Taxonomic and nomenclatural notes on Pontic-Mediterranean coastal and some Australasian taxa of Salsola (Chenopodiaceae)
Current and historical views on taxonomy and nomenclature of Pontic-Mediterranean coastal and some Australasian taxa of Salsola (Chenopodiaceae) are analyzed. Taxonomic identity and nomenclature of several names applied and misapplied to members of the species group known in recent publications ma...
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irk-123456789-1758882021-02-04T01:30:22Z Taxonomic and nomenclatural notes on Pontic-Mediterranean coastal and some Australasian taxa of Salsola (Chenopodiaceae) Mosyakin, S.L. Систематика, флористика, географія рослин Current and historical views on taxonomy and nomenclature of Pontic-Mediterranean coastal and some Australasian taxa of Salsola (Chenopodiaceae) are analyzed. Taxonomic identity and nomenclature of several names applied and misapplied to members of the species group known in recent publications mainly as Salsola pontica (Pall.) Degen sensu lato are discussed. It is demonstrated that Kali dodecanesicum C. Brullo & al. is a later synonym of Salsola squarrosa Steven ex Moq., which appears to be the earliest species-rank name available for the whole Pontic-Mediterranean group of taxa. This group is represented by Western Mediterranean (S. controversa Tod. ex Lojac.), Eastern Mediterranean (S. squarrosa s. str.), and mainly Pontic (S. pontica s. str.) geographical races. Considering the blurred morphological and geographical limits between these taxa, they are better treated as three subspecies of S. squarrosa: subsp. controversa (Tod. ex Lojac.) Mosyakin, comb. nov., subsp. squarrosa, and subsp. pontica (Pall.) Mosyakin, comb. nov., respectively. It is demonstrated that S. macrophylla R. Br. (described from Australia) is not conspecific with any of Pontic-Mediterranean coastal taxa but is probably related to the currently recognized species S. australis R. Br. The identity of S. brachypteris Moq. (described from Java, Indonesia) remains uncertain but, judging from available evidence, it is most probably either related to or conspecific with S. macrophylla, or some other insufficiently known Australasian species. The problem of conflicting typifications and application of the name S. caroliniana Walter is briefly discussed; it is concluded that the name should be proposed for rejection. The need for further morphological, molecular phylogenetic, and phylogeographic studies of coastal Eurasian and Australasian species of Salsola is emphasized. Проаналізовано сучасні та історичні погляди щодо систематики та номенклатури понтично-середземноморських та деяких австралазійських таксонів Salsola (Chenopodiaceae). Обговорюється таксономія та номенклатура декількох назв, які раніше застосовувалися до представників видової групи, що здебільшого була визнана у недавніх публікаціях під назвою Salsola pontica (Pall.) Degen s. l. Показано, що Kali dodecanesicum C. Brullo & al. є синонімом раніше описаного виду Salsola squarrosa Steven ex Moq.; остання назва є пріоритетною у ранзі виду для усієї понтично-середземноморської групи таксонів. Ця група представлена західносередземноморською (S. controversa Tod. ex Lojac.), східносередземноморською (S. squarrosa s. str.) та переважно чорноморсько-каспійською (S. pontica s. str.) географічними расами. Враховуючи нечіткі морфологічні та географічні межі між цими таксонами, їх доцільно розглядати як три підвиди виду S. squarrosa: subsp. controversa (Tod. ex Lojac.) Mosyakin, comb. nov., subsp. squarrosa, та subsp. pontica (Pall.) Mosyakin, comb. nov., відповідно. Показано, що вид S. macrophylla R. Br. (описаний з Австралії) є відмінним від усіх понтично-середземноморських прибережних таксонів; натомість він, очевидно, споріднений з нині визнаним видом S. australis R. Br. Ідентичність S. brachypteris Moq. (описаний з о. Ява, Індонезія) лишається невизначеною; судячи з наявних даних, цей таксон, ймовірно, споріднений з S. macrophylla або з деякими іншими недостатньо відомими австралазійськими видами. Стисло розглянута проблема типіфікації та застосування назви S. caroliniana Walter; зроблено висновок, що цю назву слід запропонувати до відхилення. Наголошено на потребі проведення подальших морфологічних, молекулярно-філогенетичних та філогеографічних досліджень приморських євразійських та австралазійських видів Salsola. Проанализированы современные и исторические взгляды на систематику и номенклатуру понтическо-средиземноморских и некоторых австралазийських таксонов рода Salsola ( Chenopodiaceae). Обсуждены систематика и номенклатура нескольких названий, которые ранее применялись к представителям видовой группы, которая признавалась в недавних публикациях преимущественно под названием Salsola pontica (Pall.) Degen s. l. Показано, что Kali dodecanesicum C. Brullo & al. является синонимом ранее описанного вида Salsola squarrosa Steven ex Moq.; последнее название является приоритетным в ранге вида для всей понтическо-средиземноморской группы таксонов. Эта группа представлена западносредиземноморской (S. controversa Tod. ex Lojac.), восточносредиземноморской (S. squarrosa s. str.) и преимущественно черноморско-каспийской (S. pontica s. str.) географическими расами. Учитывая нечеткие морфологические и географические границы между этими таксонами, их целесообразно рассматривать как три подвида вида S. squarrosa: subsp. controversa (Tod. ex Lojac.) Mosyakin, comb. nov., subsp. squarrosa, и subsp. pontica (Pall.) Mosyakin, comb. nov., соответственно. Показано, что вид S. macrophylla R. Br. (описан из Австралии) отличается от всех понтическо-средиземноморских прибрежных таксонов; он, очевидно, является родственным признанному сейчас виду S. australis R. Br. Идентичность S. brachypteris Moq. (описан с о. Ява, Индонезия) остается неопределенной; судя по имеющимся данным, этот таксон, вероятно, родственен S. macrophylla или некоторым другим недостаточно изученным австралазийским видам. Кратко рассмотрена проблема типификации и применения названия S. caroliniana Walter; сделан вывод о том, что это название следует предложить номенклатурно отвергнуть. Отмечена необходимость проведения дальнейших морфологических, молекулярно-филогенетических и филогеографичних исследований приморских евразийских и австралазийских видов Salsola. 2017 Article Taxonomic and nomenclatural notes on Pontic-Mediterranean coastal and some Australasian taxa of Salsola (Chenopodiaceae) / S.L. Mosyakin // Український ботанічний журнал. — 2017. — Т. 74, № 6. — С. 521-531. — Бібліогр.: 48 назв. — англ. 0372-4123 DOI: 10.15407/ukrbotj74.06.521 http://dspace.nbuv.gov.ua/handle/123456789/175888 en Український ботанічний журнал Інститут ботаніки ім. М.Г. Холодного НАН України |
| institution |
Digital Library of Periodicals of National Academy of Sciences of Ukraine |
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| language |
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| topic |
Систематика, флористика, географія рослин Систематика, флористика, географія рослин |
| spellingShingle |
Систематика, флористика, географія рослин Систематика, флористика, географія рослин Mosyakin, S.L. Taxonomic and nomenclatural notes on Pontic-Mediterranean coastal and some Australasian taxa of Salsola (Chenopodiaceae) Український ботанічний журнал |
| description |
Current and historical views on taxonomy and nomenclature of Pontic-Mediterranean coastal and some Australasian
taxa of Salsola (Chenopodiaceae) are analyzed. Taxonomic identity and nomenclature of several names applied and misapplied
to members of the species group known in recent publications mainly as Salsola pontica (Pall.) Degen sensu lato are discussed. It
is demonstrated that Kali dodecanesicum C. Brullo & al. is a later synonym of Salsola squarrosa Steven ex Moq., which appears
to be the earliest species-rank name available for the whole Pontic-Mediterranean group of taxa. This group is represented
by Western Mediterranean (S. controversa Tod. ex Lojac.), Eastern Mediterranean (S. squarrosa s. str.), and mainly Pontic
(S. pontica s. str.) geographical races. Considering the blurred morphological and geographical limits between these taxa, they
are better treated as three subspecies of S. squarrosa: subsp. controversa (Tod. ex Lojac.) Mosyakin, comb. nov., subsp. squarrosa,
and subsp. pontica (Pall.) Mosyakin, comb. nov., respectively. It is demonstrated that S. macrophylla R. Br. (described from
Australia) is not conspecific with any of Pontic-Mediterranean coastal taxa but is probably related to the currently recognized
species S. australis R. Br. The identity of S. brachypteris Moq. (described from Java, Indonesia) remains uncertain but, judging
from available evidence, it is most probably either related to or conspecific with S. macrophylla, or some other insufficiently
known Australasian species. The problem of conflicting typifications and application of the name S. caroliniana Walter is briefly
discussed; it is concluded that the name should be proposed for rejection. The need for further morphological, molecular
phylogenetic, and phylogeographic studies of coastal Eurasian and Australasian species of Salsola is emphasized. |
| format |
Article |
| author |
Mosyakin, S.L. |
| author_facet |
Mosyakin, S.L. |
| author_sort |
Mosyakin, S.L. |
| title |
Taxonomic and nomenclatural notes on Pontic-Mediterranean coastal and some Australasian taxa of Salsola (Chenopodiaceae) |
| title_short |
Taxonomic and nomenclatural notes on Pontic-Mediterranean coastal and some Australasian taxa of Salsola (Chenopodiaceae) |
| title_full |
Taxonomic and nomenclatural notes on Pontic-Mediterranean coastal and some Australasian taxa of Salsola (Chenopodiaceae) |
| title_fullStr |
Taxonomic and nomenclatural notes on Pontic-Mediterranean coastal and some Australasian taxa of Salsola (Chenopodiaceae) |
| title_full_unstemmed |
Taxonomic and nomenclatural notes on Pontic-Mediterranean coastal and some Australasian taxa of Salsola (Chenopodiaceae) |
| title_sort |
taxonomic and nomenclatural notes on pontic-mediterranean coastal and some australasian taxa of salsola (chenopodiaceae) |
| publisher |
Інститут ботаніки ім. М.Г. Холодного НАН України |
| publishDate |
2017 |
| topic_facet |
Систематика, флористика, географія рослин |
| url |
http://dspace.nbuv.gov.ua/handle/123456789/175888 |
| citation_txt |
Taxonomic and nomenclatural notes on Pontic-Mediterranean coastal and some Australasian taxa of Salsola (Chenopodiaceae) / S.L. Mosyakin // Український ботанічний журнал. — 2017. — Т. 74, № 6. — С. 521-531. — Бібліогр.: 48 назв. — англ. |
| series |
Український ботанічний журнал |
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AT mosyakinsl taxonomicandnomenclaturalnotesonponticmediterraneancoastalandsomeaustralasiantaxaofsalsolachenopodiaceae |
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2025-07-15T13:19:08Z |
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2025-07-15T13:19:08Z |
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| fulltext |
521Укр. бот. журн., 2017, 74(6)
Introduction
Salsola L. sensu lato (Chenopodiaceae: Salsoloideae:
Salsoleae) is a notoriously complicated group from the
taxonomic and phylogenetic viewpoints; it was recently
split into numerous segregate genera supported by
molecular phylogenetic and partly morphological and
biogeographical evidence (see Akhani et al., 2007; Wen
et al., 2010; Wen, Zhang, 2011, Voznesenskaya et al.,
2013; Schüssler et al., 2017, and references therein).
The process of generic rearrangements in that group
is still far from being complete, and further taxonomic
and nomenclatural changes are expected. Recent
nomenclatural and taxonomic developments resulting
from molecular phylogenetic studies and nomenclatural
controversies (Akhani et al., 2014; Mosyakin et al.,
2014) were summarized in several recent publications
(e. g., Hernández-Ledesma et al., 2015; Mosyakin
et al., 2017; Mosyakin, 2017, and references therein)
and will not be repeated here.
I provide here a brief outline of the state of
current knowledge of and remaining taxonomic and
nomenclatural problems in just one the group of coastal
taxa presumably native to the Mediterranean region
and extending to the maritime areas of the Black
and Caspian seas and the Sea of Azov. Several names © S.L. MOSYAKIN, 2017
doi: 10.15407/ukrbotj74.06.521
Taxonomic and nomenclatural notes on Pontic-Mediterranean coastal
and some Australasian taxa of Salsola (Chenopodiaceae)
Sergei L. MOSYAKIN
M.G. Kholodny Institute of Botany, National Academy of Sciences of Ukraine
2, Tereshchenkivska Str., Kyiv (Kiev) 01004, Ukraine
s_mosyakin@hotmail.com
Mosyakin S.L. Taxonomiс and nomenclatural notes on Pontic-Mediterranean coastal and some Australasian taxa of Salsola
(Chenopodiaceae). Ukr. Bot. J., 2017, 74(6): 521–531.
Abstract. Current and historical views on taxonomy and nomenclature of Pontic-Mediterranean coastal and some Australasian
taxa of Salsola (Chenopodiaceae) are analyzed. Taxonomic identity and nomenclature of several names applied and misapplied
to members of the species group known in recent publications mainly as Salsola pontica (Pall.) Degen sensu lato are discussed. It
is demonstrated that Kali dodecanesicum C. Brullo & al. is a later synonym of Salsola squarrosa Steven ex Moq., which appears
to be the earliest species-rank name available for the whole Pontic-Mediterranean group of taxa. This group is represented
by Western Mediterranean (S. controversa Tod. ex Lojac.), Eastern Mediterranean (S. squarrosa s. str.), and mainly Pontic
(S. pontica s. str.) geographical races. Considering the blurred morphological and geographical limits between these taxa, they
are better treated as three subspecies of S. squarrosa: subsp. controversa (Tod. ex Lojac.) Mosyakin, comb. nov., subsp. squarrosa,
and subsp. pontica (Pall.) Mosyakin, comb. nov., respectively. It is demonstrated that S. macrophylla R. Br. (described from
Australia) is not conspecific with any of Pontic-Mediterranean coastal taxa but is probably related to the currently recognized
species S. australis R. Br. The identity of S. brachypteris Moq. (described from Java, Indonesia) remains uncertain but, judging
from available evidence, it is most probably either related to or conspecific with S. macrophylla, or some other insufficiently
known Australasian species. The problem of conflicting typifications and application of the name S. caroliniana Walter is briefly
discussed; it is concluded that the name should be proposed for rejection. The need for further morphological, molecular
phylogenetic, and phylogeographic studies of coastal Eurasian and Australasian species of Salsola is emphasized.
Keywords: Salsola, Chenopodiaceae, Salsoloideae, nomenclature, taxonomy, biogeography
This article is dedicated to Alfred [Christian Horace Bénédict
Alfred] Moquin-Tandon (1804–1863), whose excellent
publications on Chenopodiaceae remain valuable resources for
all plant taxonomists studying this fascinating plant family
[image in Public Domain: https://fr.wikipedia.org/wiki/Alfred_
Moquin-Tandon#/media/File:Alfred_Moquin-Tandon.jpg]
https://fr.wikipedia.org/wiki/Alfred_Moquin-Tandon#/media/File:Alfred_Moquin-Tandon.jpg
https://fr.wikipedia.org/wiki/Alfred_Moquin-Tandon#/media/File:Alfred_Moquin-Tandon.jpg
522 Ukr. Bot. J., 2017, 74(6)
concluded that for establishing the priority name (or
names) for the Pontic-Mediterranean coastal species
(or a group of species) "the identity of two earlier
species-rank names (S. macrophylla described from
Australia and S. brachypteris Moq. described from Java,
Indonesia) mentioned by Rilke (1999) in synonymy of
her S. tragus subsp. pontica should be critically assessed
as well" (Mosyakin, 2017: 410).
Following these preliminary conclusions, in
this article I provide further results of my critical
assessment of available evidence on the possible
identity and application of the names S. macrophylla
and S. brachypteris and on forgotten (and almost never
taxonomically recognized until now) Mediterranean
taxa S. squarrosa Steven ex Moq. and S. controversa Tod.
ex Lojac. Additionally, it was necessary to re-evaluate
available data on the name S. caroliniana Walter, a very
obscure taxon described from North America in 1788.
Two new nomenclatural combinations are proposed for
Pontic-Mediterranean coastal taxa.
Herbarium acronyms are given following Thiers
(2017–onward). Digital images of many herbarium
specimens cited in the article are available online.
What is Salsola macrophylla R. Br.?
The type specimen of Salsola macrophylla
(BM000016766, available at https://plants.jstor.
org/stable/10.5555/al.ap.specimen.bm000016766)
has the printed label on blue paper ("R. Brown, Iter
Australiense, 1802–5. [Presented by direction of
J.J. Bennett, 1876], No. 3082" [handwritten number
added – S.M.]) and the handwritten label ("2. Salsola
macrophylla. prodr. 411. East Coast") attached to it.
There is also a handwritten pencil text in the lower right
corner of the sheet: "Brown's manuscript description
of Salsola macrophylla was based on material collected
on inner entrance [of] Thirsty Sound, Queensland
[illegible text follows, scan cut – S.M.]". A recent
map with a cross indicating the locus classicus and
Rilke's identification label ("Salsola tragus L. subsp.
pontica. Det./rev. S. Rilke. Kassel, 30.1.1999") are also
attached. In my opinion, the specimen does not belong
to S. pontica (sensu stricto or sensu lato).
The type specimen contains four plant fragments:
two larger terminal branches (mounted on the left and
right sides of the sheet), a middle portion of a branch
(central lower part of the sheet), and a small terminal
part of a branch (center of the sheet). All fragments
are morphologically similar and thus there is no reason
to suspect that they represent a mixed collection of
different taxa. Larger branch fragments are rather thick;
they belonged to a robust or probably even somewhat
lignescent plant with erect or somewhat ascending
were applied and misapplied to these coastal plants,
and some additional taxa were recently described or
recognized. Morphologically similar plants, however,
also occur in Australasia (where some might be aliens
and some native plants: see Borger et al., 2008; Hrusa,
Gaskin, 2008; Chinnock, 2010), North America (all
aliens; see Mosyakin, 1996, 2003; Rilke, 1999; Hrusa,
Gaskin, 2008; Ayres et al., 2009), and probably some
other coastal regions of the world.
In her concise monograph of Salsola sect. Salsola
sensu lato, Rilke (1999) recognized just two western
European and Mediterranean coastal taxa of Salsola:
S. kali L. sensu stricto (from the Atlantic and Baltic
coasts) and S. tragus L. subsp. pontica (Pall.) Rilke (from
shores of the Mediterranean, Black and Caspian seas,
and the Sea of Azov). She also provided extensive lists of
synonyms for each recognized taxon, in most cases with
information on their types, and relevant nomenclatural
references. The species included by Rilke (1999) in her
"Salsola sect. Salsola sensu lato" (in fact, she recognized
sect. Salsola sensu stricto, sect. Sogdiana Rilke, sect.
Androssowia Rilke, and sect. Kali Dumort. with three
subsections) are now placed in three genera: Salsola
sensu stricto, Soda (Dumort.) Fourr. (Fourreau,
1869; see Mosyakin et al., 2017), and Turania Akhani
& Roalson (Akhani et al., 2007). Additionally, one
species (S. rosacea L.) probably fits in Noaea Moq. (see
Akhani et al., 2014); as far as I am aware, no relevant
combination has been validated yet.
The present article grew from my recent paper
published with a modest intention to communicate
a new record of the alien species S. paulsenii Litv. in
Ukraine (Mosyakin, 2017). That article, despite being
rather limited in its scope, contained a brief overview
of species of Salsola sensu lato (now placed in Salsola
sensu stricto, Caroxylon Thunb., Climacoptera Botsch.,
Pyankovia Akhani & Roalson, and Soda) occurring
in Ukraine, and some notes on nomenclature and
taxonomy of coastal Salsola pontica and related taxa.
Taxa placed by Rilke (1999) in Salsola tragus subsp.
pontica: a brief overview
I already commented that "[s]ynonymization of the
Australian taxon S. macrophylla with the Pontic (and
eastern Mediterranean?) S. pontica (Pall.) Degen (=
Kali ponticum (Pall.) Sukhor., S. tragus subsp. pontica
(Pall.) Rilke) cursorily done by Rilke (1999) and later
accepted by Galasso and Bartolucci (2014), who coined
the combination Kali macrophyllum (R. Br.) Galasso
& Bartolucci, seems to be at least questionable. Their
identity has not been proven yet beyond doubt, and
until such a proof is available, I prefer to use the name
S. pontica for our plants" (Mosyakin, 2017: 410). I also
https://plants.jstor.org/stable/10.5555/al.ap.specimen.bm000016766
https://plants.jstor.org/stable/10.5555/al.ap.specimen.bm000016766
523Укр. бот. журн., 2017, 74(6)
suberectis, foliis alternis elongatis semiteretibus
spinoso-muricatis crassis carnosis rigidibus, floralibus
vix dilatatis, bracteis folio florali multò brevioribus
perigonio fructifero longioribus, floribus solitariis,
alis minutissimis squamæformibus vix distinctis
cartilagineis.
In insulà Java.
Planta sicca fragilis. Folia caduca. Facies S. Sodæ.
Fructus vix inflati. (v. s. ex itin. Belang.)".
Later Moquin-Tandon (1849: 189) provided an
updated description and additional information on his
species, including references to additional herbarium
specimens seen ("v. s. in h. Belang. Mus. paris. DC. et
Hook.").
The plant was collected in Java (now Indonesia) by
Charles Paulus Bélanger (1805–1881) during his travels
in Europe, the Caucasus, Persia, India, Indonesia,
and some other regions in 1825–1829, described in his
Voyages aux Indes orientales par le nord de l'Europe, les
provinces du Caucase, la Géorgie, l'Arménie et la Perse,
suivi de détails topographiques et autres sur le Pégou, les
îles de Java, de Maurice et de Bourbon, sur le Cap de
Bonne-Espérance et Sainte-Hélène, pendant les années
1825, 1826, 1827, 1828 et 1829; see Ross (1964) for
complicated bibliographic details. In 1828 Bélanger
visited Java, mainly areas of Batavia (now Jakarta) and
Buitenzorg (now Bogor), and some adjacent islands
(Hooker, 1836).
In subsequent publications Salsola brachypteris
was usually mentioned as a synonym of S. kali sensu
latissimo (e. g., Boerlage 1900: 38; Backer, 1949: 106),
and only rarely it was recognized as a species (e. g.,
Miquel, 1855: 1022; Hooker, 1859: xlvi).
I traced online the original specimen from Java that
was certainly studied in detail by Moquin-Tandon and
should be considered the type (P00799151; available
online from https://science.mnhn.fr/institution/mnhn/
collection/p/item/p00799151). The sheet contains two
plant fragments (one mounted in the center of the
sheet, another much smaller fragment in the envelope
in the upper right corner), the labels, and additional
annotation slips on which Moquin-Tandon wrote a
description (which was evidently used for preparing
the published descriptions) and analytical drawings
illustrating diagnostic characters of the species. The
yellow-paper label in the lower left corner of the sheet
contains the following text: "Salsola brachypteris Moq.
Java No. 4. M. Bèlangé [Bélanger – S.M.]".
There are several other available specimens in P
identified as S. brachypteris (e.g., P05196083, image
available from https://science.mnhn.fr/institution/
mnhn/collection/p/item/p05196083; P00256012, https://
branches. Stem leaves are indeed rather long; fruits
are mainly immature, but it looks like the fruits in
nodes and lower parts of branches have short-winged
or almost wingless tepals. However, a few fruits in the
terminal part of the central branch fragment are better
developed, and those fruits evidently have tepals with
quite well-developed and broad wings. One fruit is
especially well visible; as far as it can be seen in the
high-resolution digital image, its two minor wings are
rather large also, similar to those in S. australis. Judging
from that evidence, in combination with morphological
data provided by Borger et al. (2008) and Chinnock
(2010), we can conclude that S. macrophylla most
probably represents a rather tall-growing tropical and
subtropical Australasian race, probably a lignescent
short-lived perennial, which is related to S. australis. Its
taxonomic status remains obscure, but data of Borger
et al. (2008) and Chinnock (2010) indicate that some
taxonomic recognition of that plant is to be expected
(a separate species, or a subspecies of S. australis?),
following further research. Consequently, the name
S. macrophylla should not be applied to any native
Pontic-Mediterranean coastal taxa of Salsola.
What is Salsola brachypteris Moq.?
Rilke (1999: 133) listed the names Salsola
brachypteris Moq. and S. kali var. brachypteris (Moq.)
Benth. (Bentham, 1870) in synonymy of her S. tragus
subsp. pontica (Pall.) Rilke. If it is indeed so, the name
S. brachypteris could be of priority for the whole Pontic-
Mediterranean complex, or for some of its segregate
species (if recognized). Because of that I studied in
more detail the protologue, other relevant literature,
and high-resolution images of original and some other
specimens available online.
The name S. brachypteris appeared for the first
time in the article by Moquin-Tandon (1835: 214) as
nomen nudum because no description or diagnosis of
the species was provided. The name was mentioned in
the key to groups of species of Salsola and was included
in the third group, with the following diagnosis:
"Alæ crassiusculæ, brevissimæ (unguiformes). Ex.:
S. Soda L., S. collina Pall. Ill., S. brachypteris Moq. in
Belang." It is evident that this brief diagnostic statement
was applied to several species, of which only three taxa
were mentioned as examples. No diagnostic characters
distinguishing S. brachypteris from other taxa have
been provided. Consequently, the name remained
invalid until 1840, when Moquin-Tandon (1840: 147)
published the following description:
"28. S. brachypteriS (Moq. in Belang. fl. Pers.)
herbacea suberecta glauca ramosa, ramis alternis
https://science.mnhn.fr/institution/mnhn/collection/p/item/p00799151
https://science.mnhn.fr/institution/mnhn/collection/p/item/p00799151
https://science.mnhn.fr/institution/mnhn/collection/p/item/p05196083
https://science.mnhn.fr/institution/mnhn/collection/p/item/p05196083
https://science.mnhn.fr/institution/mnhn/collection/p/item/p00256012
524 Ukr. Bot. J., 2017, 74(6)
Mediterranean taxon (either sensu lato or sensu stricto).
It might be related to S. macrophylla (see above), a
predominantly tropical Australasian (or exclusively
Australian?) race of the S. australis aggregate, which
is in need of further research. It should be noted that
some Australian specimens of Salsola also have that
very peculiar branching habit, with subopposite or
opposite arrangement of lower branches and leaves
(see, for example, a specimen of "S. kali var. strobilifera
Benth."; K000899590, image available from: https://
plants.jstor.org/stable/viewer/10.5555/al.ap.specimen.
k000899590). Many intriguing questions about
that taxon remain, but, in any case, we can safely
exclude the name S. brachypteris from our further
considerations on appropriate names to be applied to
Pontic-Mediterranean taxa.
In my opinion, the amazing morphological diversity
of Australian Salsola does not fit just one native species
now recognized as S. australis. Judging from specimens
and images I have seen, and from other available
evidence (Mueller, 1891; Wilson, 1984; Rilke, 1999;
Borger et al., 2008; Chinnock, 2010, etc.), there are
at least five native Australian species of Salsola (plus
probably one or two introduced ones?). An article is
under preparation (Mosyakin et al., in progress), in
collaboration with Australian colleagues.
The problem of Salsola caroliniana Walter
It is now evident that the names S. macrophylla
and S. brachypteris should not be applied to native
Pontic-Mediterranean taxa. However, before proposing
taxonomic and nomenclatural solutions for the coastal
Pontic-Mediterranean species complex of Salsola, we
should consider in more detail another earlier name,
S. caroliniana Walter, which may compete in terms of
priority with the names of Pontic-Mediterranean taxa.
Salsola caroliniana was described by Thomas Walter in
Flora Caroliniana (Walter, 1788: 111), one of the earliest
North American floristic and taxonomic publications
consistently applying binomial nomenclature and
Linnaean taxonomy. The real identity of that species
name remains uncertain (see Botschantzev, 1974;
Tzvelev, 1993, 1996; Mosyakin, 1996, 2003; Rilke,
1999, and references therein). The standing type of
S. caroliniana (Botschantzev, 1974: 614; Rilke, 1999)
is a small fragment of a Salsola branch mounted on
page 93 (specimen 93-F) of the Walter Herbarium folio
volume kept at BM (image available from Botanica
Caroliniana – Texts, Data, and Images at http://folio.
furman.edu/botcarweb/indices?urn=urn:cite:fufolio
img:BotCarWalter.walter_v_93_01135). It definitely
belongs to the S. kali – S. tragus – S. pontica aggregate,
but is hardly identifiable precisely as a concrete species
or subspecies of that group.
science.mnhn.fr/institution/mnhn/collection/p/item/
p00256012; P00256013, https://science.mnhn.fr/
institution/mnhn/collection/p/item/p00256013),
but the two last cited specimens were collected in
Timor, and they are morphologically different from
the type specimen. The sheet P05196083 contains two
gatherings, one from Timor [four fragments, left side of
the sheet: "Herb. Mus. Paris. Salsola brachyptera Moq.
(an differt ab australi Br.?) I-le Timor"] and one from
Java (one broken branch, most probably a fragment
of the type P00799151: "Herbarium Moquinianum.
Salsola brachypteris Moq. Java (Belangé, no. 4)
A. Moq.-Tand.", with a characteristic signature of
Moquin-Tandon). Both specimens have small slips
"Herb. Al. de Bunge" attached at larger labels. Most
probably these plant fragments were provided to Bunge
by Moquin-Tandon or some other colleague, and later
returned to the P herbarium through the herbarium of
E. Cosson (see the small label in the upper right corner
of the sheet). All mentioned specimens from Timor
evidently are members of Salsola sect. Salsola ("S. kali –
S. tragus – S. pontica – S. australis etc. aggregate" in
the widest possible sense) and are morphologically
different from the type from Java (P00799151, formally
designated here as the lectotype of Salsola brachypteris
Moq., because of the existence of the second specimen
of Moquin-Tandon in P: P05196083, right-hand plant,
possibly a fragment of the type).
After studying the high-resolution image of the type
of S. brachypteris, I am certain that there is not even a
slight possibility that this name is applicable to any native
Pontic-Mediterranean taxon of the Salsola pontica
group. First, judging from the available larger fragment
of P00799151, the plant itself was probably either a
robust annual or a short-lived perennial, somewhat
lignescent at base. Leaves were probably easily detached
at base, falling off (caducous) at maturity/senescence
or under unfavorable environmental conditions
(changes of draught and rainy seasons?), which was
properly noted by Moquin-Tandon (1840, 1849) in the
original and updated descriptions. Most importantly,
the branches and bases of lower leaves in that fragment
are condensed, positioned very close to each other,
with leaves subopposite or even opposite (!). That leaf
arrangement is very peculiar, almost never occurring in
most species of Salsola sect. Salsola, and it explains why
Moquin-Tandon (1835, 1840, 1849) so consistently
compared his plant to Salsola soda L. (now Soda inermis
Fourr.).
Thus, the characters observed in the plant fragments
of the type specimen and data of the protologue
and later publications indicate that S. brachypteris
is definitely not conspecific with any native Pontic-
https://plants.jstor.org/stable/viewer/10.5555/al.ap.specimen.k000899590
https://plants.jstor.org/stable/viewer/10.5555/al.ap.specimen.k000899590
https://plants.jstor.org/stable/viewer/10.5555/al.ap.specimen.k000899590
http://folio.furman.edu/botcarweb/indices?urn=urn:cite:fufolioimg:BotCarWalter.walter_v_93_01135
http://folio.furman.edu/botcarweb/indices?urn=urn:cite:fufolioimg:BotCarWalter.walter_v_93_01135
http://folio.furman.edu/botcarweb/indices?urn=urn:cite:fufolioimg:BotCarWalter.walter_v_93_01135
https://science.mnhn.fr/institution/mnhn/collection/p/item/p00256012
https://science.mnhn.fr/institution/mnhn/collection/p/item/p00256012
https://science.mnhn.fr/institution/mnhn/collection/p/item/p00256013
https://science.mnhn.fr/institution/mnhn/collection/p/item/p00256013
525Укр. бот. журн., 2017, 74(6)
or barely developed, but this character is very variable
and fruits with winged tepals (with wings up to 1 mm
wide) sometimes occur in middle and upper parts of
branches. Both erect and prostrate-ascending plants
were observed in S. pontica, but the growth habit in this
taxon greatly depends on environmental conditions.
Besides, Pallas (1803: 37) mentioned in the protologue
that his S. kali var. pontica is a prostrate plant (at least
when young): "Planta annua patentissima, diffusa, ab
imo ramosissima, junior prostrata…"
Brullo et al. (2015b: 67) also indicated that, in their
understanding, K. ponticum has "stems and leaves green,
glabrous or subglabrous…"; but Pallas (1803: 37) reported
that the whole plant is slightly hispid-pilose ("…tota
planta subpiloso-aspera…"). Both glabrous (common)
and somewhat hispid plants of S. pontica occur in the
Black Sea area, quite often in the same populations.
Because of that Tzvelev (1993: 82) reserved the name
S. pontica var. pontica for hispid plants and proposed var.
glabra Tzvelev for glabrous forms of that species, which
are more common, at least along northern shores of the
Black Sea.
Thus, the diagnostic characters given for
K. dodecanesicum by Brullo et al. (2015b) seem to be
rather elusive and/or variable. However, the rather
short, inflated and prominently succulent bracts/
bracteoles abruptly contracted into thin and quite long
spines reported for K. dodecanesicum and illustrated in
Brullo et al. (2015b: 62, Fig. 1; 64, Fig. 3) are normally
not peculiar to specimens of S. pontica from northern
shores of the Black Sea. The reported morphological
differences between the mainly Pontic (also occurring
in some localities in the Eastern Mediterranean area)
and the eastern Mediterranean races are better suited
for subspecies rather than two separate species.
But is Kali dodecanesicum really a new species?
At the end of his treatment of Salsola in De Candolle's
Prodromus, Moquin-Tandon (1849: 190) provided a list
(with descriptions and other data) of some insufficiently
known species ("Species non satis notae"), which he,
although, accepted. One of such species was Salsola
squarrosa Steven ex Moq., for which the following
information was given:
"40. S. SquarroSa (Stev. obs. ined. in herb. Willd.
1840), foliis teretiusculis oblongis mucronatis… — In
ins. Naxo. Folia carnosa, reflexa. Flores axillares. — An
S. Kali varietas?"
As we see, the species was known to Moquin-Tandon
by only one collection in the Willdenow herbarium in
Berlin (B), which was annotated by Ch. Steven. A short
diagnostic description has been published as well, so
It was commonly assumed that specimens mounted
in the Walter Herbarium folio are types and other
original specimens used by Walter for descriptions of
new taxa in his Flora Caroliniana, but that view was
challenged by Ward (2006, 2007), who claimed that
most specimens from the Walter Herbarium cannot be
regarded holotypes and in most cases are not suitable
for designation of lectotypes. He later proposed
the "neotype" for S. caroliniana (Ward, 2008: 483),
the specimen collected in South Carolina in 1939
(GH00247988, image available from https://plants.
jstor.org/stable/10.5555/al.ap.specimen.gh00247988).
That "neotypification" is, however, ineffective because
the earlier typification has been disregarded (Art. 9.19
of ICN: McNeill et al., 2012); it is also evidently in
conflict with the protologue (Art. 9.19(b) of ICN:
McNeill et al., 2012). The name S. caroliniana is not
in current use, and was used rather inconsistently in
the past. Considering these and some other arguments,
especially the history of misapplication of the name,
its uncertain taxonomic identity, and conflicting
typifications, I believe that the best solution is to reject
the name S. caroliniana. The relevant nomenclatural
proposal has been prepared (Mosyakin, manuscript
submitted to Taxon); it contains more arguments and
detailed information on the nomenclatural problem of
S. caroliniana.
What is Kali dodecanesicum C. Brullo & al.?
Brullo et al. (2015b) described from the Dodecanese
archipelago (Greece; holotype from Rhodes) a
new species, Kali dodecanesicum, and proposed to
distinguish their new taxon and Kali ponticum (Salsola
pontica) using the following characters mentioned in
the key (Brullo et al., l.c.: 67):
Kali dodecanesicum: Plant yellowish-green, prostrate,
with branches prostrate–ascending, perianth 2.5−
3.0 mm long, anthers 2 mm long, fruiting perianth
4.0−4.5 in diameter with wings 0.5−1.0 mm wide.
Kali ponticum: Plant green to green-glaucous (often
tinged with reddish), erect with branches ascending,
perianth 2.0−2.2 mm long, anthers 0.8−1.4 mm long,
fruiting perianth 3.0−3.5 in diameter with wings 0.2−
0.4 mm wide or with tubercles only (at least in Israel).
However, judging from my experience with living
plants observed in southern Ukraine (shores of the Black
Sea from Odessa Region to Crimea) and herbarium
specimens consulted mainly in KW, LE, and in several
other herbaria, plants of S. pontica from Ukraine are
normally (or at least quite often) yellowish-green or light
green; dark green individuals sometimes also occur, but
they seem to be less common. Wings are usually short
https://plants.jstor.org/stable/10.5555/al.ap.specimen.gh00247988
https://plants.jstor.org/stable/10.5555/al.ap.specimen.gh00247988
526 Ukr. Bot. J., 2017, 74(6)
available for the latter taxon; however, it is quite possible
that it is in fact conspecific with S. controversa Tod. ex
Lojac. described from Sicily (Lojacono Pojero, 1904:
271–272), which in that case will be the correct name
for this Mediterranean species (if it is indeed specifically
distinct from S. pontica), or for the whole Pontic-
Mediterranean coastal group (if treated as one species,
incl. S. pontica)". I also indicated that S. controversa
was validated by Lojacono Pojero (1904: 271–272), not
by Nyman (see also Rilke, 1999; Domina et al., 2014).
This species was firmly forgotten even by Italian authors
(probably because its name was considered invalid),
and only occasionally S. controversa was mentioned in
literature, mainly as a synonym of S. kali or S. tragus
(see, e.g., Casu, 1910). It has not been mentioned at
all in recent Italian publication on Salsola sensu lato
(Brullo et al., 2013, 2015a, b) and in the inventory of
plant species described from Italy and their loci classici
(Peruzzi et al., 2015).
Formally, all herbarium specimens distributed by
Todaro under No. 1088 (now present in many herbaria)
are syntypes of S. controversa. However, my analysis of
high-resolution images of various specimens available
online (in particular, K000899539, K000899540,
P05157673, P05344327, etc., most of them available
from JSTOR Global Plants: https://plants.jstor.org; and
especially specimens from PAL, see below) indicate that
Todaro's plants deposited in various collections are rather
diverse morphologically. They in fact represent either a
mixed collection or several gatherings, and belong to
three entities: (1) the coastal taxon (S. controversa sensu
stricto), (2) S. tragus, and (3) forms morphologically
intermediate between these taxa. Because of that,
proper typification of S. controversa becomes a crucial
issue, especially if we intend to preserve the original
application of the name established by its validating
author, Lojacono Pojero (1904).
Sukhorukov (2014: 332) listed in synonymy of
Kali pontica the name "Salsola controversa Todaro ex
Nym., Consp. Fl. Eur. 3: 631 (1881)" and designated
its lectotype, with the following citation: "Lectotypus
(Sukhorukov, designated here): Palermo, in arenosis
maritimis, IX. [sensu anno], Todaro 1088 (K-000899539 !,
iso — K-000899540 !)" (here "sensu anno" is evidently
an error, probably meaning "sine anno", "no year
indicated"). In my opinion, this lectotype designation
was a premature decision, because of the considerations
that follow. First, Sukhorukov probably has not
seen the real protologue and original description of
S. controversa, because Flora Sicula (Lojacono Pojero,
1904) is not cited anywhere in his book (Sukhorukov,
2014), while no description of the species was provided
by Nyman, who simply cited the name S. controversa
there is no doubt that the name S. squarrosa is valid.
The plant was collected in the island of Naxos (Greece).
There is just one specimen in B (B-Willd, Willdenow
Herbarium) matching the protologue of S. squarrosa
(B-W-05383010, digital images available from http://
herbarium.bgbm.org/object/BW05383010 [image ID:
325010] and http://plants.jstor.org/stable/10.5555/
al.ap.specimen.b%20-w%2005383%20-01%200).
The text on the Willdenow Herbarium folder follows:
"Pentandria Digynia / Salsola squarrosa / foliis
teretiusculis oblongis / carnosis mucronatis reflexis /
floribus axillaribus / Habitat in insula Naxos". As we
see, that text closely matches the published description
(Moquin-Tandon, 1849: 190). The herbarium sheet
bears two labels: [Label 1] "Salsola Kali L. var.! (v.
Pounge)" and [Label 2] "In insula Naxia (Schwartz)".
There are also the following texts written directly on the
sheet: "S. squarrosa. S." (top right corner) and "Schwartz
Centuria. W." (bottom right corner). The plant fragment
is ca. 17 cm long; the plant is yellowish-green, slightly
hispid or papillose; flowers/fruits condensed; bracts
and bracteoles short (probably prominently succulent
and inflated when fresh), abruptly contracted into long
subulate spines.
Judging from the high-resolution digital image of the
type (holotype) specimen, the brief original description
(Moquin-Tandon, 1849: 190), and other available
evidence (including the obvious geographic proximity
of loci classici), there should be no doubt that Salsola
squarrosa is the priority name for the taxon described
later as Kali dodecanesicum (Brullo et al., 2015b).
Moreover, S. squarrosa seems to be the earliest avail-
able binomial applicable for the whole Mediterranean
coastal complex of races known earlier as S. pontica,
S. controversa, and under the misapplied names S. tragus
auct. and Kali macrophylla sensu Galasso & Bartolucci
(2014: 83; non S. macrophylla R. Br.). Consequently, if
just one native coastal species of Salsola is recognized
in the Pontic-Mediterranean area, it should be called
S. squarrosa (sensu lato). However, before proposing a
nomenclatural and taxonomic solution, it is necessary
to consider another name applicable to Mediterranean
plants, S. controversa.
What is Salsola controversa Tod. ex Lojac.?
In my recent article (Mosyakin, 2017: 410) I
commented that the coastal species of Salsola from the
Black Sea area (which I accepted as S. pontica) "belongs
to a problematic group of Pontic-Mediterranean
coastal taxa currently known as S. pontica (either sensu
lato or sensu stricto) and Kali dodecanesicum C. Brullo,
Brullo, Giusso & Ilardi (see Brullo et al. 2015b). No
nomenclatural combination in Salsola is currently
https://plants.jstor.org/
http://herbarium.bgbm.org/object/BW05383010
http://herbarium.bgbm.org/object/BW05383010
http://plants.jstor.org/stable/10.5555/al.ap.specimen.b -w 05383 -01 0
http://plants.jstor.org/stable/10.5555/al.ap.specimen.b -w 05383 -01 0
527Укр. бот. журн., 2017, 74(6)
was accumulated for many years, but already in 1899
Churchill was unable to add specimens to his collection
because of his failing health (see relevant details in
Hemsley, 1906). Thus, it is evident that Lojacono
Pojero has never seen the specimen K000899539 (the
lectotype designated by Sukhorukov, 2014) and did not
use it when he was preparing the validating description
of S. controversa in Flora Sicula. In contrast, Lojacono
Pojero evidently used the collections of the Herbarium
Siculum in Palermo (now Herbarium Mediterraneaum
Panormitanum, PAL) until 1913, when he left Palermo
(Domina et al., 2014).
In view of the new information about patterns of
diversity of coastal Mediterranean taxa and the need
for precise application of the name S. controversa,
there are two possible ways of coping with the uncertain
situation caused by Sukhorukov's lectotypification. The
first option is to accept his lectotypification, despite its
conflict with the protologue; in that case, a second-step
lectotypification (right-hand immature fragment?) with
simultaneous epitypification is be needed. However, in
my opinion, both plants on the sheet K000899539 do not
match the protologue in many of their morphological
characters: the plant fragments are not glaucous, their
flowers/fruits are not condensed on the branches
and not arranged in very dense and large panicle-like
inflorescences, and branches are not strongly angular.
At least one (right-side) plant fragment most probably
belongs to S. tragus. Thus, the only reasonable option
under Art. 9.19 of ICN (McNeill et al., 2012) is to have
that lectotypification (Sukhorukov, 2014) superseded
because of its evident and serious enough conflict
with the protologue (Art. 9.19(b) of ICN: McNeill
et al., 2012) and to select another lectotype among
several specimens from PAL, which perfectly match
the original detailed description provided by Lojacono
Pojero (1904).
There are four Todaro's specimens in PAL
available online and identified as S. controversa, all
with printed labels (No. 1088). A PAL specimen
on sheet with ID No. 58916 (image available from:
http://147.163.105.223/herbarium_vdetails_en2.
asp?idmode=simple&id=71449) is immature;
the plant has narrow, almost filiform leaves and
most probably belongs to S. tragus sensu stricto.
Morphological characters of plant fragments of three
specimens correspond to the characters given in the
protologue. The specimen with PAL ID No. 58918
(http://147.163.105.223/herbarium_vdetails_en2.
asp?idmode=simple&id=71451) is designated here as
the lectotype of S. controversa (see below). Two other
specimens (PAL ID Nos. 58915 and 58917; see links to
images below) are considered isolectotypes.
in synonymy. Second, the sheet K000899539 (image
available from: https://plants.jstor.org/stable/10.5555/
al.ap.specimen.k000899539) evidently contains two
morphologically different plant fragments: the left-
hand plant is pale green to yellowish green (probably
the living plant was light green to yellowish green) while
the right-hand fragment is dark green. The left-hand
plant has better developed (more mature) fruits than
the right-hand one, which is only flowering. Both plant
fragments have rather lax inflorescences, with flowers/
fruits not much condensed on branches.
Lojacono Pojero (1904: 272) in his detailed original
description reported that his species is glaucous-green
("…glaucescens…"), with flowers arranged in very
dense and large panicle-like inflorescences ("…floribus
densissimis secus ramulos alternos paniculatos crebris
magnis fere contiguis…"), and with strongly angular
branches ("…ramis <…> omnibus valide angulatis…").
Thus, morphological characters of the pale green
fragment (left-hand plant, K000899539) are evidently
in conflict with the protologue. The taxonomic identity
of the right-hand (immature) fragment is problematic,
especially if we take into consideration that the specimens
distributed by Todaro under No. 1088 represent a mixed
collection or several gatherings of rather diverse plants
belonging to two or three entities (see above). However,
the right-hand fragment is morphologically very similar
to plant fragments mounted on the sheet K000899540
(reported as an isolectotype by Sukhorukov, 2014).
Plant fragments on K000899540 (image available from
https://plants.jstor.org/stable/10.5555/al.ap.specimen.
k000899540) have tepals with rather well developed
wings and, in my opinion, definitely belong to S. tragus
(as well as P05157673: https://science.mnhn.fr/
institution/mnhn/collection/p/item/p05157673; and
some other specimens of Todaro No. 1088). Characters
of the right-hand fragment of K000899539 also
contradict the protologue; in particular, in the shape
of inflorescences (terminal branches with flowers/fruits
are not particularly dense and not "paniculate"); the
plant seems to be very sparsely short-papillose, while
Lojacono Pojero reported his species as glabrous.
The specimen K000899539 has an additional label
(in the lower right corner of the sheet) "Herbarium
Kewense. Herbarium Churchillanum Proprium.
Bequeathed, 1906" indicating that it was added
to the Kew herbarium in 1906, after the death of
G.C. Churchill. He was donating parts of his private
collection to K starting from 1884, and in 1892 "in
connection with a will he thought of making, he
announced the intention he had formed of bequeathing
to Kew his European herbarium" (Hemsley, 1906:
386). The main European part of his private herbarium
http://147.163.105.223/herbarium_vdetails_en2.asp?idmode=simple&id=71449
http://147.163.105.223/herbarium_vdetails_en2.asp?idmode=simple&id=71449
http://147.163.105.223/herbarium_vdetails_en2.asp?idmode=simple&id=71451
http://147.163.105.223/herbarium_vdetails_en2.asp?idmode=simple&id=71451
https://plants.jstor.org/stable/10.5555/al.ap.specimen.k000899539
https://plants.jstor.org/stable/10.5555/al.ap.specimen.k000899539
https://plants.jstor.org/stable/10.5555/al.ap.specimen.k000899540
https://plants.jstor.org/stable/10.5555/al.ap.specimen.k000899540
https://science.mnhn.fr/institution/mnhn/collection/p/item/p05157673
https://science.mnhn.fr/institution/mnhn/collection/p/item/p05157673
528 Ukr. Bot. J., 2017, 74(6)
Nomenclature and new combinations
I recognize here three subspecies of coastal Pontic-
Mediterranean taxa known in recent publications under
many properly applied and misapplied names, such as
Salsola pontica, S. kali subsp. pontica, S. tragus subsp.
pontica, Kali ponticum, K. dodecanesicum, S. tragus auct.,
p.p., excl. pl. etc. (see Iljin, 1936, 1952; Tzvelev, 1993,
1996; Mosyakin, 1996, 2003; Rilke, 1999; Sukhorukov,
Akopian, 2013; Sukhorukov, 2014; Brullo et al., 2013,
2015a, b, and references therein).
Salsola squarrosa Steven ex Moq. in DC., Prodr.
13(2): 190. 1849. (Moquin-Tandon, 1849: 190).
Type (holotype, the only specimen cited in the
protologue): [Greece, Naxos] "in ins. Naxo"; Herb.
Willd. 5383 (B-W-05383010, images available from
http://herbarium.bgbm.org/object/BW05383010 [image
ID: 325010] and http://plants.jstor.org/stable/10.5555/
al.ap.specimen.b%20-w%2005383%20-01%200).
Salsola squarrosa subsp. squarrosa
Type: see above.
= Kali dodecanesicum C. Brullo, Brullo, Giusso &
Ilardi, Phytotaxa 218(1): 63. 2015. (Brullo et al., 2015b:
63).
Type (holotype): "GREECE. Dodecanese: Rhodes,
sandy coast near Kattavia, 35° 58’ 35’’ N, 27° 44’ 25’’ E,
22 August 2013, S. Brullo & V. Ilardi s.n." (CAT).
Salsola squarrosa subsp. controversa (Tod. ex Lojac.)
Mosyakin, comb. nov.
Basionym: Salsola controversa Tod. ex Lojac.,
Fl. Sicula 2(2): 271. 1904. (Lojacono Pojero, 1904: 271,
description on page 272).
Type (lectotype, designated here): "Todaro. Flora Sicula
Exsiccata. Salsola controversa Tod. enum. fl. sic. ined.
In arenosis maritimis—Palermo. Septembri. n. 1088.
Legit Todaro" (PAL ID No. 58918, image available
from http://147.163.105.223/herbarium_vdetails_en2.
asp?idmode=simple&id=71451). Isolectotypes: PAL ID
No. 58915 (http://147.163.105.223/herbarium_vdetails_
en2.asp?idmode=simple&id=71448); PAL ID No.
58917 (http://147.163.105.223/herbarium_vdetails_en2.
asp?idmode=simple&id=71450); P05344327 (https://
science.mnhn.fr/institution/mnhn/collection/p/item/
p05344327).
The earlier lectotypification (Sukhorukov, 2014: 332)
is superseded following Art. 9.19(b) of ICN (McNeill
et al., 2012).
Salsola squarrosa subsp. pontica (Pall.) Mosyakin,
comb. nov.
Basionym: Salsola kali L. var. pontica Pall., Ill. Pl.:
37. 1803. (Pallas, 1803: 37).
A taxonomic and nomenclatural solution for Pontic-
Mediterranean coastal taxa of Salsola
Unfortunately, Brullo et al. (2013, 2015a, b) and
almost all other authors who recently commented on
taxonomy of Pontic-Mediterranean taxa of Salsola
(e. g., Sukhorukov, 2014) did not mention an interesting
article on phylogeography of some European coastal
plants (Kadereit et al., 2005), partly based on results
from the dissertation by Arafeh (2005). That article
provided noteworthy data and conclusions on the
genetic and geographical differentiation of five coastal
species, including the taxon listed under the name
Salsola kali. Their molecular phylogeography results
based on AFLP evidence clearly indicate that three
rather distinct but closely related lineages of coastal
Salsola exist in the Pontic-Mediterranean area.
Unfortunately, Kadereit et al. (2005) refrained from
discussing taxonomy of the taxa involved and made no
attempt to match the revealed lineages with any existing
names in Salsola.
The studied taxon (in fact, taxa) was accepted as
S. kali sensu latissimo, including at least five entities
corresponding, in my opinion, to S. kali sensu stricto
(Atlantic and Baltic race), S. tragus (inland weedy race),
S. controversa (western Mediterranean), S. squarrosa
(= Kali dodecanesicum; eastern Mediterranean), and
S. pontica sensu stricto (mainly Pontic, with extensions
to the eastern Mediterranean area). Following these
molecular results and the taxonomic discussion
provided above, one may prefer to accept S. controversa,
S. squarrosa, and S. pontica as separate species.
However, considering the evident genetic similarity
(Kadereit et al., 2005) and blurred morphological and
geographical borders of these three coastal taxa, I prefer
to treat them as three subspecies of S. squarrosa (see
new combinations below).
Further detailed morphological, molecular
phylogenetic, and phylogeographic studies of coastal
Eurasian and all Australasian species of Salsola
are needed, preferably in comparison with data on
morphologically diverse S. tragus and some other
Eurasian inland taxa. Reliable morphological characters
of the discussed taxa should be further specified using a
wider geographical coverage and the population-based
approach. It should be also taken into consideration that
plants of S. tragus quite often co-occur with S. squarrosa
sensu lato in maritime coastal habitats (at least in the
Black Sea area, but probably also in the Mediterranean
region: see comments above, under S. controversa),
where they often develop rather thick and fleshy leaves,
and can be thus confused with the coastal taxa.
http://herbarium.bgbm.org/object/BW05383010
http://plants.jstor.org/stable/10.5555/al.ap.specimen.b -w 05383 -01 0
http://plants.jstor.org/stable/10.5555/al.ap.specimen.b -w 05383 -01 0
http://147.163.105.223/herbarium_vdetails_en2.asp?idmode=simple&id=71451
http://147.163.105.223/herbarium_vdetails_en2.asp?idmode=simple&id=71451
http://147.163.105.223/herbarium_vdetails_en2.asp?idmode=simple&id=71448
http://147.163.105.223/herbarium_vdetails_en2.asp?idmode=simple&id=71448
http://147.163.105.223/herbarium_vdetails_en2.asp?idmode=simple&id=71450
http://147.163.105.223/herbarium_vdetails_en2.asp?idmode=simple&id=71450
https://science.mnhn.fr/institution/mnhn/collection/p/item/p05344327
https://science.mnhn.fr/institution/mnhn/collection/p/item/p05344327
https://science.mnhn.fr/institution/mnhn/collection/p/item/p05344327
529Укр. бот. журн., 2017, 74(6)
Fachbereich Biologie der Johannes Gutenberg-
Universität in Mainz], Mainz, 2005, 84 pp.
Ayres D., Ryan F.J., Grotkoo E., Bailey J., Gaskin J.
Tumbleweed (Salsola section Kali) species and speciation
in California. Biol. Invasions, 2009, 1: 1175–1187.
Backer C.A. Chenopodiaceae. In: Flora Malesiana, series
1: Spermatophyta. Ed. C.G.G.J. van Steenis. Batavia:
Noordhoff-Kolff N.V., 1949, vol. 4(2), pp. 99–106.
Bentham G. Chenopodiaceae. In: Flora Australiensis: a
description of the plants of the Australian territory. London:
Reeve and Co., 1870, vol. 5, pp. 150–208.
Boerlage J.G. Handleiding tot de kennis der flora van
Nederlandsch Indië. Beschrijving van de families
en geslachten der Nederl. Indische phanerogamen.
Leiden: E.J. Brill, 1900, vol. 3(1), xxxi + 418 pp.
Borger C.P.D., Yan G., Scott J.R., Walsh M., Powles S.B.
Salsola tragus or S. australis (Chenopodiaceae) in
Australia – Untangling the taxonomic confusion through
random amplified microsatellite polymorphism (RAMP)
and cytological analysis. Austral. J. Bot., 2008, 56: 600–
608. http://dx.doi.org/10.1071/BT08043
Botschantzev V.P. A synopsis of Salsola (Chenopodiaceae)
from South and South-West Africa. Kew Bull., 1974, 29:
597–614. http://dx.doi.org/10.2307/4108004
Brullo C., Brullo S., Giusso del Galdo G., Guarino R.,
Iamonico D. Il genere Salsola s. l. (Chenopodiaceae) in
Italia. In: Contributi alla ricerca floristica in Italia. Eds S.
Peccenini, G. Domina. Firenze: Soc. Bot. Italiana, 2013,
pp. 35–38.
Brullo C., Brullo S., Gaskin J.F., Giusso del Galdo G.,
Hrusa G.F., Salmeri C. A new species of Kali
(Salsoloideae, Chenopodiaceae) from Sicily, supported by
molecular analysis. Phytotaxa, 2015a, 201(4): 256–277.
http://dx.doi.org/10.11646/phytotaxa.201.4.2
Brullo C., Brullo S., Ilardi V., Giusso del Galdo G. Kali
dodecanesicum (Chenopodiaceae, Salsoloideae) a new
species from Greece. Phytotaxa, 2015b, 218(1): 61–68.
http://dx.doi.org/10.11646/phytotaxa.218.1.4
Casu A. Salsola Kali L. e Salsola Tragus L.: Specie critiche.
Memorie della Reale Accademia delle Scienze di Torino,
Ser. 2, 1910, 60: 393–407.
Chinnock R.J. Some observations on Salsola L.
(Chenopodiaceae) in Australia. J. Adelaide Bot. Gard.,
2010, 24: 75–79.
Domina G., Greuter W., Mazzola P., Raimondo F.M.
Names of Italian vascular plants published by Michele
Lojacono Pojero. Fl. Medit. 2014, 24: 215–232. http://
dx.doi.org/10.7320/FlMedit24.215
Fourreau M. Catalogue des plantes du cours du Rhone (suite
et fin). Annales de la Société Linnéenne de Lyon (nouvelle
série), 1869, 17: 89–200.
Galasso G., Bartolucci F. Notula: 2070 (Kali macrophyllum,
comb. nov.) / Notulae alla checklist della flora vascolare
italiana: 17 (2027–2070). Inform. Bot. Ital., 2014, 46(1):
83.
Hemsley W.B. [under initials "W. B. H."; an untitled note
about G.C. Churchill in:] Miscellaneous notes. LIV
[No. 54]. Bull. of Miscellaneous Information (Royal
Botanic Gardens, Kew), 1906, [Vol. of 1906] No. 9:
384–392.
Type (lectotype, designated by Rilke, 1999: 133):
[Ukraine, Crimea, Sudak] "Sudagh. Herb. Pallas"
(BM000016635, image available from: https://
plants.jstor.org/stable/10.5555/al.ap.specimen.
bm000016635).
≡ Salsola ruthenica Iljin subsp. pontica (Pall.) Iljin,
Sorn. rast. SSSR [Сорные растения СССР] 2: 140.
1934. ≡ S. pontica (Pall.) Iljin, Fl. URSS [Флора СССР]
6: 212. 1936, nom. inval. ≡ S. pontica (Pall.) Degen, Fl.
Velebit. 2: 47. 1937. ≡ S. kali L. subsp. pontica (Pall.)
Mosyakin, Ann. Missouri Bot. Gard. 83: 389. 1996. ≡
S. tragus L. subsp. pontica (Pall.) Rilke, Biblioth. Bot.
149: 133. 1999. ≡ Kali ponticum (Pall.) Sukhor., Novosti
Sist. Vyssh. Rast. [Новости систематики высших
растений] 42: 106. 2011, as "pontica". ≡ Kali tragus (L.)
Scop. subsp. ponticum (Pall.) Mosyakin, Ukrayins'k.
Bot. Zhurn. [Український ботанічний журнал]
69(3): 395. 2012, as "pontica".
Acknowledgments
I am grateful to Prof. Helmut Freitag (University of Kassel,
Germany), Dr. Ahmet Emre Yaprak (Ankara University,
Turkey), and Dr. Leila M. Shultz (Utah State University,
Logan, Utah, USA), with whom I discussed some issues of
taxonomy and nomenclature of Salsola sensu lato. Thanks
are due to an anonymous reviewer for his/her comments
on the manuscript. Victoria McMichael and Mary
Stiffler (Peter H. Raven Library, Missouri Botanical
Garden, St. Louis, Missouri, USA) provided copies of
some rare publications, and their kind help is gratefully
acknowledged. Herbarium assistance of Dr. Natalia
M. Shiyan (Head Curator, National Herbarium of
Ukraine – KW, M.G. Kholodny Institute of Botany
of the National Academy of Sciences of Ukraine) and
Alisa V. Shumilova (Curatorial Assistant, KW) is greatly
appreciated.
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531Укр. бот. журн., 2017, 74(6)
видом S. australis R. Br. Ідентичність S. brachypteris Moq.
(описаний з о. Ява, Індонезія) лишається невизначеною;
судячи з наявних даних, цей таксон, ймовірно, спорідне-
ний з S. macrophylla або з деякими іншими недостатньо
відомими австралазійськими видами. Стисло розглянута
проблема типіфікації та застосування назви S. caroliniana
Walter; зроблено висновок, що цю назву слід запропону-
вати до відхилення. Наголошено на потребі проведення
подальших морфологічних, молекулярно-філогенетич-
них та філогеографічних досліджень приморських євра-
зійських та австралазійських видів Salsola.
Ключові слова: Salsola, Chenopodiaceae, Salsoloideae,
номенклатура, систематика, біогеографія
Мосякин С.Л. Таксономические и номенклатурные
заметки о понтическо-средиземноморских и некоторых
австралазийских таксонах Salsola (Chenopodiaceae).
Укр. бот. журн., 2017, 74(6): 521–531.
Институт ботаники им. Н.Г. Холодного НАН Украины
ул. Терещенковская, 2, Киев 01004, Украина
Проанализированы современные и исторические взгля-
ды на систематику и номенклатуру понтическо-среди-
земноморских и некоторых австралазийських таксонов
рода Salsola (Chenopodiaceae). Обсуждены систематика
и номенклатура нескольких названий, которые ранее
применялись к представителям видовой группы, кото-
рая признавалась в недавних публикациях преимуще-
ственно под названием Salsola pontica (Pall.) Degen s. l.
Показано, что Kali dodecanesicum C. Brullo & al. являет-
ся синонимом ранее описанного вида Salsola squarrosa
Steven ex Moq.; последнее название является приори-
тетным в ранге вида для всей понтическо-средиземно-
морской группы таксонов. Эта группа представлена за-
падносредиземноморской (S. controversa Tod. ex Lojac.),
восточносредиземноморской (S. squarrosa s. str.) и преи-
мущественно черноморско-каспийской (S. pontica s. str.)
географическими расами. Учитывая нечеткие морфоло-
гические и географические границы между этими таксо-
нами, их целесообразно рассматривать как три подвида
вида S. squarrosa: subsp. controversa (Tod. ex Lojac.) Mo-
syakin, comb. nov., subsp. squarrosa, и subsp. pontica (Pall.)
Mosyakin, comb. nov., соответственно. Показано, что вид
S. macrophylla R. Br. (описан из Австралии) отличается
от всех понтическо-средиземноморских прибрежных
таксонов; он, очевидно, является родственным при-
знанному сейчас виду S. australis R. Br. Идентичность
S. brachypteris Moq. (описан с о. Ява, Индонезия) оста-
ется неопределенной; судя по имеющимся данным, этот
таксон, вероятно, родственен S. macrophylla или некото-
рым другим недостаточно изученным австралазийским
видам. Кратко рассмотрена проблема типификации и
применения названия S. caroliniana Walter; сделан вывод
о том, что это название следует предложить номенкла-
турно отвергнуть. Отмечена необходимость проведения
дальнейших морфологических, молекулярно-филогене-
тических и филогеографичних исследований примор-
ских евразийских и австралазийских видов Salsola.
Ключевые слова: Salsola, Chenopodiaceae, Salsoloideae,
номенклатура, систематика, биогеография
photosynthesis. J. Experim. Bot., 2013, 64: 3583–3604.
http://dx.doi.org/10.1093/jxb/ert191
Walter T. Flora Caroliniana. London, 1788. 263 pp.
Ward D.B. Thomas Walter Typification Project, I:
Observations on the John Fraser folio. Sida, 2006, 22:
1111–1118.
Ward D.B. The Thomas Walter Herbarium is not the
herbarium of Thomas Walter. Taxon, 2007, 56(3):
917–926.
Ward D.B. Thomas Walter Typification Project, V: Neotypes
and epitypes for 63 Walter names, of genera D through Z.
J. Bot. Res. Inst. Texas, 2008, 2: 475–486.
Wen Zh.-B., Zhang M.-L., Zhu G.-L., Sanderson S.C.
Phylogeny of Salsoleae s. l. (Chenopodiaceae) based
on DNA sequence data from ITS, psbB–psbH,
and rbcL, with emphasis on taxa of northwestern
China. Pl. Syst. Evol., 2010, 288: 25–42.
http://dx.doi.org/10.1007/s00606-010-0310-5
Wen Z.B., Zhang M.L. Anatomical types of leaves
and assimilating shoots and carbon 13C/12C isotope
fractionation in Chinese representatives of Salsoleae
s. l. (Chenopodiaceae). Flora – Morphology, Distribution,
Functional Ecology of Plants, 2011, 206(8): 720–730.
https://doi.org/10.1016/j.flora.2010.11.015
Wilson P.G. Chenopodiaceae. In: Flora of Australia. Ed.
A.S. George. Canberra: Austral. Government Publ.
Service, 1984, vol. 4, pp. 81–317.
Recommended for publication by Submitted 10.11.2017
Zigmantas Gudžinskas
Мосякін С.Л. Таксономічні та номенклатурні нотатки
про понтично-середземноморські та деякі австралазійські
таксони Salsola (Chenopodiaceae). Укр. бот. журн., 2017,
74(6): 521–531.
Інститут ботаніки ім. М.Г. Холодного НАН України
вул. Терещенківська, 2, Київ 01004, Україна
Проаналізовано сучасні та історичні погляди щодо сис-
тематики та номенклатури понтично-середземномор-
ських та деяких австралазійських таксонів Salsola (Che-
nopodiaceae). Обговорюється таксономія та номенклату-
ра декількох назв, які раніше застосовувалися до пред-
ставників видової групи, що здебільшого була визнана
у недавніх публікаціях під назвою Salsola pontica (Pall.)
Degen s. l. Показано, що Kali dodecanesicum C. Brullo & al.
є синонімом раніше описаного виду Salsola squarrosa Ste-
ven ex Moq.; остання назва є пріоритетною у ранзі виду
для усієї понтично-середземноморської групи таксонів.
Ця група представлена західносередземноморською
(S. controversa Tod. ex Lojac.), східносередземноморською
(S. squarrosa s. str.) та переважно чорноморсько-каспій-
ською (S. pontica s. str.) географічними расами. Врахову-
ючи нечіткі морфологічні та географічні межі між цими
таксонами, їх доцільно розглядати як три підвиди виду
S. squarrosa: subsp. controversa (Tod. ex Lojac.) Mosyakin,
comb. nov., subsp. squarrosa, та subsp. pontica (Pall.) Mosya-
kin, comb. nov., відповідно. Показано, що вид S. macro-
phylla R. Br. (описаний з Австралії) є відмінним від усіх
понтично-середземноморських прибережних таксонів;
натомість він, очевидно, споріднений з нині визнаним
http://dx.doi.org/10.1093/jxb/ert191
http://dx.doi.org/10.1007/s00606-010-0310-5
https://doi.org/10.1016/j.flora.2010.11.015
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