167 (barcode L 0056094) collected by Sphaerococcus chondracanthus Kützing 1863a: 17.
Vieillard # 89c from New Caledonia. It is not clear if the Fig. 10
material has been used by Kützing to prepare his fig. 62c, but Type is L 941,61-74 (barcode L 0194042) collected by
it is the only specimen located and thus the holotype. The Vieillard # 17.6 from New Caledonia. This is the only spec-
species was recently rediscovered from Papua New Guinea imen in the collection and is thus the holotype. Kützing
(Coppejans & Millar 2000). It is a currently accepted name (1863a) related this species to the western Atlantic Sphaero-
and species. coccus cervicornis, and the specimen was filed under the
Sphaerococcus angustifolius Kützing 1868: 34, table 99e–i. name Gracilaria cervicornis (Turner) J. Agardh (1852), which
Fig. 8 is a western Atlantic and western Indian Ocean species. The
Type is L 941,311-34 (barcode L 0194039) collected by name, S. chondracanthus, has not been cited since its proto-
Vieillard from New Caledonia. The single envelope in this logue. The New Caledonian specimen is a species of Graci-
collection contains several small plants, and these have been laria and not Chondracanthus, as cross sections reveal a cel-
used by Kützing to prepare fig. 99e–g. De Toni (1900: 397) lular medulla of large cells grading to a small celled cortex.
related this species to the genus Phyllophora, and Garrigue & Chondracanthus has a medulla of loosely entangled filaments.
Tsuda (1988) suggested this represents Gelidium samoense Based on Turner’s description and illustrations (1808–1809,
Reinbold. Kützing’s specimens are sterile, but the anatomy of pl. 121) of Fucus cervicornis, the two species are not the
cross sections of the branches lack rhizines, have a small- same. Kützing’s species looks remarkably similar to, and is
celled medulla and a thick cortex consisting of catenate files most likely conspecific with the western Pacific G. bangmei-
of four to five cells, and strongly suggest the family Phyllo- ana Zhang & I.A. Abbott in Abbott et al. (1991), which has
phoraceae as De Toni predicted. The Rhodymeniaceae, espe- been recorded from Hainan Island (China), Norfolk Island,
cially Rhodymenia species, have a thin cortex and mostly a and the east Australian mainland (Millar & Xia 1999). Both
large-celled medulla, and in our opinion are quite different. species possess characteristic short spine-like branchlets with
The habit of the New Caledonian species consists of branched acute apices. Kützing’s name has priority: Gracilaria chon-
flattened, ligulate fronds arising from a slender terete, seem- dracantha (Kützing) Millar comb. nov. [Basionym: Sphaero-
ingly creeping stolon and match well with the genus Schottera coccus chondracanthus Kützing 1863a: 17.]
Guiry & Hollenberg (1975). The only species in that genus, Sphaerococcus denticulatus Kützing 1869: 19, table 51e–g.
Schottera nicaeensis (Lamouroux ex Duby) Guiry & Hollen- Type is L938,92-252 (barcode L 0055699) collected by
berg (see Womersley 1994: 263, fig. 83C–F) consists of much Vieillard # 2089bis from Wagap, New Caledonia. This in-
larger plants that regularly have flattened blades with thin, cludes a specimen used by Kützing to prepare fig. 51e and is
terete, stolon-like apices as well as bases. Schottera angusti- thus the lectotype. This name was the intended basionym for
folia (Kützing) Millar comb. nov. [Basionym: Sphaerococcus Gracilaria denticulata (Kützing) Weber-van Bosse (1928),
angustifolius Kützing 1868: 34, table 99e–i.] which is a later homonym of Gracilaria denticulata Schmitz
Sphaerococcus canaliculatus Kützing 1868: 29, table 82d, ex Mazza (1907). This species is currently known as Graci-
e. nom. illeg. (see Silva et al. 1996). Type is L 941,61-93 laria vieillardii Silva (Silva et al. 1987: 44; Withell et al.
(barcode L 0194040) collected by Vieillard # 2128 in 1863 1994: 309).
from Wagap, New Caledonia. Kützing specifically cited the Sphaerococcus lemania Kützing 1868: 26, table 75a–c.
specimen from Wagap and used this specimen to prepare fig. Type is L 941,61-61 (barcode L 0194043) collected by
82d. It is thus the holotype. This now represents Gracilaria Vieillard # 2028bis from Wagap, New Caledonia. This is the
canaliculata Sonder (Withell et al. 1994: 301. Silva et al. specimen used by Kützing to prepare the fig. 75a and is thus
1996: 157). the holotype. May (1948) reduced this species to a forma of
Sphaerococcus caulescens Kützing 1868: 32, table 92c, d. Gracilaria lichenoides Greville (1830). The New Caledonian
Fig. 9 species appears to be conspecific with and thus a heterotypic
Type is L 941,311-36 (barcode L 0194041) collected by synonym of the earlier Gracilaria edulis (S. Gmelin) Silva
Vieillard # 1928 in 1861 from Ile des Pins, New Caledonia. (1952).
One of the three samples has been used by Kützing to prepare Sphaerococcus setaceus Kützing 1868: 26, table 73a.
fig. 92c and thus is the holotype. De Toni (1900: 398) related Type is L 941,61-95 (barcode L 0194044) collected by
this species to Rhodymenia flabellifolia (Bory) Montagne Vieillard # 2028 from Wagap, New Caledonia. Of the three
(1846) but did not place it in synonymy under that taxon’s specimens in this collection, one has been used by Kützing to
entry. Cross-sections of Kützing’s specimen confirm that the prepare fig. 73a and is thus the holotype. De Toni (1900) said

546 Phycologia, Vol. 44 (5), 2005

that he had seen this material and suggested it had the struc- with Professor Claude Payri, Université de Polynésie française.
ture of Gracilaria. We would agree and further suggest that A.J.K.M. sincerely thanks Professor Payri for reigniting the
it represents Gracilaria edulis (S. Gmelin) Silva (1952). idea of completing this project. The authors sincerely thank
Sphaerococcus spinescens Kützing 1868: 27, table 79a, b. Per Lassen from the Botanical Museum in Lund, Sweden, for
Type is L941,51-64 (barcode L 0055916) collected by information regarding Agardh’s type specimens. Professor
Vieillard # 1924 from New Caledonia. Kützing (1868: 28) Paul Silva (University of California, Berkeley) generously
specifically cited Vieillard # 1924 and used this specimen to helped with nomenclatural advice, and John Lewis (University
prepare fig. 79a. It is thus the holotype. This is the basionym of Melbourne), Dr John Huisman (Murdoch University), Dr
of Gracilaria spinescens (Kützing) J. Agardh (1876), which Frederik Leliaert (University of Gent) and Drs Giovanni Fur-
is now considered conspecific with the earlier Gracilaria ed- nari and Mario Cormaci (Università di Catania) offered their
ulis (S. Gmelin) Silva (Withell et al. 1994: 348). expertise on certain families and genera. Dr Michael Wynne
Sphaerococcus spinulosus Kützing 1868: 30, table 85a, b. made many valuable suggestions for improvement of the orig-
Type is L941,51-61 (barcode L 0056140) collected by inal manuscript.
Vieillard # 2118 in 1863 from Wagap, New Caledonia. This
is the only specimen in the collection, and Kützing used it to
prepare fig. 85a. It is thus the holotype. J. Agardh (1876: 424), REFERENCES
with a question mark, placed this species as a synonym of
Gracilaria corniculata (R. Brown ex Turner) J. Agardh (1852:
ABBOTT I.A., ZHANG J. & XIA B. 1991. Gracilaria mixta, sp. nov. and
595), which was described from Kent Island in Bass Strait, other western Pacific species of the genus (Rhodophyta, Gracilari-
and De Toni (1900: 451) later followed suit. Although Kütz- aceae). Pacific Science 45: 2–27.
ing’s specimen is sterile, it anatomically and morphologically AGARDH C.A. 1822. Species algarum, vol. 1, part 2a. Berling, Lund,
matches the distinctive (originally Japanese) species Hypnea Sweden. pp. 169–398.
saidana Holmes (1896: 256, pl. 11, fig. 3a, b) as it occurs in AGARDH J.G. 1837. Novae species algarum, quas in itinere ad oras
eastern Australia and Papua New Guinea (Millar 1990: Fig. marius Rubri collegit Eduardus Rüppell; cum observationibus non-
nullis in species rariores antea cognitas. Museum Senckenbergianum
23A; Millar et al. 1999; Coppejans & Millar 2000). Hypnea
2: 169–174.
saidana has also been recorded from the warmer waters of AGARDH J.G. 1841. In historiam algarum symbolae. Linnaea 15: 1–
the Indian Ocean (see Silva et al. 1996 for a range of refer- 50, 443–457.
ences) and the Philippines (Silva et al. 1987). Although Kütz- AGARDH J.G. 1851. Species genera et ordines algarum. Volumen se-
ing’s name is earlier, it cannot be used within the genus Hyp- cundum: algas florideas complectens, part 1. Gleerup, Lund, Swe-
nea as it would be a later homonym of Hypnea spinulosa den. pp. 1–351.
AGARDH J.G. 1852. Species genera et ordines algarum. Volumen se-
Lamouroux (1813). However, Lamouroux’s name is an in-
cundum: algas florideas complectens, part 2, pp. 337–720; part 3,
tended combination of Fucus spinulosus J.F. Gmelin (1792), pp. 701–786. Gleerup, Lund, Sweden.
which in turn was a superfluous name for Fucus spinosus S.G. AGARDH J.G. 1863. Species genera et ordines algarum. Volumen se-
Gmelin (1768), that is the basionym of Gelidium spinosum (S. cundum, part 3. Gleerup, Lund, Sweden. pp. 787–1291.
Gmelin) Silva (in Silva et al. 1996). Therefore the name Hyp- AGARDH J.G. 1876. Species genera et ordines algarum, Volumen ter-
nea saidana is available for this species. Turner’s (1809–1811, tium: de florideis curae posteriores. Epicrisis systematiis Flori-
pl. 182) description of Fucus corniculatus from Bass Strait dearum. Weigel, Leipzig, Germany. 724 pp.
AGARDH J.G. 1892. Analecta algologica. Lunds Universitets Års-Skrift,
mentioned a plant that did not adhere to paper and had a Andra Afdelningen, Kongeligen Fysiografiska sällskapets i Lund
smooth surface, features that do not match those of Kützing’s Handlingar, vol. 28(6): 1–182.
specimen of Sphaerococcus spinulosus from New Caledonia. ARDISSONE F. 1871. Revista dei Ceramii della flora Italiana. Nuova
Sphaerococcus vieillardii Kützing 1863a: 16. 1868: 26, ta- Giornale Botanico Italiano 3: 32–50.
ble 74c–e. BERGER S., FETTWEISS U., GLEISSBERG S., LIDDLE L.B., RICHTER U.,
Type is L 941,311-38 (barcode L 00556141) collected by SAWITZKY H. & ZUCCARELLO G. 2003. 18S rDNA phylogeny and
evolution of cap development in Polyphysaceae (formerly Aceta-
Vieillard # 1925 from Ile des Pins, New Caledonia. There are
bulariaceae; Dasycladales, Chlorophyta). Phycologia 42: 506–561.
two specimens and one has been used by Kützing to prepare BOISSET F., FURNARI G., CORMACI M. & SERIO D. 2000. The distinction
fig. 74c and is thus the holotype. An isotype is L 941,311-40. between Chondrophycus patentirameus and C. paniculatus (Cera-
J. Agardh (1876: 426) did not see this specimen but com- miales, Rhodophyta). European Journal of Phycology 35: 387–395.
mented that Kützing’s illustrations suggested that it was more BøRGESEN F. 1932. A revision of Forsskål’s algae mentioned in Flora
like a Dicranema. The habit of this species is very much that Aegyptiaco-Arabica and found in his herbarium in the Botanical
of a Gracilaria, and the anatomy of a cross-section of the Museum of the University of Copenhagen. Dansk Botanisk Arkiv
8: 1–14, 4 figs, 1 pl.
branches also suggests this genus. On habit alone, this species BRAND F. 1904. Über die Anheftung der Cladophoraceen und über
would appear to be representative of the earlier described Gra- verschiedene polynesische Formen dieser Familie. Beihefte zum Bo-
cilaria edulis (S. Gmelin) Silva (1952). tanischen Centralblatt 18: 165–193, pls V, VI.
CHIANG Y.-M. 1970. Morphological studies of red algae of the family
Cryptonemiaceae. University of California Publications in Botany
ACKNOWLEDGEMENTS 58: 1–83, pls 1–10.
COPPEJANS E.G.G. & MILLAR A.J.K. 2000. Marine red algae from the
north coast of Papua New Guinea. Botanica Marina 43: 315–346.
Although the idea behind this manuscript was borne several CRIBB A.B. 1958. Records of marine algae from South-Eastern
decades ago, the French Government (via the French Embassy Queensland – III. Laurencia Lamx. Papers of Department of Bot-
in Canberra, Australia) generously funded the 2002 collecting any, University of Queensland 3(19): 159–191.
expedition to New Caledonia by A.J.K.M., in collaboration CRIBB A.B. 1983. Marine algae of the southern Great Barrier Reef

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