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Notes on the samphires (Salicornioideae, Chenopodiaceae–Amaranthaceae) in Madagascar and Europa Island, with further conclusions on their chorology in Africa

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The last treatment of Chenopodiaceae in Madagascar was published in 1954 and since then, the generic circumscription of Chenopodiaceae including Salicornioideae has changed drastically. The Madagascan Salicornioideae (samphires) are still insufficiently studied. Salicornia pachystachya described from Madagascar had a turbulent nomenclatural and morphological history, frequently considered as Arthrocnemum pachystachyum due to the alleged similarity of the seed characters with other Arthrocnemum species, or considered in a broader sense including the related Salicornia perrieri. However, the reproductive features of S. pachystachya confirm its placement within Salicornia, and the thick inflorescences and longer seeds distinguish it from S. perrieri, another native Madagascan species. Two other members of Salicornioideae in Madagascar are the subshrubby Salicornia mossambicensis, erroneously named S. fruticosa or Arthrocnemum natalense, and Tecticornia indica. Herein, a diagnostic key is provided for the delimitation of four samphire species in Madagascar, and the lectotypes of Salicornia pachystachya and S. perrieri are designated. Only two species, S. pachystachya and Tecticornia indica, are recorded for Europa Island, and their habitat preferences in this location are presented, as well as a discussion of the species composition and chorology of samphires in continental Africa
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Notes on the samphires (Salicornioideae, Chenopodiaceae
Amaranthaceae) in Madagascar and Europa Island, with further
conclusions on their chorology in Africa
Заметки о солеросах (Salicornioideae, ChenopodiaceaeAmaranthaceae)
Мадагаскара и острова Европа, с заключениями об их хорологии в Африке
Alexander P. Sukhorukov1,2 *,
Maria A. Kushunina3, Vitaly Yu. Alyonkin4,
Jean Hivert5, Vincent Boullet6
1 Lomonosov Moscow State University, Biological Faculty,
Department of Higher Plants
Leninskie Gory, 1/12, Moscow, 119234, Russia
2 Tomsk State University, P. N. Krylov Herbarium
Lenina Ave., 36, Tomsk, 634050, Russia
suchor@mail.ru
3 Lomonosov Moscow State University, Biological Faculty,
Department of Plant Physiology
Leninskie Gory, 1/12, Moscow, 119234, Russia
mkushunina@gmail.com
4 The Apothecaries’ Garden, Botanic Gardens of Moscow State
University
Mira Ave., 26/1, Moscow, 129090, Russia
boraginaceae@mail.ru
5 Conservatoire Botanique National de Mascarin
2 rue du Père Georges, Les Colimaçons, 97436, Saint-Leu, île de
La Réunion, France
jhivert@cbnm.org
6 Universite de Bretagne Occidentale, EA 7462, Geoarchitecture,
29200 Brest, France;
Vourlhat, 43230, Frugières-le-Pin, France
v.boullet43@orange.fr
*Corresponding author
А. П. Сухоруков1,2 *, М. А. Кушунина3,
В. Ю. Алёнкин4 , Ж. Ивер5, В. Булле6
1 Московский государственный университет имени
М. В. Ломоносова, биологический факультет, кафедра
высших растений
Ленинские горы, 1/12, Москва, 119234, Россия
2 Томский государственный университет, Гербарий
им. П. Н. Крылова
пр. Ленина, 36, Томск, 634050, Россия
suchor@mail.ru
3 Московский государственный университет имени
М. В. Ломоносова, биологический факультет, кафедра
физиологии растений
Ленинские горы, 1/12, Москва, 119234, Россия
mkushunina@gmail.com
4 Ботанический сад Московского государственного
университета имени М.В.Ломоносова «Аптекарский Огород»
пр. Мира, 26/1, Москва, 129090, Россия
boraginaceae@mail.ru
5 Национальный ботанический институт изучения
Маскаренских островов
ул. Пэр Жорж, 2, Ле Колимасон, 97436, Сен-Ле, о.Реюньон,
Франция
jhivert@cbnm.org
6 Университет Западной Бретани, EA7462, департамент
геоархитектуры, Брест, 29200, Франция;
Вурла, Фрюжьер-ле-Пен, 43230, Франция
v.boullet43@orange.fr
*Автор для переписки
https://doi.org/10.31111/novitates/2021.52.38
Abstract. The last treatment of Chenopodiaceae in Madagascar was published in 1954 and since then, the generic circumscrip-
tion of Chenopodiaceae including Salicornioideae has changed drastically. The Madagascan Salicornioideae (samphires) are still
insuffi ciently studied. Salicornia pachystachya described from Madagascar had a turbulent nomenclatural and morphological
history, frequently considered as Arthrocnemum pachystachyum due to the alleged similarity of the seed characters with other
Arthrocnemum species, or considered in a broader sense including the related Salicornia perrieri. However, the reproductive fea-
tures of S. pachystachya confi rm its placement within Salicornia, and the thick infl orescences and longer seeds distinguish it
from S.perrieri, another native Madagascan species. Two other members of Salicornioideae in Madagascar are the subshrubby
Salicornia mossambicensis, erroneously named S.fruticosa or Arthrocnemum natalense, and Tecticornia indica. Herein, a diagnos-
tic key is provided for the delimitation of four samphire species in Madagascar, and the lectotypes of Salicornia pachystachya
and S.perrieri are designated. Only two species, S.pachystachya and Tecticornia indica, are recorded for Europa Island, and their
habitat preferences in this location are presented, as well as a discussion of the species composition and chorology of samphires
in continental Africa.
Keywords: Salicornia, Tecticornia, Arthrocnemum, Madagascar, Europa Island, taxonomy.
Поступила в редакцию | Submitted: 05.05.2021
A. P. Sukhorukov et al.
Принята к публикации | Accepted: 15.10.2021
Новости систематики высших растений
Novitates Systematicae Plantarum Vascularium
2021
52: 38–52
ISSN 0568-5443 (print)
ISSN 2687-1564 (online)
Аннотация. Последняя обработка маревых Мадагаскара опубликована в 1954 году. С тех пор родовой состав как
Chenopodiaceae в целом, так и Salicornioideae претерпел существенные изменения. Мадагаскарские Salicornioideae все
еще недостаточно изучены. Описанный с Мадагаскара вид Salicornia pachystachya имеет сложную номенклатурную
и морфологическую историю. Его часто рассматривали как Arthrocnemum pachystachyum по причине поверхностно-
го сходства признаков семян с другими видами Arthrocnemum или объединяли с другим аборигенным мадагаскарским
видом — Salicornia perrieri. Репродуктивные признаки свидетельствуют о принадлежности S. pachystachya именно к
роду Salicornia; при этом он отличается от S. perrieri мясистыми соцветиями и более крупными семенами. Остальные
два вида солеросов на Мадагаскаре — полукустарнички Salicornia mossambicensis (ошибочно называемый S. fruticosa
или Arthrocnemum natalense) и Tecticornia indica. Внастоящей работе обозначены лектотипы Salicornia pachystachya и
S.perrieri; приведен ключ для определения четырех солеросов, произрастающих на Мадагаскаре. На острове Европа об-
наружены только два вида: S.pachystachya и Tecticornia indica, приводятся описания их предпочтительных местообита-
ний на этом атолле. Также обсуждаются видовой состав и хорология Salicornioideae в континентальной Африке.
Ключевые слова: Salicornia, Tecticornia, Arthrocnemum, Мадагаскар, остров Европа, таксономия.
The last treatment of Chenopodiaceae Vent. in Ma-
dagascar included eleven species belonging to seven
genera (Cavaco, 1954). In this treatment, Chenopodi-
umL. is the only species-rich genus (5 spp.), other gen-
era (Arthrocnemum Moq., Atriplex L., Beta L., Salicor-
niaL., SalsolaL., and Suaeda Forssk. ex J. F. Gmel.) are
monospecifi c. Out of eleven species, four members (Ar-
throcnemum pachystachyum (Bunge ex Ung.-Sternb.)
A. Chev., Atriplex perrieri Leandri, Salicornia perrieri
A.Chev. and Salsola littoralis Moq.) are considered en-
demic to Madagascar (Cavaco, 1954). Suaeda monoica
Forssk. ex J. F. Gmel. is reported to be native, and the
island is the southernmost part of its wide distribu-
tion in tropical East Africa and Western Asia (Brenan,
1954a; Hedge, 1997; Sukhorukov et al., 2016).
Recently, the taxonomic status of many Chenopo-
diaceae genera has changed after extended molecular
studies. According to the latest phylogenetic revisions
(Fuentes-Bazan et al., 2012; Sukhorukov et al., 2018b;
Uotila et al., 2021), Chenopodioideae Burnett comprise
several genera with diff erent positions in the molecular
trees. In Madagascar, Chenopodium in a previous broad-
er sense is represented by Chenopodium s.str. (C.album
L. and C.amaranticolor H.J.Coste et Reyn.), Chenopo-
diastrum S. Fuentes, Uotila et Borsch (C. murale (L.)
S. Fuentes, Uotila et Borsch), and Dysphania R. Br.
(D.ambrosioides (L.) Mosyakin et Clemants, incl. Che-
nopodium integrifolium Vorosch.). Furthermore, accord-
ing to an extensive phylogenetic analysis of Salsoloideae
Raf. by Akhani et al. (2007), a part of Salsola is cur-
rently considered within Caroxylon Thunb.. Madagas-
can Salsola littoralis was not included in the phylogeny,
but it shares all the principal morphological characters
with other species of Caroxylon and should be regarded
as C. littorale (Moq.) Akhani et Roalson (Akhani et al.,
2007; Mucina, 2017). Europa Island (French South-
ern and Atlantic Lands) located in the Mozambique
Channel about 300 km west of Madagascar hosts sev-
eral Chenopodiaceae species, namely Caroxylon littorale,
Notes on the samphires (Chenopodiaceae–Amaranthaceae) in Madagascar and Europa Island
Salicornia pachystachya Bunge ex Ung.-Sternb., Suaeda
monoica, and Tecticornia indica (Willd.) K. A. Sheph. et
Paul G. Wilson (Boullet et al., 2018). Recently, C.litto-
rale was also recorded in the coastal parts of Inhambane
Province (Mozambique) as a native species (Friis, Holt,
2016). Based on these records, C.littorale and Salicornia
pachystachya cannot be considered as Madagascan en-
demics.
The taxonomy of Salicornioideae Luerss., also known
as samphires, has undergone drastic changes after mo-
lecular phylogenetic studies. All perennial Sarcocor-
nia A.J.Scott were nested within Salicornia (Kadereit
et al., 2006) and later transferred into the latter genus
(Steff en et al., 2015; Piirainen et al., 2017). Members of
the Australian Halosarcia Paul G.Wilson, Pachycornia
Hook. f., Sclerostegia Paul G. Wilson, Tegicornia Paul
G. Wilson and Tecticornia Hook. f. form a large clade
(Shepherd et al., 2004, 2005b), and they were united
into Tecticornia, the oldest generic name for this group
of genera (Shepherd, Wilson, 2007). The diversity of
Arthrocnemum in West Africa and further nomencla-
tural problems in this genus were discussed by Sukho-
rukov and Nilova (2016). Simultaneously, Piirainen et
al. (2017) described two new genera, of which Arthro-
caulon Piirainen et G.Kadereit was a replacement name
for Arthrocnemum.
Two reported Madagascan members of Salicornioi-
deae are still poorly investigated. The description of
the fi rst species, Salicornia pachystachya, was based
on a fragment of the plant collected by L. H. Boivin
in Madagascar. The species is considered as an an-
nual plant related to other Salicornia species (Ungern-
Sternberg, 1866). The name S. pachystachya had been
widely accepted (e. g., Hemsley, 1919; Toelken, 1967;
Scott, 1977; Piirainen et al., 2017). Chevalier (1922)
changed the generic name into Arthrocnemum pachys-
tachyum (Bunge ex Ung.-Sternb.) A. Chev. based on
the presence of perisperm in the ripe seeds, a charac-
ter lacking in other Salicornia. Simultaneously, he de-
Notes on the samphires (Chenopodiaceae–Amaranthaceae) in Madagascar and Europa Island
Novitates Systematicae Plantarum Vascularium | Volume 52 | 2021
39
scribed another Salicornia from Madagascar, S.perrieri
A.Chev. Cavaco (1954), following Chevalier, accepted
two Salicornioideae in Madagascar, Salicornia per-
rieri and Arthrocnemum pachystachyum, and corrected
the life form of the latter species to be shrubby. Both
names were also accepted by Moss (1954). Toelken
(1967) synonymised Salicornia perrieri with S.pachys-
tachya, while Bigot (1971) and O’Callaghan (1992)
considered both species as independent taxa imitating
“robust shrublets”. Two Arthrocnemum species, A.indi-
cum (Willd.) Moq. and A. natalense (Bunge ex Ung.-
Sternb.) Moss, were also reported in Madagascar only
several decades ago (Bigot, 1971; Duranton, 1975;
Thomasson, Théodore, 1993; Koechlin et al., 1997).
The analysis of the type material of Salicornia
pachystachya and S. perrieri, as well as the revision of
other herbarium material, led us to a distinct conclusion
on the taxonomy of the Madagascan samphires,
therefore this study aims to precise generic composition
of the samphires in Madagascar and Europa atoll, with
further notes on Salicornioideae in Africa.
Material and methods
The fi eld observations in southwest Madagascar
were made by V.Alyonkin in January 2020. The fi eld
observations in the Europa atoll were essentially
made by Vincent Boullet and Jean Hivert between
2006 and 2016 with several expeditions organised by
the Conservatoire Botanique National de Mascarin
(CBNM) to study the fl ora and plant communities.
The Scattered Islands (Fr.: les îles Éparses)
comprise fi ve islands located in the southwest Indian
Ocean, close to Madagascar, four in the Mozambique
Channel (cited here from south to north): Europa,
Bassas da India, Juan de Nova and the Îles Glorieuses,
while the fi fth, Tromelin, is located in the northern
part of the Mascarene Archipelago. All these islands
are relatively young, about 125,000 years old (Caceres,
2003) and administratively belong to the 5th district
of the French Southern and Antarctic Lands (Terres
Australes et Antarctiques Françaises, or TAAF), a
French Oversea Territory since 2007 (TAAF, 2016,
2021). The native Chenopodiaceae are only present
in the Europa atoll, not in the other Scattered Islands
(Boullet et al., 2018). The ecology of samphire
communities on the island has been specially studied
and described by Boullet (2014). The Europa atoll
is located at a low elevation (up to 12 m). Its tropical
climate is semiarid with two prevailing seasons, a dry
and relatively cool season (from May to November),
and a warm humid season (from December to April)
(Boullet et al., 2018). The island is located in the zone
of formation and circulation of tropical depressions
and cyclones (Delépine et al., 1976; Caceres, 2003)
and consists of a coral reef with a reduced terrestrial
area (30 km2) built on an ancient volcanic cone. It
is a remnant of old intra-plate volcanoes, nowadays
morphologically present in the form of atolls (Goldberg,
2016).
The ecology of some samphires was studied by
A.Sukhorukov (unpubl. data) in several parts of Africa
(Cape Verde, South Africa, and Namibia) and the type
material of the Salicornia species was investigated in the
Herbarium of Natural History Museum, Paris, France
(P). The analysis of other collections (BM, BR, K, LE)
housing abundant material from other African countries
was used for comparison.
The images of the seed ultrasculpture were taken
using a scanning electron microscope (SEM) JSM-6380
(JEOL Ltd., Japan) at 15 kV. Before SEM observations,
the seeds were soaked in a series of solvents and dried at
the critical point (see for more Sukhorukov, 2014). The
maps were prepared using the online tool SimpleMappr
(Shorthouse, 2010).
Results
Diversity of samphires in Madagascar and the Europa
Island
Our investigations show that the data on the di-
versity of the Salicornioideae in Madagascar should be
corrected. Four samphire species present in Madagas-
car are three Salicornia members (S. mossambicensis,
S.pachystachya, S. perrieri) and Tecticornia indica. Two
species, Salicornia pachystachya and S. perrieri, are an-
nuals, whereas S.mossambicensis and T.indica are pros-
trate or ascending subshrubs. Salicornia pachystachya
and S.perrieri diff er in seed length, with S.pachystachya
having longer (1.2–1.4 mm) seeds than those of S.perri-
eri (0.7–0.9 mm) (Fig. 1: A, B). Besides, the dried infl o-
rescences of S.pachystachya are thicker, reaching ca.6
mm diam., compared to those of S.perrieri (less than 3
mm across). The third samphire species is a subshrub-
by Salicornia mossambicensis with papillate seeds (Fig.
1: C, D). The fourth species, previously confused with
S.pachystachya, is Tecticornia indica, another subshrub
with distinctly smooth or slightly rugose seeds (Fig.
1: E, F). Only Salicornia pachystachya and Tecticornia
indica are present on Europa Island. A checklist of all
Salicornioideae found in the territory under study is
provided below.
Taxonomic account
Salicornia pachystachya Bunge ex Ung.-Sternb.
1866, Vers. Syst. Salicorn.: 51. Arthrocnemum pachy-
stachyum (Bunge ex Ung.-Sternb.) A.Chev. 1922, Rev.
Bot. Appl. Agric. Colon. 2 (16): 748. Arthrocnemum
A. P. Sukhorukov et al.
Новости систематики высших растений | Том 52 | 2021
40
Fig. 1. SEM micrographs of samphire seeds.
A— Salicornia pachystachya; B— S. perrieri; C, D— S. mossambicensis; E, F— Tecticornia indica. Scale bars: A–C—
100 μm, D— 50 μm, E, F— 200 μm.
Origin of the material: A: Madagascar, Baie de Diego-Suarez, II 1848, L.H.Boivin, №2370 (P); B: Madagascar, Anony
Lake, 26 II 1931, M.Decary, №8569 (BM); C, D: Madagascar, delta of the Linta River, VIII 1928, H.Humbert,
C.F.Swingle, № 5450 (K); E: Madagascar, near Toliara, IX 1924, H.Humbert, H.Perrier de la Bathie, №2478
(P04618457); F: Angola, Mossamedes [Namibe], F.Texeira, №2059 (BM).
Notes on the samphires (Chenopodiaceae–Amaranthaceae) in Madagascar and Europa Island
Novitates Systematicae Plantarum Vascularium | Volume 52 | 2021
41
pachystachyum (Bunge ex Ung.-Sternb.) Moss, 1954,
J.S. African Bot. 20: 16, isonym.
Lectotype (designated here): Nord de Mada-
gascar, [Diana Region] baie de Diego-Suarez [Antsira-
nana], II 1848, M. [L.H.] Boivin, №2370 (P: without
barcode).— Fig.2.
Ungern-Sternberg (1866: 51) attributes the name
Salicornia pachystachya to Bunge’s “Salicornia pachys-
tachya Bunge in herb.” not validly published by Bunge.
According to ICN (Turland et al., 2018: Art.46.2), the
authority of the name must be cited as Salicornia pa-
chystachya Bunge ex Ung.-Sternb.
Ungern-Sternberg (1866) indicates that the speci-
mens he examined are from the herbarium of Boivin (two
sheets in total), and no collection numbers are cited in
the protologue. Moss (1954) indicated that the type is
in P, while Toelken (1967) reported the location of the
type specimen in LE. In the latter Herba rium, no type
collection of Salicornia pachystachya was traced. One of
the two sheets (№ 2370) kept in P was labelled by an
anonym as “type” (P00487041), and the other specimen
labelled as “co-type” (also anonymously). The specimens
correspond with each other in gene ral morphological
characters and belong to the same plant. However, the
“type” contains a short fragment of the plant with sever-
al infl orescence branches and has been supplied with two
labels: (1)“Madagascar. Coll. Boivin”, and (2)“Zanzi-
bar. Boivin” (Bunge’s writing). The latter region is lo-
cated in Tanzania and not mentioned in the protologue
(Ungern-Sternberg, 1866). Due to the ambiguous loca-
tion of the specimen (Mada gascar versus Tanzania) and
its incompleteness, we chose a lectotype that contains a
fully developed twig collected in Madagascar.
The authentic material of Salicornia pachystachya
was collected at the fruiting stage. The seeds with hair-
like outgrowths of testa cells (Fig. 1: A) are very similar
to those of other annual Salicornia investigated earlier
(Shepherd et al., 2005a; Zare, Keshavarzi, 2007; Sukho-
rukov, 2014; Gasparri et al., 2016). This character has
evolved in all annual and some perennial species of Sali-
cornia (in its current circumscription), hence consid-
ered as a unique generic trait among all Salicornioideae.
Besides, almost none of Salicornia species in its current
circumscription have nutritive tissue (perisperm) in the
seeds. The complete fusion of perianth segments and
hyaline pericarp also unite all Salicornia species in its
recent circumscription (Shepherd et al., 2005a; Sukho-
rukov, 2014; Fuente et al., 2016). Consequently, S.pa-
chystachya cannot be considered within Arthrocnemum
(Arthrocaulon) based on the seed structure and should
be left in Salicornia.
Description. See Ungern-Sternberg (1866)
and Chevalier (1922).
The species is easily distinguished by thick (5–7
mm) infl orescences and relatively large (1.2–1.4 mm
long) seeds.
Habitat. Salicornia pachystachya is reported
to be one of the dominants of the halophytic vegeta-
tion near Tsimanampetsotse Lake (Rauh, 1995; Eve,
Pers, 2014), but it seems that these records belong to
Tecticornia indica, an abundant plant around the lake
(V. Alyonkin, pers. obs.). On the Europa Island, Sali-
cornia pachystachya inhabits salt marshes of low topo-
graphic level, located at the borders of the mediolitto-
ral and supralittoral domains (Fig.3:A), which leads to
weak and brief tidal fl ooding (up to a few centimetres
deep or sometimes just a moistening of the substrate).
These salt marshes have developed on coral-derived
muddy substrate surrounding the rugged karstic rocky
surface or rarely they occur on karst reef clogged with
muds (Boullet, 2014). The plant stature varies ac-
cording to the ecological conditions (see also Toelken,
1967), in particular according to the duration of tides
and the exposure to the tidal fl ow. The plants of the in-
tertidal habitats of the lagoon have an erect stem and
obliquely orientated lateral branches (usually at an an-
gle less than 45°). In the mangroves, on the edge of the
supratidal habitat complex experiencing irregular tidal
conditions, the plants often become decumbent and
more irregularly branched (Fig.3:B).
For Madagascar and the East Coast of Africa, detailed
ecological data are lacking, and a possibility of confusion
between Salicornia pachystachya and S.perrieri is higher.
Besides, S.pachystachya is also morphologically similar
to S.brachiata Roxb. known from the southern coast of
the Indian subcontinent. Bigot (1971) described the
surroundings of Tulear (Madagascar) as “Salicornia pa-
chystachya zone”, referring to abundant dry and highly
saline muddy coastal depressions.
Rarity status. The rarity status of Salicor-
nia pachystachya in Madagascar has not been evaluated
so far. On the Europa Island, S. pachystachya is pres-
ent in seven cells of 100× 100m, in three geographi-
cally distinct stations. This local rarity renders its rarity
status as “Exceptional”. Its population on the island is
estimated at around 1000 individuals and this popula-
tion covers a total area of about 5800m2 (Hivert, 2021;
Boullet, Hivert, pers. obs.). According to the criteria as
proposed by the Conservatoire Botanique National de
Mascarin (CBNM), and inspired by the IUCN Region-
al Red List (UICN France, 2011), but adapted to the
case of small territories, S. pachystachya is considered
“Vulnerable” on the Europa Island and the Scattered
Islands (Hivert et al., 2018; Boullet, Hivert, pers. obs.).
There are seemingly no direct threats to the Europa Is-
land population, except for putative sea-level changes.
A. P. Sukhorukov et al.
Новости систематики высших растений | Том 52 | 2021
42
Fig. 2. Lectotype of Salicornia pachystachya (P).
Notes on the samphires (Chenopodiaceae–Amaranthaceae) in Madagascar and Europa Island
Novitates Systematicae Plantarum Vascularium | Volume 52 | 2021
43
Distribution (Fig. 4: A). Madagascar: Diana
Region (see lectotype); [Boeny Region] Antongomena
Bevary, 23 VI 1965, R.Capuron, №24275 (P04618438,
K); [Mahajanga Region] Andranonomby, XII 1928,
H.Poisson, №378 (P04618440). Europa Island: Petite
Mangrove, 7 IV 2011, J.Hivert (CBNM-IE 92, CBNM,
MO).
General distribution. Madagascar,
French Southern and Atlantic Lands (Europa Island:
Boullet et al., 2018) and the East Coast of tropical Af-
rica (Kenya, Tanzania, and Mozambique). Also reported
from the East Coast of KwaZulu-Natal, South Africa
(Toelken, 1967; Germishuizen, Meyer, 2003).
Salicornia mossambicensis (Brenan) Piirainen et
G. Kadereit, 2017, Taxon, 66 (1): 125. Sarcocornia
mossambicensis Brenan, 1988, Fl. Zambes. 9 (1): 153.
Salicornia fruticosa auct., non (L.) L.
Arthrocnemum natalense auct., non (Bunge ex
Ung.-Sternb.) Moss.
Holotype: Mozambique, Inhambane Bay,
Mocucuni Island, 26 IX 1958, Mogg, № 29306
(LISC— photo!; isotype — SRGH).
Description. A detailed description is pro-
vided in the protologue (Brenan, 1988). It is distin-
guished from the related perennial Salicornia species
by the seeds with papillate (not hairy) outgrowths.
S.mossambicensis is mentioned by Leandri (1931) and
later by Cavaco (1954) under S. fruticosa (L.) L., or
as Arthrocnemum natalense (Bunge ex Ung.-Sternb.)
Moss by Toelken (1967). The examined specimens from
Madagascar correspond in their characters, especially
papillate seed surface (Fig.1: C, D), to S.mossambicen-
sis described from Mozambique (Brenan, 1988, as Sar-
cocornia mossambicensis Brenan).
Habitat. According to the herbarium labels,
the species occupies intertidal coastal salt marshes and
depressions.
Distribution (Fig. 4: B). Madagascar:
[Boeny Region] Majunga [Mahajanga], 24 VI 1912,
Fig. 3. Habitat and general view of the plants.
A, BSalicornia pachystachya, Europa Island, 27 V 2016, photo by Vincent Boullet; C, DTecticornia indica,
Madagascar, near Tsimanampetsotse Lake, I 2020, photo by Vitaly Alyonkin.
A. P. Sukhorukov et al.
Новости систематики высших растений | Том 52 | 2021
44
K. Afzelius (P051594880); [Atsimo-Andrefana Re-
gion] delta of the Linta River, VIII 1928, H. Hum-
bert, C. F. Swingle, № 5450 and 5451 (P04618441,
P04618442, P04618496, P04618497, K); [Atsimo-
Andrefana Region] Tsimanampetsotsa, 17 III 1962,
J. Bosser, № 15431 (P04618443); Atsimo-Andrefana
Region, Toliara, Morombe Distr., Befandriana Sud
commune, Andrianala, 8 VI 2015, C.Rakotovao et al.,
№6640 (P00956436).
Note. The data on the presence of “Salicornia fruticosa
at Manampetsa Lake, Toliara Province of Madagascar (Lean-
dri, 1931; Cavaco, 1954) may apply to S.mossambicensis (the
herbarium sheets might not exist).
General distribution.Madagascar and
Mozambique. Collections from the Scattered Islands
are still lacking.
Salicornia perrieri A. Chev. 1922, Rev. Bot. Appl.
Agric. Colon. 2: 749.
Lectotype (designated here): Madagascar,
[Boeny Region] Soalala (Ambongo = Mahajanga II
District), marais salants des environs du Soalala, VIII
1905, H.Perrier de la Bathie, №1580 (P: P00157180!;
isolectotype: P04618498!).
Description. See Chevalier (1922). The most
indicative characters of this species are annual life his-
tory, infl orescences up to 3(4–5) mm thick, and seeds
up to 1 mm long.
Habitat. Saline habitats, but a true nature of
the habitats still remains elusive.
Distribution (Fig. 4: C). Madagascar: [Boe-
ny Region] near Majunga [Mahajanga], 24 VI 1912,
K. Afzelius, № 256 (P05158071); [Atsimo-Andrefana
Region] near Toliara, IX 1924, H. Humbert, H. Per-
rier de la Bathie, № 2479 (P05267613); [Androy Re-
gion] Anony Lake, 26 II 1931, M. Decary, № 8569
(BM); [Bongolava Region] Bevato, s. d., B. Koechlin
(P05047688); [Atsimo-Andrefana Region] Toliara, Mo-
rombe, 27 VIII 2014, M.L.Rabarivola et al., №561 (K,
P00910878); Atsimo-Andrefana Region, Toliara Prov.,
Tsimanampetsotsa National Park, 1 II 2018, U.Swen-
son et al., №302 (P01182779).
General distribution. Madagascar.
Reported from Mozambique (Brenan, 1988; Lebrun,
Stork, 1991) and South Africa (O’Callaghan, 1992;
Germishuizen, Meyer, 2003; Slenzka et al., 2013), but
these records require confi rmation.
Tecticornia indica (Willd.) K. A. Sheph. et Paul
G. Wilson, 2007, Austral. Syst. Bot. 20 (4): 327.
Salicornia indica Willd. 1799, Neue Schriften Ges.
Naturf. Freunde Berlin, 2: 111. Arthrocnemum indicum
(Willd.) Moq. 1840, Chenop. Monogr. Enum. 111.
Halosarcia indica (Willd.) Paul G. Wilson, 1980,
Nuytsia, 3 (1): 63.
Holotype: India [Tamil Nadu State], Wepan-
scheri [Veppancheri], prope Trankenbar [Tharangam-
badi] [1797?], J.G.Klein, №3601 (B-W00054-010!).
Description. The morphological characters
of Madagascan Tecticornia indica specimens agree with
all previous descriptions of this species from other
territories (e. g., Moquin-Tandon, 1840; Backer, 1948,
both as Arthrocnemum indicum; Wilson, 1980). The
most indicative characters of T. indica are: glabrous
subshrubs forming sprawling mats (Fig.3: C, D); stems
prostrate or ascending, rooting at their base or not;
annual shoots upright, up to 20 cm tall, fl eshy, often
reddish or reddish-green, producing 10–15 pairs of
cuspidate leaves and terminating with dense cylindrical
infl orescences; bracts similar to the leaves, each bract
encloses three immersed fl owers; fruit with indurated
pericarp, contains yellowish-brown glabrous seed.
Habitat. Salt marches with Tecticornia indica
are found in the western and southwestern regions
of Madagascar. T. indica is found there on saline,
sandy substrates, often in fringing mangroves and in
temporary saline ponds (Bigot, 1971; Duranton, 1975;
Thomasson, Théodore, 1993; Koechlin et al., 1997).
The species is common around Tsimanampetsotsa
Lake (Atsimo-Andrefana Region), where it grows
with Caroxylon littorale (V. Alyonkin, pers. obs.). On
the Europa Island, Tecticornia indica is a native species
considered “relatively common” (i. e. present in 523
to 1062 (out of 3373) cells of 100 × 100 m; Boullet,
Hivert, 2021). Tecticornia indica is a dominant species
of the coastal paleotropical saltmarshes (Suaedo
monoicae-Tecticornietea indicae Knapp ex Boullet
2014) in East Africa (from southern Somalia to
southern Mozambique), in western and southwestern
Madagascar, and in the Europa Island. The species
grows in salt marshes at slightly elevated topography,
corresponding to the upper part of the supratidal
coastal border, that is fl ooded only at a very high tide
(Salsolo littoralis-Tecticornion indicae (Boullet,
2014)). This succulent subshrub usually forms dense,
low (not exceeding 15 cm) mats, showing spectacular
purplish-red colour (Bigot, 1971; Koechlin et al., 1997;
Boullet, 2014). On the Europa Island, the salt marshes
with T. indica are formed on saline coral limestone
muds around rugged stony karstic relief or surface
karstic pavement, and on the median part of supratidal
terraces, dry most of the year, except for the time of
spring tides (Boullet, 2014). On the East Coast of
Africa, the presence of the salt marshes with T. indica
was recorded long time ago (Knapp, 1965, 1973) but
their detailed ecological description is still lacking.
Notes on the samphires (Chenopodiaceae–Amaranthaceae) in Madagascar and Europa Island
Novitates Systematicae Plantarum Vascularium | Volume 52 | 2021
45
Rarity status. The rarity status of
Tecticornia indica is LC (Least Concern) at the scale
of both the Europa Island and the Scattered Islands
(Hivert et al., 2018; Boullet, Hivert, pers. obs.). No
direct threats seem to impact the populations on the
Europa Island; nevertheless it might be aff ected by
the sea level fl uctuations in future. The accompanying
species are Caroxylon littorale (Chenopodiaceae) and
Sesuvium portulacastrum (L.) L. subsp. portulacastrum
(Aizoaceae).
Distribution (Fig. 4: D). Madagascar:
[Atsimo-Antsiranana Region] Ambongo, 1841,
N. O. Pervillé, № 661 (P04618444–P04618446,
P04618448); [Sofi a Region] way to Befandriana, s.d.,
herb. J. Decuaire, № 27485 (P04618447); [Atsimo-
Andrefana Region] South of Toliara, 30 VII 1909,
F. Geay, № 6054 (P04618449); [Atsimo-Andrefana
Region], Toliara, 9 X 1921, H. Poisson, № 347
(P04618465); [Atsimo-Andrefana Region], near
Toliara, Befotaka, 12 V 1922, Petit, №6 (P04618499);
[Atsimo-Andrefana Region, near Toliara] Ihodo, 8 V
1924, R. Decary, № 2793 (P04618461); [Atsimo-
Andrefana Region] near Toliara, IX 1924, H.Humbert,
H. Perrier de la Bathie, № 2478 (P04618457,
P04618460); [Atsimo-Andrefana Region], delta of the
Linta River, VIII 1928, H. Humbert, C. F. Swingle,
№ 5452 (P04618452); [Boeny Region] Baie de Baly,
9 VI 1930, R.Decary, №7838 (P04618458); [Melaky
Region] Besalampy, 1932, H. Perrier de la Bathie,
№8639 (P04618450, P04618463); [Atsimo-Andrefana
Fig. 4. Distribution maps of Salicornia pachystachya (A), S.mossambicensis (B), S.perrieri (C), and Tecticornia indica
(D). Scale bars— 250 km.
A. P. Sukhorukov et al.
Новости систематики высших растений | Том 52 | 2021
46
Region], Tsimanampetsotsa, II 1947, H. Humbert,
№20244 (P04618462); [Menabe Region] Morondava,
IX 1956, J. Bosser, № 10041 (P04618464); [Atsimo-
Andrefana Region] near Toliara, 13 XI 1960, J.Leandri,
R. J. de Dieu, № 3836 (P04618451); Toliara, 26 II
1966, M. Peltier, № 4925 (P04618453); [Atsimo-
Andrefana Region] Tsimanampetsotsa Lake, 8 IX 1994,
R. Ranaivojaona et al., № 29 (P06795100). Europa
Island: Plaine centrale, 5 IV 2011, J.Hivert, V.Boullet,
L.D.B.Gigord (CBNM-IE 44, CBNM, MO, P).
General distribution. Madagascar,
French Southern and Atlantic Lands (Europa
atoll: Boullet et al., 2018), tropical Africa (Kenya,
Tanzania, Mozambique, Angola, and Senegal), Indian
subcontinent, South-East Asia, North Australia.
It has also been listed for the fl ora of South Africa
(Germishuizen, Meyer, 2003, as Halosarcia indica).
A key to the samphire species of Madagascar
This key only includes the most remarkable
characters of the plants. Anatomically, Tecticornia indica
diff ers from other samphires by having only a single
palisade layer of the stem chlorenchyma (see for more
De Fraine, 1913; Wilson, 1980; Voznesenskaya et al.,
2008). Some reproductive features are also diff erent,
the pericarp of T.indica is indurated and the seed has a
slightly bent embryo and abundant perisperm, whereas
all Madagascan Salicornia have a thin parenchymatous
pericarp, seeds with horseshoe-shaped embryo and
without nutritive tissue (perisperm). However, some
Australian Salicornia have seeds with a perisperm
(Shepherd et al., 2005a).
1. Erect or rarely decumbent annuals; seeds with hair-like
hooked outgrowths .................................................................. 2.
+ Prostrate or ascending subshrubs; seeds smooth or with
conical papillae .......................................................................... 3.
2. Infl orescences thick, obtusely conical; seeds 1.2–1.4 mm
long .................................................... Salicornia pachystachya.
+ Infl orescences thin, narrowly cylindrical; seeds < 1 mm
long ................................................................ Salicornia perrieri.
3. Perianth tip seen by the naked eye. Seeds papillate .............
........................................................... Salicornia mossambicensis.
+ Perianth completely hidden in bract. Seeds glabrous ..........
.......................................................................... Tecticornia indica.
Discussion
The reproductive characters of the Salicornioideae
genera in their recent circumscription were only
precised recently (Shepherd et al., 2005a; Sukhorukov,
2014; Sukhorukov, Nilova, 2016), and they play
a major role supporting the existing phylogenetic
subdivision of this subfamily. Insuffi ciently known
characters of each genus and taxonomic chaos often
led to misidentifi cations. The observations on the
presence of perisperm in the seeds (Chevalier, 1922)
and subshrubby life form (Cavaco, 1954) in Salicornia
pachystachya (as Arthrocnemum pachystachyum)
were erroneous and apply to Tecticornia indica, the
only samphire in Madagascar with both traits. Figure
II, images 6–8 of Salicornia pachystachya in Cavaco
(1954, as Arthrocnemum pachystachyum) depict
Tecticornia indica. Moreover, Salicornia perrieri cannot
be synonymised with S. pachystachya as proposed by
Toelken (1967) due to diff erent seed lengths, hence
they are treated here as separate species.
Distribution of Tecticornia indica in Africa
Tecticornia with approximately 33 species in its
recent circumscription is predominantly distributed
in Australia and adjacent parts of Malesia (Shepherd,
Wilson, 2007). T. indica is the most widespread
paleotropical species of the genus found in the
mangroves, on tidal fl ats and other saline (mostly
coastal) habitats of Australia, South Asia, and Africa
including Madagascar. The distribution of T. indica
in Africa was insuffi ciently known. According to the
APD (2021), it is present in North Africa (Tunisia,
Libya, Algeria), and West Africa (Senegal, Mauritania,
Morocco, Western Sahara), East Africa (Somalia,
Kenya, Tanzania, Mozambique), and South Africa. We
agree with Jafri and Rateeb (1978, as Arthrocnemum
indicum) that T. indica is absent in entire North
Africa. All specimens from this region identifi ed as
Arthrocnemum indicum” and cited in the literature
(Maire, 1962) belong to Arthrocaulon macrostachyum
and to other perennial samphires.
The presence of Tecticornia indica in West Africa
used to be a matter of debate. It has been reported for
Senegal (seashores of St. Louis town) by Hutchinson
and Dalziel (1927, as Salicornia indica), but Brenan
(1954b) and later Friis and Gilbert (1993) suggested
that T. indica was confused with Arthrocaulon
macrostachyum (cited as Arthrocnemum glaucum and
A. macrostachyum, respectively). These doubts were
also expressed by Lebrun and Stork (2003).
According to the herbarium specimens examined
and partially revised by the fi rst author (APS), Tecti-
cornia indica is present in continental East Africa (So-
malia, Kenya, Tanzania, and Mozambique: BM!, BR!,
K!, P!), on the Europa Island (Boullet et al., 2018) and
Madagascar. We can also confi rm the presence of T.in-
dica on the western coast of tropical Africa, namely in
Angola and Senegal (Hiern, 1900; Baker Clarke, 1913,
as Arthrocnemum indicum; Figueiredo, Smith, 2008, as
Halosarcia indica), but the herbarium collections from
these areas are scarce. The seeds of the Angolan plants
Notes on the samphires (Chenopodiaceae–Amaranthaceae) in Madagascar and Europa Island
Novitates Systematicae Plantarum Vascularium | Volume 52 | 2021
47
are slightly smaller with a rugose (not smooth) surface
(Fig. 3: F), but this may also be attributed to the infra-
specifi c variability. The specimens from KwaZulu-Natal
Province, South Africa (Germishuizen, Meyer, 2003, as
Halosarcia indica) were not seen by us. T.indica is not
known from other countries on the western coast of
Africa, and this huge disjunction in the distribution of
T.indica is highly intriguing.
The chorology of samphires in Africa
There are fi ve genera of samphires in Africa, Ar-
throcaulon (2–3 species), formerly known as Arthro-
cnemum (Piirainen et al., 2017), Salicornia which today
includes also Sarcocornia with ca. 20 species (Steff en et
al., 2015; Piirainen et al., 2017), Tecticornia (1species)
in its recent circumscription (Shepherd, Wilson, 2007),
Halocnemum represented by subshrubby H. cruciatum
(Forssk.) Tod. (Biondi et al., 2013; Sukhorukov, 2014)
and two Halopeplis Bunge ex Ung.-Sternb., an anuual
H. amplexicaulis Ung.-Sternb. ex Ces., Pass. et Gibelli
and subshrubby H. perfoliata Bunge ex Ung.-Sternb.
(Maire, 1962; Boulos, 1999).
Salicornioideae are unevenly distributed in Africa.
The recent records of Arthrocaulon south of the Equa-
tor (Figueiredo, Smith, 2008, as Arthrocnemum macro-
stachyum) belong to the perennial Salicornia species
known from Namibia, South Africa and Mozambique
(Scott, 1977; Brenan, 1988; O’Callaghan, Oliver, 1992;
Craven, 1999; Steff en et al., 2010; all as Sarcocornia). In
fact, Senegal and Cape Verde in the west and Horn of
Africa in the east are the southernmost regions where
Arthrocaulon is present, and the genus is mainly associ-
ated with the subtropical regions of North Africa and
Western Asia. It is represented in Senegal by A. fran-
zii (Sukhor.) Piirainen et G.Kadereit (Sukhorukov et
al., 2018a), the species described from Cape Verde Ar-
chipelago (Sukhorukov, Nilova, 2016) and previously
identifi ed as Arthrocnemum macrostachyum (Moric.)
K.Koch or A.glaucum (Delile) Ung.-Sternb. (e.g., Bre-
nan, 1954b; Berhaut, 1974; Martins, 2002). Both Arth-
rocnemum franzii and A. macrostachyum (incl. A.glau-
cum) now belong to the recently established genus
Arthrocaulon Piirainen et G.Kadereit (Piirainen et al.,
2017). A. macrostachyum, a species widely distributed
on the seashores of North Africa and Western Asia (Fri-
is, Gilbert, 1993; Sukhorukov, Nilova, 2016; Ramírez et
al., 2019), has the southernmost records in Mauritania
and Somalia. The ranges of both Arthrocaulon and Tec-
ticornia overlap in Senegal in the west and Somalia in
the east (Berhaut, 1974; Friis, Gilbert, 1993). Arthro-
caulon meridionalis Est. Ramírez, Rufo, Sánchez Mata
et Fuente is present in North Africa, at least in Moroc-
co (Ramírez et al., 2019). In contrast to Arthrocaulon,
the range of Tecticornia indica is limited entirely to the
tropical Africa (Brenan, 1954a; O’Callaghan, 1992).
Salicornia (incl. Sarcocornia) is sometimes consid-
ered to be cosmopolitan (O’Callaghan, 1992), but is
absent in some regions of Africa, especially in the Gulf
of Guinea and Horn of Africa (Cavaco, 1963; Friis,
Gilbert, 1993; APD, 2021). The annual samphires are
represented in North Africa by S.perennans Willd. and
probably also by S.procumbens Sm. subsp. procumbens
(Kadereit et al., 2012). Other annual species in Africa
include S. meyeriana Moss (incl. S. unifl ora Toelken)
found in South Africa and Namibia (Slenzka et al.,
2013), S. senegalensis A. Chev. in Senegal (Chevalier,
1922; Berhaut, 1974; APD, 2021) and probably also in
Mauritania (Lebrun, 1998), S. pachystachya from Ma-
da gascar, Mozambique, Tanzania, and Kenya (Brenan,
1954a; Cavaco, 1954, as Arthrocnemum pachystachyum;
Brenan, 1988), the Europa Island (Boullet, 2014) as
well as South Africa (Lebrun, Stork, 2003), and S.perri-
eri from Madagascar and Mozambique (Brenan, 1988).
S. pachystachya and S. perrieri are tropical elements
that are absent in the subtropics of southern Africa
(O’Callaghan, 1992).
Although Salicornia pachystachya was included in
the molecular analysis (Steff en et al., 2015; Piirainen et
al., 2017; Ball et al., 2017), the samples originated from
South Africa (KwaZulu-Natal) and belong in fact to a
perennial Salicornia (MJG— images seen!), not a “true”
S. pachystachya. Thus, the phylogenetic position of
S.pachystachya, as well as that of S.perrieri and S.sene-
galensis, has not been evaluated yet.
The perennial Salicornia (formerly Sarcocornia)
are found mainly in warm-temperate regions of Af-
rica. At least two species were recorded for North Af-
rica, S. alpini Lag. and S. pruinosa (Fuente, Rufo et
Sánchez Mata) Piirainen et G. Kadereit (Fuente et
al., 2016, as Sarcocornia; Piirainen et al., 2017), which
comprise a Mediterranean clade of the genus. They
may belong to the Salicornia subgen. Arthrocnemoides
Ung.-Sternb., which incorporates the morphologically
similar S. fruticosa and S. perennis Mill. (Piirainen et
al., 2017). Two further species were described by Che-
valier (1934) from South Algeria, but they have not
been properly studied yet. The fi rst one, S. deserticola
A.Chev., is not an annual (Chevalier, 1934) but rather
a suff ruticose perennial (Piirainen, 2015); it was ac-
cepted by Maire (1962) but synonymised with S. fru-
ticosa (Piirainen, 2015, as Sarcocornia fruticosa). The
other species, S.longispicata A.Chev., was also accepted
by Maire (1962) but merged with S. perennis Mill. in
APD (2021). The plants with the same small habit as
in S.longispicata were collected by V.P.Bochantsev at
the fruiting stage in the neighbouring regions of Algeria
A. P. Sukhorukov et al.
Новости систематики высших растений | Том 52 | 2021
48
(LE!, as Salicornia arabicaL.) and found to belong to
Arthrocaulon. Both Salicornia deserticola and S.longispi-
cata need further investigation.
The samphires in southern Africa are represented
by perennial species belonging to the Salicornia subgen.
Afrocornia Piirainen et G.Kadereit and all are endemic
to this region (Steff en et al., 2010, 2015; Piirainen et
al., 2017). The Mediterranean S.fruticosa is not found
in Madagascar and southern Africa, and all its records
belong to diff erent species, such as S. natalensis Bunge
ex Ung.-Sternb., S. mossiana (Toelken) Piirainen et
G.Kadereit and S.mossambicensis.
In total, Salicornia in Africa is represented by ca. 20
species, with at least four of them known in the North
and 13 in the southern Africa.
Acknowledgements
The authors thank the French Southern and Antarc-
tic Territories (TAAF) for their support for the inven-
tory programs of the fl ora and vegetation of the Scat-
tered Islands (and in particular the Europa Island). We
also thank Ladislav Mucina and anonymous reviewer
for the comments and suggestions on the fi rst draft of
the paper. The research of APS and MAK was support-
ed by the scientifi c programmes 121032500084-6 and
121032300068-8 of the Department of Higher Plants
and Department of Plant Physiology, respectively (Lo-
monosov Moscow State University).
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Новости систематики высших растений | Том 52 | 2021
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Salicornioideae (Amaranthaceae/Chenopodiaceae) comprise 11 genera and ca. 100 species of succulent halophytes distributed worldwide in coastal and inland saline habitats. Most species have peculiar articulated, seemingly leafless stems and spike-like sessile thyrses with highly reduced flowers hidden by fleshy bracts. We analysed sequence data of four markers (ETS, ITS, atpB-rbcL, matK-trnK) from 57 species, 9 subspecies, 1 variety and 2 hybrids representing all curently accepted genera using maximum likelihood, Bayesian analysis, fossil-calibrated molecular dating and ancestral area analysis. All molecular markers gave similar phylogenetic signals. Salicornioideae probably originated in Eurasia during the late Eocene–early Oligocene. The divergence of the early main clades took place in the middle Oligocene with the separation of Allenrolfea/Heterostachys, Halocnemum/Halopeplis/Halostachys/Kalidium and Arthrocnemum/Microcnemum/Tecticornia/Salicornia/Sarcocornia lineages. The latter lineage diversified most and comprises ¾ of the species of the subfamily. The Arthrocnemum macrostachyum/Microcnemum lineage diverged at the turn of the Oligocene/Miocene – while the Arthrocnemum subterminale, Tecticornia and Salicornia/Sarcocornia lineages were all present by the middle Miocene. Long-distance dispersal has taken place several times to the Americas (Allenrolfea/Heterostachys, Arthrocnemum subterminale, Salicornia/Sarcocornia), South Africa (Halopeplis, Salicornia/Sarcocornia) and Australia (Tecticornia, Salicornia/Sarcocornia). Most currently accepted genera were resolved as monophyletic. However, Sarcocornia is paraphyletic in relation to Salicornia, and Arthrocnemum proved to be polyphyletic. In addition, the name Arthrocnemum is a nomenclatural synonym of Salicornia. As a consequence, we propose the merging of Sarcocornia under Salicornia and provide 19 new nomenclatural combinations and one replacement name. For Salicornia (incl. Sarcocornia) we propose a new infrageneric classification to accommodate the four phylogenetically and geographically well-supported sublineages of the genus. We also propose abandoning the name Arthrocnemum in its current use, and describe two new genera, Arthrocaulon (Eurasia, Africa) and Arthroceras (North America), and provide three new nomenclatural combinations.