A synopsis of Euphorbia (Euphobiaceae) for the Caucasus Обзор видов Euphorbia (Euphorbiaceae) Кавказа

Abstract

Euphorbia L. (Euphorbiaceae), one of the largest genera of angiosperms, is represented in the Caucasus by 80 species. This paper contains a taxonomic treatment for these species and a key for their determination. For every species the following information is provided: accepted name and major synonyms with typification where possible; brief characteristics of habitats; distribution in the Caucasus and indication of geographical element (for indigenous species); and taxonomic and geographical notes if necessary. Lectotypes of Euphorbia arvalis Boiss. et Heldr., E. damascena Boiss., E. fossulata Boiss. et Gaill., E. iberica Boiss. var. intermedia Boiss., E. kemulariae Ter-Chatsch., E. notadenia Boiss. et Hohen., E. platyphyllos L., E. paralias L. and E. ruderalis Sheele are newly designated. The indigenous component of Caucasian Euphorbia comprises 73 species. The majority belong to three geographical elements: Euro-Siberian (41.10%), Irano-Turanian (20.54%) and Submediterranean (15.07%). There are also 7 alien species mostly of North American origin. Taxonomically, most of Caucasian Euphorbia species (70 or 87.5%) belong to subgen. Esula and represent 16 of the 21 currently accepted sections. The remaining ten species belong to subgen. Chamaesyce.

Full text

A synopsis of Euphorbia (Euphobiaceae) for the Caucasus
Обзор видов Euphorbia (Euphorbiaceae) Кавказа
D. V. Geltman
Поступила в редакцию | Submitted: 29.07.2020 Принята к публикации | Accepted: 30.11.2020
D. V. Geltman
Komarov Botanical Institute of the Russian Academy of Sciences
Professora Popova Str., 2, St. Petersburg, 197376, Russia
geltman@binran.ru
Д. В. Гельтман
Ботанический институт им. В.Л. Комарова РАН
ул. Профессора Попова, 2, Санкт-Петербург, 197376, Россия
geltman@binran.ru
https://doi.org/10.31111/novitates/2020.51.43
Abstract. Euphorbia L. (Euphorbiaceae), one of the largest genera of angiosperms, is represented in the Cau casus by 80 species.
This paper contains a taxonomic treatment for these species and a key for their determination. For every species the following
information is provided: accepted name and major synonyms with typifi cation where possible; brief characteristics of habitats;
distribution in the Caucasus and indication of geographical element (for indigenous species); and taxonomic and geographical
notes if necessary. Lectotypes of Euphorbia arvalis Boiss. et Heldr., E.damascena Boiss., E. fossulata Boiss. et Gaill., E.iberica
Boiss. var. intermedia Boiss., E.kemulariae Ter-Chatsch., E.notadenia Boiss. et Hohen., E.platyphyllos L., E.paralias L. and E.ru-
deralis Sheele are newly designated. The indigenous component of Caucasian Euphorbia comprises 73 species. The majority be-
long to three geographical elements: Euro-Siberian (41.10%), Irano-Turanian (20.54%) and Submediterranean (15.07%). There
are also 7 alien species mostly of North American origin. Taxonomically, most of Caucasian Euphorbia species (70 or 87.5%)
belong to subgen. Esula and represent 16 of the 21 currently accepted sections. The remaining ten species belong to subgen.
Chamaesyce.
Keywords: Euphorbia, spurge, Caucasus, taxonomy.
Аннотация. Euphorbia L. (Euphorbiaceae)— один из крупнейших родов цветковых растений— представлен на Кавказе
80 видами. Статья включает ключ для их определения и таксономическую обработку, в которой для каждого вида при-
водится принятое название, важнейшие синонимы (по возможности с типификацией), краткая характеристика место-
обитаний, распространение на Кавказе, для аборигенных видов— географический элемент, необходимые примечания.
Обозначены лектотипы Euphorbia arvalis Boiss. et Heldr., E.damascena Boiss., E.fossulata Boiss. et Gaill., E.iberica Boiss.
var. intermedia Boiss., E.kemulariae Ter-Chatsch., E.notadenia Boiss. et Hohen., E. platyphyllos L., E.paralias L. и E.rudera-
lis Sheele. Аборигенная фракция включает 73 вида; большинство их принадлежит к 3 географическим элементам: евро-
сибирскому (41.10%), ирано-туранскому (20.54%) и субсредиземноморскому (15.07%). Семь видов являются чужерод-
ными, большинство их имеет североамериканское происхождение. Таксономически большинство кавказских Euphorbia
принадлежат к подроду Esula (70, или 87.5%) и относятся к 16 секциям из 21 принятой в настоящее время. К подроду
Chamaesyce принадлежат 10 видов.
Ключевые слова: Euphorbia, молочай, Кавказ, систематика.
Euphorbia L. (Euphorbiaceae) is one of the largest
genera of angiosperms. Its species occur almost
worldwide, from the moist tropics to the Polar Circle,
and from lowlands to extreme high mountain areas.
Preparation of a complete monograph for the genus,
or even a synopsis, is an ambitious task which can be
achieved only by joint eff orts of several research teams
working in diff erent parts of the world. Signifi cant
steps towards solving this task were done during
the implementation of the “Euphorbia planetary
biodiversity” project (Riina, Berry, 2020).
The Caucasus, a mostly mountainous area located
between the Black and the Caspian Seas, is now
recognized as one of 36 world biodiversity hotspots
(Hoff mann et al., 2016) with the number of vascular
plant species between 5500 (Menitsky, Popova, 2006)
and 6350 (Gagnidze et al., 2000). The aim of this
paper is to present a synopsis of Euphorbia for this
area that could be a signifi cant step both towards the
preparation of a worldwide Euphorbia account and to
the completion of a regional taxonomic monograph.
Background to this study
Marschall von Bieberstein (1808) published the fi rst
more or less comprehensive synopsis of vascular plants
of Crimea and the Caucasus in which he listed 24 Eu-
phorbia species. Lipsky (1899) in his mostly compilato-
ry “Flora of the Caucasus” cited 42 species. The famous
researcher of the Caucasian fl ora, A. A. Grossheim in
the fi rst edition of his “Flora of the Caucasus” (Gross-
2020
51: 43–78
Новости систематики высших растений
Novitates Systematicae Plantarum Vascularium
ISSN 0568-5443 (print)
ISSN 2687-1564 (online)

heim, 1932) listed 66 species, and later in the “Guide to
plants of the Caucasus” (Grossheim, 1949) 64 species.
Prokhanov (1949) listed 159 species in his Euphor-
bia account for the “Flora of the USSR”, with 63 of
them recorded for the Caucasus. This treatment until
now has been a very important source for Euphorbia
taxonomy because of the excellent morphological de-
scriptions and accompanying comments. The Caucasian
species in this treatment belong to subgenus Esula Pers.
(then treated as subgen. Paralias (Raf.) Prokh.), with
10 sections, and to subgen. Chamaesyce Raf.
Tamamschjan (1962) prepared the Euphorbia
treatment for the second edition of Grossheim’s “Flora
of the Caucasus”. She recorded 68 species and unlike
Prokhanov (1949) did not accept subgenera, instead
following the sectional subdivisions of Boissier (1862).
For every species, a short description and detailed
distribution in the Caucasus is provided as well as
distribution maps for almost all species.
Publication of the “Flora of the USSR” stimulated
preparation of “Floras” of Soviet republics (now
independent states) including those of Transcaucasia.
The Euphorbiaceae treatments for the “Floras” of
Georgia (two editions: Ter-Chatschaturova (1950);
Chinthibidze, Gvinianidze (1983)), Azerbaij an
(Khalilov, 1955) and Armenia (Ter-Chatschaturova,
Tamamschjan, 1973) generally followed Prokhanov
(1949), but contained important additional
information, especially on species distribution because
they were based on regional herbarium collections that
were not fully consulted in the preparation of the “Flora
of the USSR”. Two editions of “Flora of Abkhazia”
(Kolakovsky, 1948, 1982) cover very important areas
with several endemic and subendemic species. Recently,
new synopses on the genus for Armenia (Geltman,
Tamanyan, 2016) and Azerbaij an (Geltman, 2016)
were published. The Russian part of the Caucasus was
covered by Galuschko (1980), but, unfortunately,
not all of his records are confi rmed by herbarium
specimens. Some important novelties in the distribution
of Euphorbia in the Russian part of the Caucasus
are found in recent regional “Floras” and checklists
(Ivanov, 1997; Zernov, 2000, 2006; Murtazaliev, 2009;
Zernov, Onipchenko, 2011; Shilnikov, 2010, etc.).
Recently Ivanov (2019) tried to summarize data on
the Russian Caucasus. For spurges he accepted genera
Agaloma Raf., Chamaesyce Raf., Poinsettia Graham and
Tithymalus Gaertn.; unfortunately, in this treatment
he repeated some evident mistakes made by Galushko
(1980) and published several invalid nomenclatural
combinations.
Since the late 1980s I have been working on the
Euphorbia taxonomy for temperate Eurasia, with
special reference to the Caucasus. As a result, reviews
of sections Peplus Lázaro (Geltman, 2000), Esula
(Pers.) Dumort. (Geltman, 2002) and Paralias Dumort.
(Geltman, 2005) (according to the Prokhanov’s (1949)
system) for the Caucasus, Crimea and Asia Minor were
published. These long-term studies were summarized
in the Euphorbiaceae account for the “Caucasian
fl ora conspectus” (Geltman, 2012). This current
paper is based mainly on that treatment, but contains
some novelties in species composition, as well as a
determination key, data on habitats, and more extensive
synonymy and commentary.
Material and methods
The area of study follows the limits accepted in
the “Caucasian fl ora conspectus” (Takhtajan, 2003).
The northern border follows the southern limit of
Europe as usually accepted in the Russian geographical
tradition. The southern border coincides with frontiers
of Georgia, Armenia and Azerbaij an with Turkey and
Iran. It is important to mention that these limits diff er
from the circumscription of the Caucasian ecoregion
according to the World Wildlife Foundation (WWF)
and the area accepted in Solomon et al. (2014).
The work is based foremost on the Euphorbia
holdings of the Caucasian section of the Herbarium of
the Komarov Botanical Institute (LE) — about 3500
specimens. Material of other herbaria having important
Caucasian collections also have been carefully studied
(BAK, ERE, KBHG, KW, LENUD, MHA, MW, TBI,
TGM, SBG, SUCH, PGFA). For typifi cation, data of
other herbaria (BM, G, K, P, W, etc.) as well as internet
resources on type specimens (mainly Global Plants
on JSTOR — plants.jstor.org and Virtual Herbaria—
https://herbarium.univie.ac.at/database/search.php)
were important. I also made several fi eld trips to various
parts of the Caucasus in 1988–1991, 2002–2004, and
2015–2016.
Morphological terminology used in the key, espe-
cially that about synfl orescences, generally follows
Riina et al. (2013: Fig.3). Subgeneric and sectional de-
limitations follow those of Yang et al. (2012) and Riina
et al. (2013), with minor modifi cations. For proper cita-
tions of protologues, consulting the Euphorbiaceae ac-
count of the “World checklist of selected plant families”
by Govaerts et al. (2020) was very important.
The taxonomic treatment includes all indigenous
and alien species (recorded at least once) as well as
ornamental plants with tendency to naturalization.
For each species, the following information is provided:
accepted name and synonyms related to the area in
question, with typifi cation whenever possible; brief
characteristics of habitats; distribution in the Caucasus
D. V. Geltman
Новости систематики высших растений | Том 51 | 2020
44

Fig. 1. Scheme for indicating the geographical distribution (accoring to Menitsky (1991)and Takhtajan (2003).
1. WCC— Western Ciscaucasia: 1а. Az.-Kub.— Azov-Kuban, 1b.W.Stavr.— Western Stavropol; 2.ECC— Eastern Ciscauca-
sia: 2a.E. Stavr.— Eastern Stavropol, 2b.Ter.-Kuma— Terek-Kuma, 2c.Ter.-Sul.— Terek-Sulak; 3.WC— Western Caucasus:
3a.Adag.-Pshish— Adagum-Pshish, 3b.Bel.-Laba— Belaja-Laba, 3c.Urup-Teb.— Urup-Teberda, 3d.U.Kub.— Upper Kuban;
4.CC— Central Caucasus: 4a.U.Kuma— Upper Kuma, 4b.Malka— Malka, 4c.U.Ter.— Upper Terek; 5.EC— Eastern Caucasus:
5a.Assa-Arg.— Assa-Argun, 5b.U.Sulak— Upper Sulak, 5c.Man.-Samur— Manas-Samur, 5d.Kubin.— Kubinsky; 6.NWTC—
North-Western Transcaucasia: 6a.Anapa-Gel.— Anapa-Gelendzhik, 6b.Pshada-Dzhubga— Pshada-Dzubga; 7.WTC— Western
Transcaucasia: 7a.Tuap.-Adl.— Tuapse-Adler, 7b.Abkh.— Abkhasia, 7c.Ing.-Rioni— Inguri-Rioni, 7d.Rioni-Kvir.— Rioni-Kviri-
li, 7e.Adzh.— Adzharia; 8.CTC— Central Transcaucasia: 8a.Kart.-S.Oss.— Karthalinia-South Ossetia, 8b.Trial.-L.Kart.— Tri-
aletia-Lower Karthalinia, 8c.Lori— Lori; 9.ETC— Eastern Transcaucasia: 9a.Alaz.-Agrich.— Alazan-Agrichay, 9b.Shirv.— Shir-
van, 9c.Iori-Sheki— Iori-Sheki, 9d.Murgh.-Murovd.— Murghuz-Murovdagh, 9e.L.Kura— Lower Kura, 9f.Karab.— Karabagh;
10.SWTC— South-Western Transcaucasia: 10a.Meskh.— Meskhetia, 10b.Dzhav.-U.Akh. — Dzhavachetia-Upper Akhurjan,
10c.Arag.— Aragatz; 11.STC— Southern Transcaucasia: 11a.Erev.— Erevan, 11b.Sevan— Sevan, 11c.Dar.— Daraleghiz,
11d.Nakh.— Nakhitshevan, 11e.Zang.— Zangezur, 11f.Megri-Zan.— Megri-Zangelan, 11g.S.Karab.— SouthernKarabagh;
12.T— Talysch.
A synopsis of Euphorbia (Euphobiaceae) for the Caucasus
Novitates Systematicae Plantarum Vascularium | Volume 51 | 2020
45

according to the scheme adopted in the “Caucasian fl ora
conspectus” (Takhtajan, 2003) (Fig. 1)1 followed by
indication of the level 3 units of “World geographical
scheme for recording plant distribution” (Brummitt,
2001)2 and for indigenous species— indication of their
geographical element according to Geltman (2015a);
and fi nally, any pertinent taxonomic or geographical
notes. All photos of the plants are made by the author.
Determination key of Caucasian Euphorbia species
1. Upper stem, ray and raylet leaves with wide white margin
............................................................................ 80. E.marginata.
+ Upper stem, ray and raylet leaves concolorous, without
white margin .............................................................................. 2.
2. Cyathia with a single gland .............................. 79. E.davidii.
+ Cyathia with 4–5 glands ........................................................ 3.
3. Stem leaves without stipules, usually alternate, rarely op-
posite. Cyathia in prominent rays, forming apical pseu-
doumbel (rarely reduced to a single cyathium), surround-
ed by ray leaves, and also located in axils of stem leaves.
Cyathial glands with or without horn-like appendages ......
........................................................................................................ 4.
+ Leaves opposite, with stipules, cyathia axillary or in short
axillary sprouts, cyathial glands with petaloid appendages
...................................................................................................... 76.
4. Stem leaves opposite; capsules indehiscent, with spongy
mesocarp ................................................................. 1. E.lathyris.
+ Stem leaves alternate; capsules dehiscent, mesocarp not
spongy ......................................................................................... 5.
5. Perennials or small shrubs ..................................................... 6.
+ Annuals or biennials .............................................................. 55.
6. Caruncle at least 2/5 total length of seed .................................
........................................................................... 43. E.heteradena.
+ Caruncle small, less than 2/5 total length of seed ............. 7.
7. Cyathial glands semilunate, always with more or less
prominent horn-like appendages .......................................... 8.
+ Cyathial glands elliptic, oblong-elliptic, reniform or trap-
ezoid, sometimes with or without horn-like appendages ....
...................................................................................................... 25.
8. Raylet leaves free ...................................................................... 9.
+ Raylet leaves connate in pairs ............................................. 22.
9. Stem leaves 5 or more times as long as wide, linear,
oblong-linear, narrow oblanceolate or narrow elliptic ... 10.
+ Stem leaves 2.5–5(8) times as long as wide, ovate, elliptic,
oblong, oblanceolate, oblong-linear ................................... 14.
1 This scheme was developed by Menitsky (1991) with con-
sultations from A. L. Takhtajan. It was an attempt to combine a
natural division of the Caucasus (fi rst of all, mountain systems
and river basins) with existing administrative borders. Initially
it was based on the administrative divisions of the former So-
viet Union. For its adaptation to modern administrative divi-
sions and relation to other systems used for the recording of
plant distribution in the Caucasus, see https://www.binran.
ru/resursy/informatsionnyye-resursy/tekuschie-proekty/
caucasian-fl ora/contentkav/departments.php
2 NCS — Northern Caucasus (territory of the Russian Federa-
tion in the Caucasus); TCS — Transcaucasia (all other coun-
tries).
10. Stems (15)20–60 cm tall, 2–6 mm thick. Stem leaves
light-green, raylet leaves yellow or green-yellow when
plants are in fl ower, diff erent in colour from stem leaves ....
.................................................................................. 62. E.virgata.
+ Stems (3)5–40 cm tall, up to 2 (2.5) mm thick. Stem
leaves grey-green, raylet leaves concolorous with stem
leaves when plants are in fl ower ......................................... 11.
11. Plants with axillary vegetative sprouts and/or numerous,
long axillary rays .................................................................... 12.
+ Plants without axillary vegetative sprouts, sometimes
with few, short axillary rays ................................................. 13.
12. Stems (8)10–20(30) cm tall and (1)1.5–2 mm thick.
Apical rays (4)5–10. Axillary sprouts without rays. Root
more or less horizontal, nearly the same thickness as stem
at bottom. Plants of mountain areas .......................................
....................................................................... 66. E.daghestanica.
+ Stems 5–15 cm tall and 1–1.5(2.5) mm thick. Apical rays
3(4). Axillary sprouts often with their own axillary and
apical rays. Root vertical, noticeably thicker than stem at
bottom. Plants of lowlands of Ciscaucasia ..............................
........................................................................ 68. E.astrachanica.
13 (11). Stem leaves linear or linear-subulate, shortly acute at
apex, (3)4–7 cm long, 1–2(3) mm wide, (12)15–30(40)
times as long as wide .................................... 67. E.leptocaula.
+ Stem leaves linear, narrow elliptic or narrow oblanceolate,
shortly acute or rounded at apex, 2.5–4 cm long, 2–4 mm
wide, 6–15 times as long as wide ..................... 65. E.subtilis.
14 (9). Stems (15)20–120 cm tall, 2–12 mm thick. Stem
leaves 2–12 cm long, bright-green or grey-green, raylet
leaves yellow, green-yellow or grey-green when plants are
in fl ower, concolorous or not concolorous with stem leaves
...................................................................................................... 15.
+ Stems (3)5–20 cm tall, up to 2 mm thick. Stem leaves 0.5–
3 cm long, grey-green, raylet leaves concolorous with stem
leaves when plants are in fl ower ......................................... 19.
15. Stems 70–120 cm tall, 5–12 mm thick, often lignescent
at base. Stem leaves more or less coriaceous, shiny, closely
spaced, elliptic or ovate-elliptic, rounded or truncate at
base, 5–15 cm long and 1–3.5 cm wide, 4–8 times as long
as wide. Wetland and coastal plants ................ 61. E.lucida.
+ Stems 15–80 cm tall, usually not lignescent at base. Stem
leaves not shiny, more sparse, elliptic or oblong-elliptic,
usually rounded or cordate at base, 3.5–5.5(6.5) cm
long and 1–1.8 cm wide, 2.5–5(7) times as long as wide.
Steppe, meadow, forest or rock plants ............................... 16.
16. Stem leaves cordate at base, sometimes almost amplexi-
caul ......................................................................... 60. E.agraria.
+ Stem leaves rounded at base ................................................ 17.
17. Stem leaves greyish, raylet leaves greyish or yellow-grey-
ish, more or less concolorous with stem leaves when plant
are in fl ower .............................................. 63. E.pseudagraria.
+ Stem leaves green, raylet leaves yellow or greenish-yellow,
diff erent in colour from stem leaves when plant are in
fl ower ......................................................................................... 18.
18. Plants glabrous .................................................... 58. E.iberica.
+ Plants at least partly pilose ....................... 59. E.dubovikiae.
19 (14). Plants with numerous comparatively long axillary
vegetative sprouts, often exceeding length of stem and
apical rays ................................................................................. 20.
D. V. Geltman
Новости систематики высших растений | Том 51 | 2020
46

+ Plants without axillary vegetative sprouts ...................... 21.
20. Stems ascendent, stem leaves oblong-ovate or oblong-el-
liptic, 0.7–0.8 cm long and 0.3–0.7 cm wide ..........................
............................................................................. 70. E.buschiana.
+ Stems usually erect, stem leaves linear or oblong-linear,
1.2–2.5 cm long and 0.1–0.5 cm wide ......................................
....................................................................... 65. E.daghestanica.
21 (19). Stems up to 10 cm tall. Stem leaves oblong-elliptic,
undulate at margin, rounded at apex .......... 69. E.undulata.
+ Stems 8–25 cm tall. Stem leaves obovate or elliptic, not
undulate at margin, often emarginated at apex .....................
............................................................................. 64. E.sareptana.
22 (8). Flowering stem annual, without a pseudorosette of
leaves ......................................................................................... 23.
+ Flowering stem biennial, fi rst year leaves in a pseudoro-
sette or at least with very short internodes ..................... 24.
23. Stem leaves elliptic or obovate-elliptic, attenuate or cu-
neate at base, with petioles 4–10 mm long. Mostly forest
plants .............................................................. 45. E.macroceras.
+ Stem leaves oblong or oblong-ovate, base truncate, round-
ed or cordate, sessile or with petiole up to 3 mm long.
Mostly plants of subalpine meadows ..... 46. E.oblongifolia.
24 (22). Plants glabrous, leaves of pseudorosette caducous,
leaving noticeable leaf scars ....................... 47. E.glaberrima.
+ Plants at least partly pubescent, leaves of pseudorosette
persistent, wintering .............................. 48. E.amygdaloides.
25 (7). Cyathial glands oblong-elliptic, always without ap-
pendages. Capsules mostly verrucose or tuberculate,
sometimes cristate. Leaves usually serrulate or dentate in
upper part, venation pinnate, prominent ......................... 26.
+ Cyathial glands trapezoid or reniform, sometimes with
appendages. Capsules smooth. Leaves entire, venation ob-
scurely palmate ....................................................................... 42.
26. Shrub with lignescent branches ...... 12. E.hierosolimitana.
+ Perennials ................................................................................. 27.
27. Cyathia solitary or few ................................ 20. E.ardonensis.
+ Cyathia in apical or axillary rays ........................................ 28.
28. Raylet leaves 2 ........................................................................ 29.
+ Raylet leaves 3–4 (sometimes 2 on axillary rays) .......... 37.
29. Rootstock tuberous. Stem leaves cordate or amplexicaul at
base ............................................................... 8. E.condylocarpa.
+ Rootstock not tuberous. Stem leaves rounded or cuneate
at base ........................................................................................ 30.
30. Stems and leaves glabrous .................................................... 31.
+ Stems and leaves pubescent, at least partly ..................... 32.
31. Plants 10–30 cm tall . Stem leaves usually oblong, 1.4–3.7
cm long. Stigma sessile, style absent. Plants of Lesser Cau-
casus ................................................................. 13. E.wittmannii.
+ Plants 40–60 cm tall. Stem leaves elliptic, (3.5)4–6 cm
long. Style ca. 1.5 mm. Plants of Western Transcaucasia ....
............................................................................... 25. E.eugeniae.
32 (30). Plants densely covered with hairs ca. 1 mm long, oc-
curring on littoral and coastal habitats ............ 7. E.hirsuta.
+ Plants covered by hairs less than ca. 0.5 mm long, occur-
ring in forests, montane meadows and steppes ............... 33.
33. Rays up to 1 cm long, always non-branched, usually equal
to ray leaves ............................................................................. 34.
+ Rays (2)3–10(15) cm long, branched or non-branched,
usually longer then ray leaves ............................................. 35.
34. Capsules with more or less long warts (ca. 1.5 mm high),
seeds 2.2–2.6 × 1.8–2 mm ................................. 19.E.scripta.
+ Capsules with very short, punctate warts (0.1–0.2 mm
high), seeds (2.3)2.8–3 × (2)2.3–2.5 mm ................................
............................................................................ 18.E.djimilensis.
35 (33). Capsules ca. 4 mm diam., seeds 2.2–2.8 mm long .........
............................................................................. 16. E.squamosa.
+ Capsules 7–10 mm diam., seeds 3–4 mm long ................ 36.
36. Capsules subglobose, stem leaves at base cuneate. Plants
of forests ....................................................... 17. E.czerepanovii.
+ Capsules trilobate, stem leaves at base truncate or round-
ed, seldom subcuneate. Plants of open habitats .....................
.......................................................................... 15. E.macrocarpa.
37 (28). Stems and leaves pubescent, at least partly ............ 38.
+ Stems and leaves always glabrous ...................................... 39.
38. Capsules at least partly cristate, (3.5)4–5.5 mm long,
seeds (2.5)2.9–3.2 mm long. Stems glabrous or pubescent.
Stem leaves (6)6.5–8.5 cm long, (3.5)4–7 times as long as
wide. Plants occurring throughout the Caucasus .................
.................................................................................. 23. E.procera.
+ Capsules verrucose, usually not cristate, 3.5–4 mm long.
Seeds 2.5–2.8 mm long. Stems always glabrous. Stem
leaves 4–6(7) cm long, 2–4(4.5) times as long as wide.
Plants of North-West Transcaucasia .......... 24. E.tauricola.
39 (37). Plants always without axillary vegetative sprouts ......
............................................................................... 14. E.orientalis.
+ Plants with axillary vegetative sprouts ............................ 40.
40. Capsules without warts. Stem leaves usually shortly
aristate at apex ................................................... 22. E.aristata.
+ Capsules warty. Stem leaves usually rounded at apex, at
least not aristate ..................................................................... 41.
41. Wetland plants (60)80–170(200) cm tall. Capsules 3.5–5
× 5–6 mm, with warts 0.15–0.25 mm high, seeds 3–4 mm
long ...................................................................... 21. E.palustris.
+ High mountain plants 40–60 cm tall. Capsules 2.5–3 ×
3–4 mm, with warts 0.5–1 mm high, seeds 2.5–2.6 mm
long ..................................................................... 25. E.eugeniae.
42 (25). Glands without appendages ....................................... 43.
+ Glands with appendages (at least very short) ................ 46.
43. Cyathia more or less broadly campanulate, their length
equal to width in upper part or slightly shorter, usually
with triangular, often bifi d lobes. Stems thin, 1.5–2 mm
diam., virgate, leaves glaucous, usually linear and oblong-
linear, sometimes linear-elliptic, (3)4–14 times as long as
wide ................................................................ 33. E.seguieriana.
+ Cyathia narrowly campanulate, 1.4–2 times as long as wide
in upper part, with obtuse, irregular, usually entire lobes.
Stems 1–5 mm diam., leaves yellowish-green, usually ellip-
tic, broad elliptic, obovate, 2–6 times as long as wide .. 44.
44. Stems 10–20 cm tall, 1.5–2(2.2) mm diam., apical
pseudoumbel of 3–6 rays ................................ 37. E.glareosa.
+ Stems (20)25–70 cm tall, (2.3)2.5–4 mm diam., apical
pseudoumbel of (5)7–16 rays .............................................. 45.
45. Ovary and capsule usually pilose. Stems and leaves at least
partly with minutely papillose (“glareose”) indumentum
(usually visible on magnifi cation) ............ 35. E.pannonica.
+ Ovary and capsule glabrous. Stems and leaves without
minutely palillose (“glareose”) indumentum, only leaf
margins sometimes glareose-dentate ........... 36. E.stepposa.
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46 (42). Capsule deeply trisulcate, wider than long. Seeds
almost rounded in cross-section, with a small complanate
caruncle. Plants of sandy sea shores ............ 52. E.paralias.
+ Capsule weakly sulcate, longer than wide. Seeds quadran-
gular or subquadrangular (two-faceted from ventral side
and almost rounded from dorsal side) in cross-section,
with a noticeable conical or cylindrical caruncle ........... 47.
47. Capsule conical; male fl owers with bracteoles; seeds
subquadrangular in cross-section ....................................... 48.
+ Capsule resembling a truncated tetrahedron; male fl owers
without bracteoles; seeds quadrangular in cross-section ....
...................................................................................................... 52.
48. Ovary and capsule pilose. Seeds smooth. Plants 20–70 cm
tall .................................................................. 34. E.macroclada.
+ Ovary and capsule glabrous. Seeds foveolate or smooth.
Plant small, 10–15 cm tall ................................................... 49.
49. Seeds usually 2.5–2.6 mm long. Stem leaves (1.5)2–3 cm
long ..................................................................... 38. E.smirnovii.
+ Seeds usually 2.2–2.4 mm long. Stem leaves 0.5–2 cm long
...................................................................................................... 50.
50. Apical rays 10–80 mm, usually longer than ray leaves, if
shorter, stem in upper part sparsely leaved. Seeds always
foveolate .......................................................... 39. E.petrophila.
+ Apical rays 0.5–5(7) mm, usually equal or shorter than ray
leaves, stem in upper part densely leaved. Seeds foveolate
or smooth .................................................................................. 51.
51. Perennial herb (sometimes with lignescent stem base),
annual shoots dying after fl owering ........ 40. E.erythrodon.
+ Small semishrublet, annual shoots lignescent and
continuing growth after fl owering ............. 41.E.panjutinii.
52 (47). Stem leaves linear, rarely linear-obovate, long acute
at apex, (2.5)3–5 times as long as wide ........... 27. E.rigida.
+ Stem leaves oblong-elliptic, obovate, sometimes almost
rounded, shortly acute at apex, (1.2)1.5–2.5 times as long
as wide ....................................................................................... 53.
53. Seeds always rugulose, caruncle truncate-conical or
almost spherical, obtuse ............................... 30. E.myrsinites.
+ Seeds rugulose or smooth, caruncle acute-conical ......... 54.
54. Apical pseuboumbel of 2–6(7) rays ............... 28. E.armena.
+ Apical pseudoumbel of (7)8–20 rays .......................................
...................................................................... 29. E.marschalliana.
55 (6). Ovary and capsule with warts ...................................... 56.
+ Ovary and capsule smooth, sometimes with minute
scattered warts (E.microsphaera) ...................................... 58.
56. Plants up to 14 cm tall, glabrous. Stem usually equal or
shorter than apical and axillary rays. Seeds ecarunculate ..
............................................................................3. E.coniosperma.
+ Plants 10–40 cm tall, at least partly pubescent. Stem
usually longer than apical and axillary rays. Seeds
carunculate .............................................................................. 57.
57. Capsule prominently trisulcate, 1.6–2 × 2–2.2 mm, with
warts located mainly in median part of the lobes. Seeds
1.3–1.7 × 1–1.2 mm. Apical pseudoumbel of 3, rarely 5
rays ............................................................................. 9. E.stricta.
+ Capsule obscurely trisulcate or subspherical, 2.3–3 ×
3–3.5 mm with warts located more or less sparsely. Seeds
1.9–2.2 × 1.6–1.9 mm. Apical pseudoumbel of 5, rarely 3
rays ................................................................ 10. E.platyphyllos.
58 (55). Seeds smooth .................................................................. 59.
+ Seeds ornamented .................................................................. 62.
59. Caruncle large, equal or longer than seed itself .....................
........................................................................... 42. E.grossheimii.
+ Caruncle much smaller than seed itself or absent .......... 60.
60. Glands with long horn-like appendages .... 50. E.terracina.
+ Glands without appendages ................................................ 61.
61. Plants moderately pilose. Capsule spherical, seeds
carunculate ............................................... 11. E.microsphaera.
+ Plants densely pilose. Capsule clearly trilobate, seeds
ecarunculate ...................................................... 4. E.eriophora.
62 (58). Plants at least partly pilose. Stem leaves, ray and
raylet leaves serrate near apex ............................................ 63.
+ Plants usually glabrous. Stem leaves, ray and raylet leaves
entire .......................................................................................... 64.
63. Seeds foveolate ............................................... 6. E.helioscopia.
+ Seeds striate-rugulose ......................... 5. E.rhabdotosperma.
64 (62). Raylet leaves linear, oblong-linear, narrowly elliptic
or ovate, at least 3 times longer than wide ....................... 65.
+ Raylet leaves ovate, rhombic-ovate, elliptic or almost
rounded, 1.5–2(2.5) times longer than wide ................... 68.
65. Seeds hexagonal in cross-section, longitudinally narrowly
sulcate and transversely plicate-rugulose on the facets .......
.................................................................. 49. E.aserbajdzhanica.
+ Seeds quadrangular or subquadrangular (two-faceted from
ventral side and almost rounded from dorsal side) in cross-
section, foveolate, minutely tuberculate or pustulate-
rugulose ..................................................................................... 66.
66. Stem leaves narrowly oblong-obovate, early deciduous.
Seeds tetrahedral with more or less equal facets,
transversely pustulate-rugulose .................... 44. E.szovitsii.
+ Stem leaves linear, persistent. Seeds quadrangular
with irregular facets or subquadrangular, foveolate or
tuberculate ............................................................................... 67.
67. Stem leaves, ray and raylet leaves linear, not enlarged at
base. Seeds foveolate, subquadrangular in cross-section .....
............................................................................. 54. E.ledebourii.
+ Stem leaves linear, ray and raylet leaves narrowly
triangular, slightly enlarged at base. Seeds tuberculate,
quadrangular in cross-section, with unequal facets: those
adjacent to raphe concave, other slightly convex .................
................................................................................... 51. E.exigua.
68 (64). Seeds hexangular or quadrangular in cross-section,
with prominent edges and fl at facets ................................. 69.
+ Seeds subquadrangular in cross-section, only the facets
adjacent to raphe fl at ............................................................. 73.
69. Seeds hexangular in cross-section ...................................... 70.
+ Seeds quadrangular in cross-section ................................. 71.
70. Capsule alate. Seeds 1.2–1.5 mm long, longitudinally
sulcate on facets adjacent to raphe, foveolate on other
ones .......................................................................... 55. E.peplus.
+ Capsule not alate. Seeds 1.6–1.7 mm long, on all facets
longitudinally sulcate ............................ 57. E.aulacosperma.
71 (69). Stem leaves almost rounded. Glands minute,
narrowly elliptic, without appendages. Seeds ca. 2.5 mm
long. Endemic to Talysh .................................... 2. E.hyrcana.
+ Stem leaves elliptic or spathulate. Glands more or less
prominent, semilunate or trapezoid, with or without ap-
pendages. Seeds 1–2 mm long. More widely distributed
plants ......................................................................................... 72.
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72. Capsule conical. Seeds dorsiventrally compressed, shal-
lowly quadrangular in cross-section, transversely sulcate
on facets ................................................................ 31. E.falcata.
+ Capsule subovoid. Seeds regularly quadrangular in cross-
section, transversely pustulate-rugulose on facets ...............
................................................................................. 44. E.szovitsii.
73 (68). Seeds foveolate or minutely tuberculate ................. 74.
+ Seeds tuberculate-rugulose or longitudinally rugulose .......
...................................................................................................... 75.
74. Stem leaves fi liform, numerous, imbricate, 0.5–1.5 mm
wide, usually early cauducous. Seeds tuberculate, 1.3–1.7
mm long .............................................................. 26. E.aleppica.
+ Stem leaves obovate or oblong, not imbricate, more or less
persistent, 2–3 mm wide. Seeds foveolate, 1.6–2 mm long
........................................................................... 53. E.taurinensis.
75. Raylet leaves ovate or ovate-elliptic. Seeds irregularly
rugulose or tuberculate-rugulose, 1.6–1.7 mm long,
caruncle prominent, more or less persistent. Plants of
Transcaucasia and Talysh .................................. 56. E.arvalis.
+ Raylet leaves elliptic or oblong. Seeds longitudinally
sulcate-rugulose, 1.3–1.4 mm long, caruncle minute, early
caudicous. Endemic to Stavropol Heights ..............................
............................................................................. 32. E.normannii.
76 (3). Stems erect or ascendent, leaves 1–4 cm long, with
hairs 0.7–1.5 mm long ........................................ 71. E.nutans.
+ Stems prostrate, leaves 0.3–1 cm long, with hairs 0.2–0.7
mm long or glabrous .............................................................. 77.
77. Capsules appressedly pilose ................................................. 78.
+ Capsules glabrous or squarrosely pilose ........................... 79.
78. Stems in upper part usually squarrosely pilose. Seeds 0.8–
0.9 mm long ...................................................... 77. E.maculata.
+ Stems in upper part usually crispate pilose. Seeds 1–1.2
mm long ............................................................ 78. E.forskaolii.
79. Leaves entire or subentire .................................................... 80.
+ Leaves dentate or crenulate ................................................. 81.
80. Seeds ovate-conical, smooth, ripe ones sometimes papil-
late, 2–3.5 mm long. Plants of sea shores ........ 72. E.peplis.
+ Seeds narrowly ovoid-cylindric, quadrangular, irregularly
foveolate-rugulose, 1–1.4 mm long. Plants of steppes and
semideserts ...................................................... 76. E.granulata.
81. Seeds ovoid-quadrangular, smooth, sometimes papillate
when ripe .......................................................... 73. E.humifusa.
+ Seeds transversely rugulose or cristate ............................. 82.
82. Seeds irregularly transversely rugulose on facets. Style
0.3–0.5 mm long, stigma not sessile ....... 74. E.chamaesyce.
+ Seeds with 3–5 transverse almost regular ribs on facets.
Style 0.1–0.2 mm long, stigma subsessile ................................
....................................................................... 75. E.glyptosperma.
Taxonomic treatment
Euphorbia L.
Subgen. 1. Esula Pers.
Sect. 1. Lathyris Dumort.
1. E. lathyris L. 1753, Sp. Pl.: 457. ≡ Tithymalus
lathyris (L.) Hill, 1768, Hort. Kew.: 172.3.— Lecto-
type (Geltman, 2015b: 129): “Herb. Linn. № 630.32”
(LINN!).
Roadsides, sea shores, human settlements; alien.
WTC: 7b, 7c, 7d, 7e; ETC: 9d.— TCS.
The origin of E. lathyris is not yet clear. It is now
found in both the Mediterranean and neighboring areas
as well as in Eastern Asia. Most of the available herba-
rium specimens from the Caucasus were collected in the
19th or early 20th centuries. Pasta and Troìa (2019) stat-
ed that this species was much more frequent and com-
monly used as a medicinal plant in southern Italy and
Sicily until 2–3 centuries ago, as well as in the classical
ages.
Euphorbia lathyris was included in the “Cultivated
fl ora of the USSR” as an oil-producing plant (Sharapov,
1941). Sinskaya (1969: 246) believed that E.lathyris oc-
cured wild in the mountains of Western China and was
cultivated in Eastern Asia as an oil plant; she also men-
tioned that its culture in Japan has now entirely ceased.
Sect. 2. Lagascae Lázaro
2. E. hyrcana Grossh. 1920, Trudy Tifl issk. Bot.
Sada, 2 (1): 7. — Lectotype (Geltman, 1991a:
112): “Prov. Baku, dist. Lenkoran (Talysh), Lerik, in
faucibus fl . Jataganovtshaj, in fruticetis, 21 IV 1915,
A.Grossheim” (TBI!).
Stony montane slopes, bush thickets.
T.— TCS.— Irano-Turanian element.
Known only from the vicinity of the village Le-
rik near Lenkoran by a few gatherings. Morphologi-
cally is very similar to E.phymatosperma Boiss. et Gaill.
Prokhanov (1949) treated this species as a synomym of
E. peplus, but his opinion was based on misinterpreta-
tion of the original material of E.hyrcana— see details
in Geltman (1991a). Unfortunately, such misinterpre-
tation was recently repeated by Pahlevani et al. (2020)
that led to wrong sectional attribution.
Sect. 3. Helioscopia Dumort.
3. E. coniosperma Boiss. et Buhse, 1860, Nouv.
Mém. Soc. Nat. Moscou, 12: 196. — Lectotype
(Rechinger, Schiman-Czeika, 1964: 26): “Persia, 1847,
Buhse” (G!; isolectotype— LE01070910!).
= E.ancyrensis Azn. ex M. S. Khan, 1964, Notes Roy.
Bot. Gard. Edinburgh, 25: 106. ≡ Tithymalus ancyrensis
(Azn. ex M. S. Khan) Soják, 1972, Čas. Nár. Mus., Odd.
Prír. 140: 170.— Holotype: “Pl. de Turquie, coll.
Frères, in Hb Aznavour” (G!; isotype— E00362396!).
Dry steppes on clayey soils.
ETC: 9c; SWTC: 10a; STC: 11a, 11c, 11d. —
TCS.— Irano-Turanian element.
The lectotype of E. coniosperma has a standard
printed label “Persia, Buhse”, although in fact the plant
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was collected in present-day Armenia. The isolectotype
(LE) has two labels: the original “Gamarlu” and the
later “In der Araxesebene bei Erevan u. Gamarlu, April
1847, Buhse” (similar to the text from the protologue).
There is little doubt that the G and LE sheets are parts
of a single gathering, because there is a record in the
protologue that the species was known by a single spe-
cimen (Boissier, Buhse, 1860: 196).
4. E.eriophora Boiss. 1844, Diagn. Pl. Orient., sér.
1, 5: 51. ≡ Tithymalus eriophorus (Boiss.) Klotzsch et
Garcke, 1860, Abh. Königl. Akad. Wiss. Berlin, 1859:
65.— Type unknown.
= E. lasiocarpa K. Koch, 1849, Linnaea, 21: 721,
nom. illeg., non Klotzsch, 1843. — T ype: “in herb.
Gundelsheimer” (Koch, 1849: 721).
Clayey steppes, fallow lands and cultivated grounds.
STC: 11a, 11c, 11d.— TCS.— Irano-Turanian ele-
ment.
Regarding the record for NWTC: 6a (Zernov, 2000,
2006)— see comment under E.hirsuta.
The holotype of E.eriphora is likely missing (Khan,
1964).
5. E. rhabdotosperma Radcl.-Sm. 1975, Notes
Roy. Bot. Gard. Edinb. 34, 1: 129. ≡Tithymalus
rhabdotospermus (Radcl.-Sm.) Holub, 1977, Folia
Geobot. Phytotax. 12: 428.— Holotype: “Elmali-
Korkuteli, 8 km from Elmali, in dry steppe, 1120 m,
31 III 1962, Dudley (Davis 35223)” (E 00359917!;
isotype— K 001080014!).
Stream banks, especially on gravel substrate, pas-
tutres and agricultural lands, roadsides.
ECC: 2c; CC: 4c; EC: 5a (vicinity of Grozny), 5b
(north), 5c, 5d; CTC: 8a; ETC: 9b, 9c, 9d, 9e; SWTC:
10a, 10b; STC: 11a, 11b, 11d, 11f, 11g; T. — NSC,
TCS.— Submediterranean element.
6. E. helioscopia L. 1753, Sp. Pl.: 459. ≡ Tithyma-
lus helioscopius (L.) Hill, 1768, Hort. Kew.: 172.3. —
Lectotype (Jafri, El-Gadi, 1982: 33): “Herb. Linn.
№630.49” (LINN!).
Cultivated grounds, roadsides, stream banks and
beds, forest margins.
WCC: 1a; WC: 3c; EC: 5c, 5d; NWTC: 6a; WTC:
7a, 7b, 7c, 7e; CTC: 8a, 8b; ETC; STC: 11d, 11e, 11f,
11g; T.— NCS, TCS.— European-Pantethyan element.
Two subspecies are accepted here. Their distribution
in the Caucasus is almost identical, although in more
northern areas only the type subspecies is found. Subsp.
helioscopioides is restricted to more natural habitats and
usually has an autumn— spring life cycle. Both subspe-
cies are evidently undercollected in the area.
Key to the subspecies
1. Stems solitary, erect, rarely ascendent .. subsp. helioscopia.
+ Stems numerous, ascendent at the base ...................................
................................................................... subsp. helioscopioides.
6a. E.helioscopia L. subsp. helioscopia.
Cultivated grounds, roadsides, sometimes by stream
banks.
WCC: 1a; WC: 3c; EC: 5c, 5d; NWTC: 6a; WTC:
7a, 7b, 7c, 7e; CTC: 8a, 8b; ETC; STC: 11d, 11e, 11f,
11g; T.— NCS, TCS.
6b. E. helioscopia L. subsp. helioscopioides (Los-
cos et J. Pardo) Nyman, 1881, Consp. Fl. Europ. 3:
651. ≡ E. helioscopioides Loscos et J. Pardo, 1863, in
H. M. Willkomm (ed.), Ser. Inconf. Pl. Aragon.: 93.
≡Tithymalus helioscopius (L.) Hill subsp. helioscopioides
(Loscos et J. Pardo) Soják, 1972, Čas. Nár. Mus., Odd.
Prír. 140: 173. ≡ T. helioscopioides (Loscos et J. Par-
do) Holub, 1977, Folia Geobot. Phytotax. 12: 428. —
Syntype (?): “Castelseras in Aragonia, ex herb. Los-
cos” (MPU 014450— image!).
= E. helioscopia L. subsp. hiemalis A. P. Khokhr.
1989, Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol.
94, 6: 96. — Holotype: “Аджарская АССР,
Хелвачаурский р-н, Эрге, осыпь под скалами, 1 III
1988, А. П. Хохряков [Adjara ASSR, Khelvachauri
district, Erge, scree under the rocks, 1 III 1988,
A.P.Khokhryakov]” (MW 0593507!).
Stream banks and beds, forest margins, less often as
a weed in cultivated grounds.
WCC: 1a; EC: 5c, 5d; WTC: 7a, 7b, 7c, 7e; CTC: 8a;
ETC: 9b, 9c, 9d, 9e; STC: 11e, 11f, 11g; T.— NCS, TCS.
7. E.hirsuta L. 1759, Amoen. Acad. 4: 483. — Type
not designated.
= E. pubescens Vahl, 1791, Symb. Bot. 2: 55.
≡Tithymalus pubescens (Vahl) Samp. 1931, Anais
Fac. Sci. Porto, 17: 46.— Lectotype (Radcliff e-
Smith, 1982): “circa Zowan, herb. Vahl [2201/28]”
(С10000684— image!).
— E.verrucosa auct. non L.: M. S. Khan, 1964, Notes
Royal Bot. Gard. Edinb. 25 (2): 103.
Gravel beaches and wetlands near seashore.
?NWTC: 6a; WTC: 7b, 7c, 7e. — ?NCS, TCS. —
Macaronesian-Mediterranean element.
Zernov (2000, 2006) collected in NWTC: 6a
(Краснодарский край, Анапа, песчаный пляж, 22
VIII 1998, А.С.Зернов, № 311 [Krasnodar Territory,
Anapa, sand beach, 22 VIII 1998, Zernov 311] (MW
0690215!)) a small undeveloped densely pubescent Eu-
phorbia plant which he identifi ed as E. eriophora. How-
ever this specimen could belong to E.hirsuta as well and
its habitat is more typical for the latter than for the for-
D. V. Geltman
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mer. Until now there is no reliable records of the occur-
rence of normally developed E. hirsuta in this area, al-
though it is quite probable because this species recently
was found in Crimea (Geltman, Shatko, 2012).
8. E. condylocarpa M. Bieb. 1808, Fl. Taur.-Cauc.
1: 377. ≡ Tithymalus condylocarpus (M.Bieb.) Klotzsch
et Garcke, 1860, Abh. Königl. Akad. Wiss. Berlin 1859:
78. — Holotype: “Ex Caucaso cabardinico, circa
Narzana lecta, 1800 [Bieberstein]” (LE 01070909!).
= E.amplexicaulis Ledeb. 1850, Fl. Ross. 3 (2): 567,
nom. illeg., non Hook. f. 1847. ≡Tithymalus amplexicau-
lis Klotzsch et Garcke, 1860, Abh. Königl. Akad. Wiss.
Berlin 1859: 80. — Holotype: “Schuscha, herb.
Ledebour, 830.35” (LE 01057260!).
= E.cardiophylla Boiss. et Heldr. 1853, in Boiss., Di-
agn. Pl. Orient., sér. 1, 12: 107. ≡ Tithymalus cardiophyl-
lus (Boiss. et Heldr.) Klotzsch et Garcke, 1860, Abh.
Königl. Akad. Wiss. Berlin 1859: 78.— Holotype:
“Mt. Solima reg. infer. en montant a Kartsibahir, in fru-
ticeti, 1 V 1845 [Heldreich], № 1058” (G-BOISS!).
Steppes and meadows, sometimes shrub thickets or
sparse forests, usually on clacareous soils or rocks; 300–
2400 m.
WCC: 1b; WC: 3a, 3b, 3c; CC; EC; NWTC: 6a;
WTC: 7c, 7d, 7e; CTC; ETC; SWTC: 10b, 10c; STC;
T.— NCS, TCS.— Submediterranean element.
9. E. stricta L. 1759, Syst. Nat., ed. 10, 2: 1049.
≡ Tithymalus strictus (L.) Klotzsch et Garcke, 1858, Fl.
N. Mitt.-Deutschland, ed. 4: 290. ≡ Euphorbia platy-
phyllos L. var. stricta (L.) Fiori, 1901, Fl. Italia, 2:
281. — Lectotype (Radcliff e-Smith, 1982: 593):
“Herb. Linn. № 630.54” (LINN!).
= E.serrulata Thuill. 1799, Fl. Env. Paris, ed. 2: 237.
≡ E. platyphyllos L. var. serrulata (Thuill.) Pers. 1806,
Syn. Pl. 2: 18. ≡ Tithymalus serrulatus (Thuill.) Holub,
1970, Preslia, 42: 94.— Type unknown.
= E. micrantha Stephan ex Willd. 1799, Sp. Pl. 2:
905. ≡ Tithymalus micranthus (Stephan ex Willd.) Raf.
1838, Fl. Tellur. 4: 115. ≡ Euphorbia stricta L. subsp. mi-
crantha (Stephan ex Willd.) Nyman, 1881, Consp. Fl.
Eur.: 651. — Lectotype (Geltman, 2015b: 130):
“Habitat in Persia, Stephan” (B-W 09318!).
= E. densifolia K. Koch, 1849, Linnaea, 21: 722. —
Lectotype (Geltman, 2011: 120): “Caucasus, Wil-
helms” (LE 01070914!).
Meadows, forest glades and margins, pastures, sand
and gravel river banks and sea shores, roadsides, culti-
vated grounds; very common; 0–2100 m.
WCC; ECC; WC; CC; EC; NWTC; WTC; CTC;
ETC; SWTC: 10a, 10b; STC: 11e, 11f; 11g; T.— NCS,
TCS.— European-Pantethyan element.
10. E.platyphyllos L. 1753, Sp. Pl.: 460. ≡ Tithyma-
lus platyphyllos (L.) Hill, 1768, Hort. Kew.: 172.4. —
Lectotype (Geltman, designated here): “Herb.
Linn. №630.61” (LINN!).
Naturalized in botanical gardens.
?CC: 4a (Zheleznovodsk); STC: 11a (Yerevan bo-
tanical garden). — ?NCS, TCS. — Euro-Siberian ele-
ment.
The only reliable record of this species for the Cau-
casus is one from the Yerevan Botanical Garden where
it is evidently alien and naturalized. There is also a
specimen: “Железноводск [Zheleznovodsk], 17 V 1887,
Herb. Akinfi ewi” (TBI) which could be referred to this
species. Most probably this specimen was wrongly la-
beled, especialy taking into account that another spe-
cies with dubious presence in the Caucasus (E.agraria)
also is known only from Zheleznovodsk according to
Akinfi evʼs gatherings. Сhintibidze and Gvinianidze
(1983: 160) mention E. platyphyllos for WTC: 7b but
the voucher herbarium specimens (SUKH) in my opin-
ion belong to E.stricta. At the same time it is impossible
to exclude that E.platyphyllos could be found as native
in the Caucasus since it also occurs in Crimea.
11. E. microsphaera Boiss. 1846, Diagn. Pl. Ori-
ent., sér. 1, 7: 87. ≡ Tithymalus microsphaerus (Boiss.)
Klotzsch et Garcke, 1860, Abh. Königl. Akad. Wiss.
Berlin 1859: 74. — Lectotype (I: Rechinger,
Schiman-Czeika, 1964: 26; II: Geltman, designated
here): “In humidis ad rad. m. Sabst-Buschom prope
Schiraz, 31 V 1842, Kotschy, № 448” (W 0031040!;
isolectotypes: A 00277273 — image!, BM 000951541!,
CAS 00123963— image!, FI 011590 — image!, G-DC
00311672!, G 00441434!, G-BOISS!, GOET 003734—
image!, JE 00002899, 00002900— image!, K001080030!,
L 0150814 — image!, LE 01071213!, MO-260133–
260135!, P 00702735!, 00702737–00702740— images!,
WAG0004311 — image!, W 0031041, 0031042, 1889-
0154945, 1889-0159509!, WU 0069098!).
= E. subtuberculata C. A. Mey. ex Boiss. 1862, in
DC., Prodr. 15, 2: 118. — Lectotype (Geltman,
2015b: 131): “Rarior ad vias circa Khoi prov. Aderbaid-
zhan, 15 VI 1828, Szovits” (LE 01071317!).
Coastal cliff s.
T.— TCS.— Submediterranean element.
Known for the area from a single specimen:
“Ленкорань, близ сел. Н. Нюады, береговой обрыв,
11 VI 1931, Н. Шипчинский, № 147 [Lenkoran, near
Nizhniye Nyuady, 11 VI 1931, N. Schipczinsky, № 147]”
(LE 01071326!).
Rechinger and Schiman-Czeika (1964) unintention-
ally designated a specimen kept in W as the lectotype
of E.microsphaera. However, there are several suitable
A synopsis of Euphorbia (Euphobiaceae) for the Caucasus
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51

specimens in W, so the next step of lectotype selection
is done here. The selection of the lectotype of this name
done by me (Geltman, 2012: 501) now appears to be su-
perfl uous.
12. E.hierosolymitana Boiss. 1853, Diagn. Pl. Ori-
ent., sér. 1, 12: 110. ≡ E.thamnoides Boiss. var. hierosoly-
mitana (Boiss.) Boiss. 1862, in DC., Prodr. 15, 2: 131.—
Lectotype: (Geltman, 2008: 150): “Palaestina,
Eirusalem, IV–V 1846, E. Boissier” (G-BOISS!; isolec-
totypes: G-DC 00312179!, G 00441419!).
= E. dumosa Boiss. 1853, Diagn. Pl. Orient., sér. 1,
12: 110, nom. illeg., non A.Rich. 1850. ≡ E.thamnoides
Boiss. 1860, Cent. Euphorb.: 33. ≡ Tithymalus tham-
noides (Boiss.) Soják, 1972, Čas. Nár. Mus., Odd. Prír.
140: 177. — Lectotype (Geltman, 2006b: 163):
“Syria, Tripoli, V–VII 1846, E. Boissier” (G-BOISS!,
isolectotypes: G-DC!, K 001080021!, LE 01071250!,
01071251!).
Rocks.
WTC: 7e (Chorokh valley, Erge).— TCS.— Medi-
terranean element.
This Mediterranean species is known only from a
single specimen in the Caucusus: “Аджария, Батумский
р-н, Эрге, 15 XI 1969, А. Дмитриева [Adjara, Batumi
district, forest rocks of Chorokh gorge, 15 XI 1969, A.
Dmitrieva]” (LE 01071325!). This specimen has part
of certainly lignescent shoot, which is characteristic of
E.hierosolimitana. This locality is very disjunct from the
main part of the species distribution area.
13. E. wittmannii Boiss. 1860, Cent. Euphorb.:
3. — Lectotype (Radcliff e-Smith, 1982: 590):
“Südrussland (Atskur, Osurgeti, etc.), Wittmann” (LE
01070961!).
Stony slopes and rocks.
WTC: 7d (Ozurgeti); CTC: 8a (Borjomi, Bakuri-
ani); STC: 10a. — TCS. — Euro-Siberian (Caucasian)
element.
There are some morphological characters, espe-
cially a very short style, which this species has in com-
mon with European species of the E.verrucosa L. group
(E.fl avicoma DC., E.polygalifolia Boiss. et Reut., E.spi-
nosa L., etc.).
The lectotype selection done by me (Geltman, 2008:
122) appears to be superfl uous.
14. E.orientalis L. 1753, Sp. Pl.: 460. ≡ Tithymalus
orientalis (L.) Hill, 1768, Hort. Kew.: 172.3. — Lec-
totype (Croizat, 1938: 98): “Herb. Linn. №630.60”
(LINN!).
= E.notadenia Boiss. et Hohen. 1853, in Boiss., Di-
agn. Pl. Orient., sér. 1, 12: 111. ≡ Tithymalus nota denius
(Boiss. et Hohen.) Klotzsch et Garcke, 1860, Abh.
Königl. Akad. Wiss. Berlin 1859: 78.— Lectotype
(Geltman, designated here): “In monte Elbrus [El-
burs] pr. Derbend, 30 VI 1843, Th. Kotschy, № 420 (Pl.
Pers. bor. Ed. R. F. Hohenacker, 1846)” (G-BOISS!,
isolectotypes: FI 011589— image!, G!, LE 01071139!,
01071310!, P 00606917!).
= E. artvinensis Bornm. et Woronow, 1912, Vestn.
Tifl issk. Bot. Sada, 26: 3.— Lectotype (Geltman,
2015b: 127): “Артвинский окр., долина Аргамуг-су
близ уроч. Горгота-ханч, 12 VII 1911, Ю. Воронов,
№ 5681 [Artvin district, valley of the river Arganuch-
su near Gorgota-khanch terrain feature, 12 VII 1911,
Woronow, № 5681]” (TBI 28701!).
Stony slopes, wadies, eroded sandy and clayey plac-
es; 1300–2100 m.
SWTC: 10c; STC: 11a, 11b, 11c, 11d, 11e, 11f. —
TCS.— Irano-Turanian element.
15. E. macrocarpa Boiss. et Buhse, 1860, Nouv.
Mém. Soc. Nat. Moscou, 12: 197. ≡Tithymalus macro-
carpus (Boiss. et Buhse) Prokh. 1949, Fl. URSS, 14:
350, nom. illeg., non (Bentham) Croizat, 1937. ≡ T.no-
tabilis Soják, 1972, Čas. Nar. Mus., Odd. Prír. 140:
174.— Lectotype (Khan, 1964: 94): “Persia, Ssa-
mam [1 VI 1848], Buhse” (G-BOISS!, isolectotypes:
BR 0000005106370— image!, LE 01071134!, W 1889-
0080953, 1963-0023445!).
Clayey steppes, sparse forests on rocks.
ETC: 9b (Gobustan area); STC: 11f (between
Karchavan and Agarak).— TCS.— Irano-Turanian ele-
ment.
16. E. squamosa Willd. 1799, Sp. Pl. 2, 2: 918. ≡
Tithymalus squamosus (Willd.) Klotzsch et Garcke,
1860, Abh. Königl. Akad. Wiss. Berlin 1859: 78. —
Holotype: “Herb. Willd. № 9358” (B-W 09358-
010!).— Fig. 2: A–C.
= E. aspera M. Bieb. 1808, Fl. Taur.-Cauc. 1: 377.
≡ Tithymalus asper (M. Bieb.) Klotzsch et Garcke, 1860,
Abh. Königl. Akad. Wiss. Berlin 1859: 78. — Lec-
totype (Geltman, 2006: 162): “Ex Caucaso demis-
siore ruthenico, 1800, [Bieberstein]” (LE-Bieb.: LE
01070983!).
= E. muricata M. Bieb. 1808, Fl. Taur.-Cauc. 1:
378.— Lectotype (Geltman, 2006: 163): “Ex Ibe-
ria, 1806, Steven” (LE-Bieb.: LE 01071029!).
= E. muricata M. Bieb. var. wilhelmsiana K. Koch,
1849, Linnaea, 21: 725. ≡ E. squamosa Willd. var. wil-
helmsiana (K. Koch) Oudejans, 1992 (publ. 1993),
Collect. Bot. (Barcelona), 21: 188. — Lectotype
(Geltman, 2015b: 130): “Caucasus, Wilhelms” (LE
01070940!).
D. V. Geltman
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52

Fig. 2. Caucasian species of Euphorbia section Helioscopia.
Euphorbia squamosa: A, B— apical synfl orescences, C— habit; E.procera: D— habit, E,F— apical synfl orescences.
A synopsis of Euphorbia (Euphobiaceae) for the Caucasus
Novitates Systematicae Plantarum Vascularium | Volume 51 | 2020
53

= E.aspera M. Bieb. var. serrata Boiss. 1862, in DC.,
Prodr. 15, 2: 124.— Holotype: “Imeretien, Eich-
wald (Herb. Ledebour)” (LE 01057264!).
= E. talyschensis Boiss. et Buhse, 1860, Nouv.
Mém. Soc. Nat. Moscou, 12: 196. ≡ E.aspera M. Bieb.
var. oligadenia Boiss. 1862, in DC., Prodr. 15, 2: 124
(1862). — Lectotype (Geltman, 2015b: 131):
“Massula, IV 1848, Buhse” (LE 01070957!, isolecto-
type: G-BOISS).
= E. abchazica Woronow, 1912, Vestn. Tifl issk.
Bot. Sada, 22: 3. — Lectotype (I: Chinthibidze,
Gvinianidze, 1983: 157; II: Geltman, designated here):
“Abchazia occid., locus Mombeschta inter mm. Mam-
dzyschcha et Kutysch, pascua alpina, 6500′, 28 VII/ 10
VIII 1905, G. Woronow, № 356” (TBI 1025054!, isolec-
totype: LE01057259!).
Forests, rarely subalpine meadows and krummholz;
50–2400 m.
WCC: 1a (Stavropol Heigth); WC; CC; EC: 5a, 5b,
5c, [?5d]; NWTC; WTC; CTC: 8a, 8c [?8b]; ETC: 9a,
9c, 9d, 9e, [?9f]; SWTC: 10a; STC: 11a, 11b, 11f, 11g;
T. — NCS, TCS. — Euro-Siberian (Caucasian-Euxini-
an-Hyrcanian) element.
Typical element of broadleaved (sometimes mixed)
forests, also penetrates into the subalpine belt. Variable,
especially in the length of warts on capsules, but this
variation does not correlate with geographical distribu-
tion or habitat preferences.
17. E. czerepanovii Geltman, 1997, Bot. Zhurn.
82 (3): 122. — Holotype: “Дагестанская АССР,
Гумбетовский р-н, между сел. Буртанай и перевалом
к сел. Данух; опушка букового леса по склону к
речке, 17 VI 1961, Н. Н. Цвелёв, С. К. Черепанов,
Г. Н. Непли, А. Е. Бобров, № 1220 [Daghestan
ASSR, Gumbetovsky district, between village Burta-
nai and pass to village Danukh, margin of beech forest
on the slope to the stream, 17 VI 1961, Tzvelev et al.,
№1220]” (LE 01070911!).
Forests.
EC: 5b (Salatau range); ETC: 9a (Lagodekhi, Kho-
chaldagh mountain). — NCS, TCS. — Euro-Siberian
(Caucasian) element.
Very similar to E. squamosa, but diff ers in larger
seeds (ca. 3 mm long). The type specimen also diff ers
by its larger raylet leaves. The second known specimen
was collected on the south slope of Mt. Khochaldagh
located just at the border between Russia and Georgia
(Georgia, Kachetia, pr. pag. Lagodechi, m. Choczal-
dag, 6 VIII 1935, W. Kozlowski (TBI)). This species is
likely a relict restricted to the Eastern Caucasus. Prob-
ably, it gradually dissappears due to hybridization with
E.squamosa.
18. E. djimilensis Boiss. 1879, Fl. Or. 4: 1104.
≡ Tithymalus djimilensis (Boiss.) Soják, 1972, Čas.
Nár. Mus., Odd. Prír. 140: 171.— Lectotype (I:
Khan, 1964: 93; II: Geltman, designated here): “Envi-
rons de Djimil (Lazistan), vers 2200 metres d’altitude,
VII 1866, Balansa, № 1440” (G-BOISS; isolectotypes:
F 0056676F— image!, G 00441427!, GOET 003712—
image!, JE 00004090 — image!, K 001080026!, LE
0107128!, 01071283!, P 00606718–00606720!, US
00109333!, W 1889-0042024!, 0031053!).
Rhododendron caucasicum Pall. and Juniperus hemi-
sphaerica C. Presl subalpine krummholz; 2000–2300 m.
WTC: 7b, 7e.— TCS.— Euro-Siberian (Euxinian)
element.
Khan (1964) selected the “type” specimen from
G. However, there are more than one specimens in G,
so additional selection is necessary. In the Caucasus
E.djimilensis mostly occurs in Adjara, but one specimen
is also known from Abkhasia.
19. E.scripta Sommier et Levier, 1892, Trudy Imp.
S.-Peterburgsk. Bot. Sada, 12 (5): 159. ≡ Tithymalus
scriptus (Sommier et Levier) Prokh. 1949, Fl. URSS,
14: 348, nom. altern. — Lectotype (Geltman,
2015b: 131): “Svanetia libera, in jugo Utviri inter fl umi-
na Nakra et Nenskra, 2500 m, 19 VIII 1890, Sommier et
Levier, № 1186” (FI, sheet with handwritten species—
image!, isolectotype: FI— image!).
Grass communities in subalpine and alpine belt;
1800–2300 m.
WC: 3b (Mt. Bolshoi Bambak); 3c (vicinity of
Dombai, the Gonachkhir River); WTC: 7a, 7b, 7c. —
NCS, TCS.— Euro-Siberian (Euxinian) element.
Similar to E. djimilensis, but diff ers in the length
of processes on the capsule and in seed size, as well as
slightly diff ererent habitat preferences. See more in
Geltman (1991b).
20. E. ardonensis Galushko, 1976, Novosti Sist.
Vyssh. Rast. 13: 210. ≡ Tithymalus ardonensis (Galushko)
Galushko, 1979, Fl. Severn. Kavkaza Vopr. Ist. 3: 56.—
Lectotype (Geltman, 2002a: 122): illustration in
Galushko, 1976, Novosti Sist. Vyssh. Rast. 13: 211.
Rocks; 1200 m.
CC: 4c (valley of the Ardon River).— NCS.— Euro-
Siberian (Caucasian) element.
Although Galushko (1976) mentioned that the type
of E.ardonensis should be in LE, it was not found there
despite careful search, so the illustration was designated
as lectotype.
Very rare species known from a single locality— the
valley of the Ardon River in the area of Skalistyi Range
(Terekbaev, 1991). The apical synfl orescence is reduced
D. V. Geltman
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54

to a single cluster of few cyathia (similar to E.capitulata
Rchb.).
21. E. palustris L. 1753, Sp. Pl.: 462. — Lec-
totype (Polyatschek, 1971: 190): “Herb. Linn.
№630.69” (LINN!).
Wetlands, river banks.
WCC: 1a; NWTC: 6a; WTC: 7b, 7c. — NCS,
TCS.— Euro-Siberian element.
Galushko (1980) recorded this species for ETC, but
without precise localities. It is quite probable taking
into account habitat preferences of the species, although
there are no herbarium specimens from this area.
22. E. aristata Schmalh. 1892, Ber. Deutsch. Bot.
Ges. 10: 292. ≡ Tithymalus aristatus (Schmalh.) Prokh.
1933, Sist. Obzor Moloch. Sredn. Azii: 108. ≡ Euphorbia
soongarica Boiss. subsp. aristata (Schmalh.) Prokh. et
Kuzm. 1961, Izv. Bot. Inst. (Sofi a), 8: 148. ≡Tithymalus
soongaricus (Boiss.) Prokh.subsp. aristatus (Schmalh.)
Soják, 1972, Čas. Nár. Mus., Odd. Prír. 140: 176. —
Lectotype (Geltman, 2008: 139): “Auf steinigen
Böden in Полковничий яр, 20 VI 1878, 20 V 1879,
A.Normann [Auf steinigen Böden in Polkovnichy Yar,
20 VI 1878, 20 V 1879, A. Normann]” (KW 147557!).
Wet saline places on slopes of ravines.
WCC: 1b (Stavropol Heights).— NSC.— Euro-Si-
berian (Caucasian) element.
This species is a member of the complex of closely
related species (E. aggr. soongarica — see Geltman,
2008) which includes also Central Asian E. soonga-
rica Boiss., E. lamprocarpa (Prokh.) Prokh. and E. ve-
lenowskyi Bornm. from the Balkans. E. aristata diff ers
from the others by its long-aristate leaf apex (especially
leaves of axillary vegetative shoots), but this character
is not very stable. The complex defi nitely needs careful
revision and phylogeographic studies.
23. E.procera M. Bieb. 1808, Fl. Taur.-Cauc. 1: 378.
≡ Tithymalus procerus (M. Bieb.) Klotzsch et Garcke,
1858, Fl. N. Mitt.-Deutschland, ed. 4: 291. ≡ Euphorbia
pilosa L. var. procera (M. Bieb.) Nyman, 1881, Consp.
Fl. Eur.: 650.— Lectotype (Geltman, 2006: 163):
“Ex Tauria et Caucaso demissiora” (LE-Bieb.: LE
01071040!).— Fig. 2: D–F.
= Euphorbia caucasica Dubovik, 1977, Novosti Sist.
Vyssh. Nizsh. Rast. (Kiev), 1976: 96.— Holotype:
“Краснодарский край, Геленджикский горсовет,
Архипо-Осиповка, в сосновом лесу из Pinus pithyusa,
27 V 1975, О. Дубовик [Krasnodar Region, Gelendzhik
city area, Arkhipo-Osipovka, in pine forest of Pinus
pithyusa, 27 V 1975, Dubovik]” (KW02226!, isotype:
KW 02225!).
— E.villosa auct. non Waldst. et Kit.: Prokh. 1949,
Fl. URSS, 14: 359; Khalilov, 1955, Fl. Azerb. 6: 114;
Ter.-Chatsch. et Tamamsch. 1973, Fl. Armenii, 6: 102;
Chint. et Gvinian. 1983, Fl. Georgiae, ed. 2, 8: 158.
Forest glades and margins, meadows in forest belt;
300–2200 m.
WCC: 1a; WC: 3a, 3b, 3c; CC; EC; NWTC; CTC;
ETC: 9c, 9e, 9f; SWTC: 10b, 10c; STC: 11a, 11b. —
NCS, TCS.— Euro-Siberian (Caucasian) element.
Very similar to European E.illirica Lam. (= E.villosa
Waldst. et Kit.), but diff ers by generally larger seeds
((2.5)2.9–3.2 mm vs. 2.2–2.9 mm in E. illirica) and
fruits often with cristate plicae.
24. E. tauricola Prokh. 1949, Fl. URSS, 14: 736.
≡ Tithymalus tauricola (Prokh.) Prokh. 1949, l.c.: 736,
nom. altern. ≡ E. austriaca A. Kern. subsp. tauricola
(Prokh.) Chrtek et Křisa, 1970, Preslia, 42: 263. —
Holotype: “Крым, Бельбек, на опушке леса,
14 V 1898, К.Л. Гольде [Crimea, Belbek, forest mar-
gin, 14 V 1898, Golde]” (LE 01071096!, isotype: LE
01071101).
Forests; 300–800 m.
NWTC: 6a.— NCS.— Euro-Siberian element.
25. E. eugeniae Prokh. 1949, Fl. URSS, 14: 735. ≡
Tithymalus eugeniae (Prokh.) Prokh. 1949, l. c.: 735,
nom. altern. — Holotype: “Красная Поляна,
субальпийский луг при подъеме на г. Ачишхо, 10
VIII 1946, Е. Победимова [Krasnaya Polyana, subal-
pine meadow on the slope of Achishkho mountain, 10
VIII 1946, E.Pobedimova]” (LE 01070915!).— Fig. 3.
Subalpine and alpine meadows, gravel screes on acid
substrate, can be dominant in the plant cover; 1900–
2100 m, sometimes at lower elevation near streams.
WC: 3b (upper reaches of the Bolshaya Laba River
basin); WTC: 7a, 7b. — NCS, TCS. — Euro-Siberian
(Euxinian) element.
Subendemic to Abkhasia.
Sect. 4. Myrsiniteae (Boiss.) Lojac.
26. E. aleppica L. 1753, Sp. Pl.: 458. ≡ Tithymalus
aleppicus (L.) Klotzsch et Garcke, 1860, Abh. Königl.
Akad. Wiss. Berlin 1859: 84. — Lectotype (Tur-
land, 1995: 136): “Herb. Linn. № 630.46” (LINN!).
= E. condensata Fisch. ex M. Bieb. 1819, Fl. Taur.-
Cauc. 3: 322. ≡ E.aleppica L. var. condensata (M. Bieb.)
K. Koch, 1848, Linnaea, 21: 729. — Lectotype
(Geltman, 2015b: 128): “Ex Iberia, circa Tifl is, comm.
Fischer [Wilhelms]” (LE-Bieb.: LE 01070990!; isolecto-
types: LE 01071361–01071363!).
Roadsides, fi elds, fallow lands, dry montane slopes.
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CTC: 8a; ETC: 9a, 9b, 9c, 9e; STC: 11a, 11g; T.—
TCS.— Mediterranean element.
27. E. rigida M. Bieb. I 1808, Fl. Taur.-Cauc. 1:
375. ≡ Tithymalus rigidus (M. Bieb.) Soják, 1972, Čas.
Nár. Mus., Odd. Prír. 140: 176.— Lectotype (I:
Khan, 1964: 147; II: Geltman, 2004: 162): “Ex Tauria
meridionali [Marschall Bieberstein]” (LE-Bieb.: LE
01071047!).— Fig. 4: A.
= E. biglandulosa Desf. VIII 1808, Ann. Mus. Hist.
Nat. (Paris), 12: 114. ≡ Tithymalus biglandulosus (Desf.)
Haw. 1812, Syn. Pl. Succ.: 141. — Lectotype
(Geltman, 2009: 188): icon in Desf., 1808, Ann. Mus.
Paris, 12, tab. 14; epitype (Geltman, 2009: 188):
“Herb. Tournefort, № 122” (P!).
Coastal cliff s.
NWTC; WTC: 7a (Sochi, vicinity of Razdol-
noye).— NCS, TCS.— Submediterranean element.
The species locality in 7a could be an escape from
cultivation (Portenier, Solodko, 2006).
28. E. armena Prokh. 1949, Fl. URSS, 14: 741.
≡ Tithymalus armenus Prokh. 1949, l. c.: 741, nom. al-
tern. ≡ Euphorbia marschalliana Boiss. subsp. armena
(Prokh.) Oudejans, 1992 (publ. 1993), Collect. Bot.
(Barcelona), 21: 186. — Holotype: “Закавказье,
Эчмиадзин, 25 IV 1910, А. А. Гроссгейм [Transcau-
casia, Echmiadzin, 25 IV 1910, A.A.Grossheim]” (LE
01057262!).
Sparse forests, stony or clayey steppes and semide-
serts.
STC: 11a, 11c, 11d.— TCS.— Irano-Turanian ele-
ment.
Closely related to E.marschalliana and recently re-
garded as its subspecies (Pahlevani et al., 2011). The
record for CTC: 8a (Chinthibidze, Gvinianidze, 1983)
belongs to E.marschalliana.
29. E. marschalliana Boiss. 1846, Diagn. Pl. Ori-
ent., ser. 1, 7: 94. ≡ Tithymalus marschallianus (Boiss.)
Klotzsch et Garcke, 1860, Abh. Königl. Akad. Wiss.
Berlin 1859: 86.— Lectotype (I: Rechinger, Schi-
man-Czeika, 1964: 45; II: Geltman, 2004: 164): “in ari-
dis arenosis prope Tatuni ditionis Swant, Georg. cauc.,
VI 1836, R. F. Hohenacker” (G-BOISS!; isolectotypes:
G 00441442, K 001080049!, P 00606900!, 00606901!).
= E. woronowii Grossh. 1916, Trudy Tifl issk. Bot.
Sada, 14: 26. ≡ E.marschalliana Boiss. subsp. woronowii
(Grossh.) Prokh. 1964, Novosti Sist. Vyssh. Rast. 1964
[1]: 232.— Holotype: “Prov. et distr. Erivan, Ar-
asda, an mons Dagna, 10 V 1914, A. Grossheim” (TBI
1025067!).
Stony slopes, steppes and semideserts; 500–2400 m.
CTC: 8c; STC: 11a, 11b, 11c, 11d, 11f; T.— TCS.—
Irano-Turanian element.
30. E. myrsinites L. 1753, Sp. Pl.: 461. ≡ Tithymalus
myrsinites (L.) Hill, 1768, Hort. Kew.: 172.4 (“myrsini-
tis”). — Lectotype (Pahlevani et al., 2011: 490):
“Herb. Linn. №630.68” (LINN!).
= E.pontica Prokh. 1949, Fl. URSS, 1949, 14: 740.
≡ Tithymalus ponticus Prokh. 1949, l. c.: 740, nom. al-
Fig. 3. Euphorbia eugeniae.
A— habit; yellow aspect of plant cover is formed by this species; B— apical synfl orescence.
D. V. Geltman
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Fig. 4. Caucasian species of Euphorbia sections Myrsiniteae and Pithyusa.
A— Euphorbia rigida, habit; E. stepposa: B— habit, C— apical synfl orescence; D— E.petrophila, habit.
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tern. ≡ Euphorbia myrsinites L. subsp. pontica (Prokh.)
R. Turner, 1995, Europ. Gard. Guide: 136.— Holo-
type: “Батумская обл., Артвинский окр., Ардаун,
каменистые склоны горы Варцхет, 26 V 1914,
С.Туркевич, № 452 [Batum region, Artvin district, Ar-
daung, stony slopes of Vartskhet mountain, 26 V 1914,
S. Turkevich, № 452]” (LE 01070946!).
= E. pectinata Albov, 1894, Bull. Herb. Boiss. 2:
640.— Lectotype (Geltman, 2009: 189): “Artwin,
12/25 V 1893, G. R[adde]” (TGM 29031!).
Stony places, montane steppes and pastures, sparse
forests; 300–1100 m.
WTC; CTC: 8a; SWTC: 10a, 10b.— NCS, TCS.—
Submediterranean element.
Prokhanov (1949) recorded E.myrsinites for Crimea,
and E.pontica for the Black Sea coast (mainly Turkey).
I failed to fi nd any signifi cant morphological diff erences
between the Black Sea coast populations from Crimea
and from the Caucasus, especially having in mind the
diversity of E. myrsinites throughout its entire distri-
bution area. However, Falch et al. (2019) discovered
genetic diff erences between Adriatic and some Cauca-
sian populations of E. myrsinites (the Crimean plants
were not studied), but did not propose any taxonomic
changes. In light of these data, the independent status
of E.pontica could be re-evaluated.
Sect. 5. Pithyusa (Raf.) Lazaro
31. E. falcata L. 1753, Sp. Pl.: 456, nom. cons.
≡ Tithymalus falcatus (L.) Klotzsch et Garcke, 1858,
Fl. N. Mitt.-Deutschland, ed. 4: 292.— Lectotype
(Molero, 1992: 715): “Arduino, Herb. Linn. № 630.26”,
right specimen (LINN!).
= E.acuminata Lam. 1788, Encycl. 2: 427. ≡ E.falca-
ta L. var. acuminata (Lam.) St.-Amans, 1818, Fl. Agen.:
189. ≡ Tithymalus falcatus (L.) Klotzsch et Garcke
subsp. acuminatus (Lam.) Soják, 1972, Čas. Nár. Mus.,
Odd. Prír. 140: 172. — Holotype: “Herb. La-
marck” (P-LA 00381891!).
= E.galilaea Boiss. 1853, Diagn. Pl. Orient. 12: 116.
≡ Tithymalus galilaeus (Boiss.) Klotzsch et Garcke, 1860,
Abh. Königl. Akad. Wiss. Berlin 1859: 83. ≡ Euphorbia
falcata L. var. galilaea (Boiss.) Boiss. 1862, in DC., Pro-
dr. 15, 2: 140.— Holotype: “Palaestina, Planitie
Esdraelon, IV–V 1846, E. Boissier” (G-BOISS!).
= E. falcata L. var. ecornuta Boiss. 1879, Fl. Ori-
ent. 4: 1111.— Lectotype (Geltman, 2015b: 128):
“In deserto fl . Chabur, V 1867, Haussknecht, № 863”
(G-BOISS!).
= E. falcata L. var. macrostegia Bornm. 1908,
Mitth. Thüring. Bot. Vereins, n.f., 24: 111. ≡ E.falcata
L. subsp. macrostegia (Bornm.) O. Schwartz, 1934,
Repert. Spec. Nov. Regni Veg. 36: 129. ≡ Tithymalus
falcatus (L.) Klotzsch et Garcke subsp. macrostegius
(Bornm.) Soják, 1972, Čas. Nár. Mus., Odd. Prír. 140:
172.— Lectotype (Geltman, 2015b: 128): “Lydia:
Sinus Smyrnaeus, Ilidja, ad rivilum, 29 V 1906, J.
Bornmüller, № 9961” (B 10 0367965!; isolectotypes:
HBG 516252— image!, LE 01071316!).
Open stony slopes, river banks, dry meadows,
steppes, semideserts and open woodlands, agricultural
and fallow lands, roadsides, human settlements.
WCC; WC: 3a, 3d; CC; EC; NWTC; WTC; CTC;
ETC; SWTC; STC; T.— NCS, TCS.— European-Pan-
tethyan element.
Plants known as E. falcata subsp. macrostegia can be
found throughout the species distribution area, so in my
opinion they do not merit subspecies rank.
This species is probably expanding its distribution
in the Caucasus, because only recently it was found in
the vicinity of Pyatigorsk (EC: 4a), which is a very well
studied area.
32. E. normannii Schmalh. ex Lipsky, 1891, Zap.
Kievsk. Obshch. Estestvoisp. 11 (2): 57. ≡ Tithyma-
lus normannii (Schmalh. ex Lipsky) Prokh. 1949, Fl.
URSS, 14: 466, nom. altern.— Lectotype (Gelt-
man, 2000: 103): “Auf salzigem, sandigem Lehmboden,
beim armenischem Friedhofe um Sengileyevschen See
(Рыбное озеро), 17 V 1879, 6 VI 1883, A. Normann
[Auf salzigem, sandigem Lehmboden, beim armenisch-
em Friedhofe um Sengileyevschen See (Rybnoye Lake),
17 V 1879, 6 VI 1883, A. Normann]” (KW 147556!).
Steppes.
WCC: 1b.— NCS.— Euro-Siberian (Caucasian) el-
ement.
Local endemic, restricted to Stavropol Heights. Ga-
lushko (1980) listed this species for ECC, but without
precise locality.
Recent molecular studies (Frajman, Geltman, in re-
view) have shown that E. normannii is likely an inter-
sectional hybrid, because according to nuclear mark-
ers it is nested in section Myrsiniteae, but according to
chloroplast markers it falls into section Pithyusa. It is
proposed to place it in the latter because of its morpho-
logical similarity to E.falcata.
33. E.seguieriana Necker, 1770, Hist. Commentat.
Acad. Elect. Sci. Theod.-Palat. 2: 493. ≡ Tithymalus se-
guierianus (Necker) Prokh. 1933, Sist. Obzor Moloch.
Sr. Azii: 163. — Lectotype (Geltman, 2006a:
1099): icon in Séguier, 1745, Pl. Veron. 1, tab. 3, f. 1;
epitype (Frajman et al., 2019: 248): “Germany,
Rheinland-Pfalz, Rheintal: Sandhausener Dünen, W of
the village Sandhausen; 111 m, sand dunes; 8°38′16″E,
D. V. Geltman
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49°21′8″N”, 13 VIII 2015, B. Frajman, P. Kirschner”
(WU0098407).
= E. gerardiana Jacq. 1778, Fl. Austriac. 5: 17.
≡ Tithymalus gerardianus (Jacq.) Steud. 1841, No mencl.
Bot., ed. 2, 2: 689. ≡ Euphorbia seguieriana Necker var.
gerardiana (Jacq.) Fiori, 1901, Fl. Italia, 2: 286. —
Lectotype (Geltman, 2015b: 129): “Crescit in pra-
tis Danubialibus passim et in locis arenosis hinc inde,
N.J.Jacquin” (W-Jacq. 0049519!).
= E.fi rma Ledeb. 1850, Fl. Ross. 3: 563. ≡ Tithymalus
fi rmus (Ledeb.) Klotzsch et Garcke, 1860, Abh. Königl.
Akad. Wiss. Berlin 1859: 73. ≡ Euphorbia gerardiana
Jacq. var. fi rma (Ledeb.) Boiss. 1862, in DC., Prodr.
15, 2: 167. ≡ E. seguieriana Necker var. fi rma (Ledeb.)
Oudejans, 1992 (publ. 1993), Collect. Bot. (Barcelo-
na), 21: 187.— Lectotype (Geltman, 2015b: 129):
“Djup Karagan, Eichwald, № 447, Herb. Ledebour” (LE
01071114!, isolectotypes: LE 01071115!, 01071116!).
= E. gerardiana Jacq. var. hohenackeri Boiss. 1862,
in DC., Prodr. 15, 2: 167. ≡ E.seguieriana Necker subsp.
hohenackeri (Boiss.) Rech.f. 1948, Ann. Naturhist. Mus.
Wien, 56: 212.— Lectotype (Geltman, 2006: 163):
“Ad margines Georg. Cauc. VI 1836, R.F.Hohenacker”
(G-BOISS 00418676!, isolectotypes: HBG 516246 —
image!, LE 01071364!, NY 00263339!, P00606937–
00606939!, HAL 118608— image!).
= E. seguieriana Necker var. petrogena Tamamsch.
1944, Dokl. Akad. Nauk Armyanskoi SSR, 1: 46. —
Lectotype (Geltman, 2015b: 131): “Armenia, pr. et
distr. Erivan, 1 VI 1922, S. Tamamschan” (ERE 8672!).
= E. seguieriana Necker subsp. armeniaca Frajman,
2019, Mol. Phylogen. Evol. 134: 249.— Holotype:
“Flora of Armenia: Vajots’dzor, along the street between
villages Gnishik and Agarakadzor (S of Jeghegnaz-
dor), 2127 m; mountain steppe, limestone, 45°18′24″ E,
39°40′32″ N, 28 VII 2012, P. Schönswetter, B. Frajman”
(WU 0098408, isotype: IB 93171).
Sandy and stony steppes and semideserts, pastures,
rocks, sandy and coquina sea shores, river beds, fallow
lands and agricultural fi elds.
WCC; ECC; CC: 4a, 4b; EC; NWTC: 6a; WTC:
7c (Poti); CTC: 8a, 8b; ETC; SWTC; STC; T.— NCS,
TCS.— West-Palearctic element.
A very variable species, especially in leaf shape and
ploidy level (Frajman et al., 2019). It is very likely
could be split into several taxa of varietal, subspecies
or even species rank, although until now no convinc-
ing variants of such splitting have been proposed. The
plants recently described from Armenia as E. segui-
eriana subsp. armeniaca Frajman do have some diff e-
rences according RAPD analysis (Frajman et al., 2019)
and can be found in other parts of the E.seguieriana dis-
tribution area, at least in its eastern part.
E. seguieriana subsp. hohenackeri (Boiss.) Rech. f.
was described from the Caucasus and is accepted by
Govaerts et al. (2020), and Frajman et al. (2019) dis-
tinguished it by elliptic raylet leaves. This character is
likely an abnormality and can be found in plants from
various parts of the species distribution area. This ab-
normality is occasionally found in other species of the
section as well (e.g. E.nicaeënsis All.).
The closely related species E. niciciana Borbás
(=E.seguieriana subsp. niciciana (Borbás) Rech.f.) oc-
curs in the Balkans and West Anatolia and is not pres-
ent in the Caucasus (Frajman et al., 2019).
34. E. macroclada Boiss. 1844, Diagn. Pl. Orient.,
sér. 1, 5: 54. ≡ Tithymalus macrocladus (Boiss.) Klotzsch
et Garcke, 1860, Abh. Königl. Akad. Wiss. Berlin
1859: 97. — Lectotype (Khan, 1964: 119): “Ca-
ria, Denisleh, ad collibus argillosis [Boissier], VI 1842”
(G-BOISS).
= E. schizoceras Boiss. 1844, Diagn. Pl. Orient.,
sér.1, 5: 54. ≡ Tithymalus schizoceras (Boiss.) Klotzsch
et Garcke, 1860, Abh. Königl. Akad. Wiss. Berlin 1859:
98. ≡ Euphorbia tinctoria Boiss. et Huet var. schizoceras
(Boiss.) Boiss. 1862, in DC., Prodr. 15, 2: 166.— Lec-
totype (Geltman, 2015b: 130): “Kurdistan, Berg
Gara, 3 VIII [1841], Th. Kotschy, № 570” (G-BOISS!,
isolectotypes: BM 000951553!, G-DC!, K 001080072!,
LE 01071163!, 01071180!).
= E. syspirensis K. Koch, 1849, Linnaea, 21: 727.
≡ Tithymalus syspirensis (K. Koch) Klotzsch et Garcke,
1860, Abh. Königl. Akad. Wiss. Berlin 1859: 97.— Type
unknown.
= E. damascena Boiss. 1853, Diagn. Pl. Orient.,
sér. 1, 12: 113. ≡ Tithymalus damascenus (Boiss.)
Klotzsch et Garcke, 1860, Abh. Königl. Akad. Wiss.
Berlin 1859: 96. — Lectotype (Geltman, desig-
nated here): “Syria, Damasci collis, E. Boissier, V–VII
1846” (G-BOISS).
= E. tinctoria Boiss. et Huet, 1862, in DC., Prodr.
15, 2: 166. — Lectotype (Geltman, 2006b: 164):
“Elmali, Gémichem quine dans les ravins [Bourgeau],
№ 598, 9 VII 1860” (G-BOISS!, isolectotypes: G-DC
00313297!, MPU014638— image!).
Rocks, stony steppes and semideserts.
STC: 11a.— TCS.— Irano-Turanian element.
Ter-Chatschaturova and Tamamschjan (1973) men-
tioned this species also for SWTC: 10b and STC: 11c,
but these records are not supported by herbarium ma-
terials.
35. E. pannonica Host, 1831, Fl. Austriac. 2: 566.
≡ E. glareosa Pall. ex M. Bieb. var. pannonica (Host)
Nyman, 1881, Consp. Fl. Eur.: 653. ≡ Tithymalus pan-
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nonicus (Host) Á. Löve et D. Löve, 1961, Bot. Not.
114: 40. ≡ Tithymalus nicaeënsis All. subsp. pannonicus
(Host) Soják, 1983, Čas. Nár. Mus., Odd. Prír. 152:
22.— Lectotype (Geltman, 2015b: 130): “Schwe-
chat, 1817, Host” (W 0495171!).
= E.glareosa Pall. ex M. Bieb. var. lasiocarpa Boiss.
1862, in DC., Prodr. 15, 2: 166. ≡E. nicaeënsis All.
var. lasiocarpa (Boiss.) Radcl.-Sm. et Govaerts, 1996,
Kew Bull. 51: 542.— Lectotype (Geltman, 2009:
178): “In campis arenosis Banatue, Heuff el” (G-DC
00313307!).
= E. novorossica Dubovik, 1976, Novosti Sist.
Vyssh. Nizsh. Rast. (Kiev), 1975: 111. — Holo-
type: “Краснодарский край, Анапский р-н, На-
тухаевское лесничество, Лысая гора, опушка, 10
VI 1973, О. Дубовик, А. Краснова [Krasnodar Terri-
tory, Natukhaevskoye forestry, Lysaya mountain, forest
margin, 10 VI 1973, O.Dubovik, A. Krasnova]” (KW
022214!).
Open steppe slopes and forest margins.
NWTC: 6a.— NCS.— Euro-Siberian element.
E. pannonica and the next species, E. stepposa, are
undoubtedly very closely related. E.pannonica is char-
acterized by the presence of a pubescent ovary (capsule
surface) and/or pubescence with short papillary (“car-
tilaginous”) hairs, at least on some parts of the plant. It
occurs in the Pannonian Plain, the Balkans, the north of
Asia Minor; in the Caucasus, the only known location
is the vicinity of Anapa, from where it was described as
E.novorossica Dubovik.
E. stepposa always has a glabrous ovary, and the
leaves and stems without short papillary pubescence.
However, sometimes similar plants can also be found
within the distribution area of E.pannonica. This gives
some reason for considering these taxa as subspecies of
the same species, which in this case would receive the
priority name E.pannonica.
In a number of publications ([Radcliff e]-Smith, Tu-
tin, 1964; Radcliff e-Smith, 1982) E.pannonica, E.step-
posa, as well as E. glareosa Pall. ex M. Bieb. are con-
sidered subspecies, varieties or even synonyms of the
western Mediterranean E.nicaeënsis All. In my opinion,
this point of view is not suffi ciently substantiated, since
E.nicaeënsis is well diff erentiated from the types men-
tioned above by the presence of horn-like appendages
on the glands, and it also has a very distinct geographi-
cal distribution.
36. E. stepposa Zoz ex Prokh. 1949, Fl URSS, 14:
738. ≡ Tithymalus stepposus (Zoz ex Prokh.) Prokh.
1949, l.c.: 738, nom. altern. ≡ Euphorbia glareosa Pall.
ex M. Bieb. subsp. stepposa (Zoz ex Prokh.) Kuzm.
1963, Izv. Bot. Inst. (Sofi a), 12: 126.— E.nicaeënsis All.
subsp. stepposa (Zoz ex Prokh.) Greuter et Burdet, 1981,
Willdenowia, 11: 278. ≡ Tithymalus nicaeënsis Klotzsch
et Garcke subsp. stepposus (Zoz ex Prokh.) Soják, 1983,
Čas. Nár. Mus., Odd. Prír. 152: 23. — Neotype
(Geltman, 1998a: 200): “Харьковская губерния,
близ. г. Старобельска, на меловых горах, 6 VII 1904,
И.Шираевский [Kharkov governance, near Starobelsk,
on chalk hills, 6 VII 1904, I. Shiraevsky]” (LE 01071100!,
isoneotype: LE 01071096!).— Fig. 4: B, C.
= E. klokoviana Railjan, 1973, Bot. Zhurn. 58 (7):
1019. ≡ E.bessarabica Klokov, 1955, Fl. URSR, 7: 629,
nom. illeg., non E.basarabica Prod. 1930. ≡ Tithymalus
klokovianus (Railyan) Holub, 1974, Folia Geobot.
Phytotax. 9: 273. — Holotype: “Бессарабия,
Кринички, 11 VI 1886, В. Липский [Bessarabia, Kri-
nichki, 11 VI 1886, V. Lipsky]” (KW 022206!).
Steppes and steppifi ed meadows, rarely fallow lands;
0–1200 m.
WCC; ECC: 2a; WC: 3c; CC; NWTC: 6a. —
NCS.— Euro-Siberian element.
Records of this species for Daghestan (EC) (Mur-
tazaliev, 2009) likely belong to E.glareosa.
37. E. glareosa Pall. ex M. Bieb. 1808, Fl. Taur.-
Cauc. 1: 373. ≡ Tithymalus glareosus (Pall. ex M. Bieb.)
Prokh. 1949, Fl. URSS, 14: 402, nom. altern. ≡ Euphor-
bia nicaeënsis All. subsp. glareosa (Pall. ex M. Bieb.)
Radcl.-Sm. 1968, Feddes Repert. 79, 1–2: 55.— Lec-
totype (Geltman, 1998a: 199): “in rupestribus ad
Tschorgun, Pallas” (LE 01071068!); epitype (Gelt-
man, 1998a: 199): “ex Tauria et Caucaso [Bieberstein]”
(LE 10070917!).
= E. glareosa Pall. ex M. Bieb. var. minor Boiss. 1879,
Fl. Orient. 4: 1129.— Lectotype (Geltman, 2009:
180): “Georg. cauc., R. F. Hohenacker, V 1838” (G-
BOISS!).
= E. volgensis Krysht. 1929, Izv. Glavn. Bot. Sada
SSSR, 28: 375. ≡ Tithymalus volgensis (Krysht.) Prokh.
1949, Fl. URSS, 14: 403, nom. altern. ≡ Euphorbia nica-
eëensis All. subsp. volgensis (Krysht.) Oudejans, 1992
(publ. 1993), Collect. Bot. (Barcelona), 21: 186. —
Lectotype (Geltman, 1998a: 201): “Саратовская
губ., Камышинский у., меловой холм, версты
полторы к югу от с. Чухонастовки, 27 VI 1926,
Ю. Григорьев, № 1040 [Saratov Governance, Ka-
myshin district, chalk hill, ca. 1.5 versts south of Chuk-
honastovka, 27 VI 1926, Yu. Grigoryev, № 1040]” (LE
01071111!, isolectotype: LE 01071112!).
= E. maleevii Tamamsch. 1944, Dokl. Akad. Nauk.
Armyanskoi SSR, 1 (1–2): 45. ≡ E. glareosa Pall. ex
M.Bieb. subsp. maleevii (Tamamsch.) Tamamsch. 1973,
Fl. Armenii, 6: 108. — Lectotype (Geltman,
2015b: 130): “Distr. Akhty, prope pag. Solak, ad rip.
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dextr. fl . Zanga, in vallecul., schist. et sabulosis, 30 VIII
1942, S. Tamamschan” (ERE 29421!).
Stony slopes, outcrops and rocks, both acid and
limestone, rarely steppes; 400–2000 m.
WCC: 1b; WC: 3d (vicinity of Kichibalyk); EC: 5b,
5c; CTC: 8a, 8b; ETC: 9b, 9c, 9d; SWTC: 10c; STC:
11a, 11b, 11c, 11d.— NCS, TCS.— Submediterranean
element.
In my opinion, to E.glareosa belong only plants with
stems 9–15(19) cm tall and 1.5–2(2.2) mm thick, with
only 3–6 rays of apical synfl orescens. It is a strict petro-
phyte generally avoiding true steppe communities.
38. E. smirnovii Geltman, 1996, Bot. Zhurn. 81
(11): 102.— E.petrophila C. A. Mey. var. armena Boiss.
1866, in DC., Prodr. 15, 2: 1268. — Lectotype
(Khan, 1964: 122): “In ruperstribus vallicum pr. Bai-
burt, 20 VI 1862, E. Bourgeau, № 242” (G!; isolec-
totypes: BR 0000005106028 — image!, E 00202633!,
LD1677755— image!, LE 01071337!, MPU 014224—
image!).
Gravelly slopes.
SWTC: 11a (Arzakan). — TCS. — Irano-Turanian
element.
A rare species, restricted mainly to the periphe-
ry of the Armenian Highlands. From this area it is
known from a single specimen: “Окрестности оз. Гокча,
Арзакенд, щебнистый склон по берегу р. Занги, 9
VIII 1929, П. Смирнов, № 703 [vicinity of the lake
Gokcha [Sevan], Arzakend, gravelly slopes by the bank
of the Zanga river, 9 VIII 1929, P. Smirnov, №703]”
(MW 0690596!).
39. E. petrophila C. A. Mey. 1850, Kleine Beitr.:
9. ≡ Tithymalus petrophilus (C. A. Mey.) Soják, 1972,
Čas. Nár. Mus., Odd. Prír. 140: 175. — Lecto-
type (Geltman, 1998a: 199): “legi in montib. Tau-
riae, Eski-Krym et Tschatyrdagh, 1818, Meyer” (LE
01071082!).— Fig. 4: D.
= E. petrophila C. A. Mey. var. colchica Litv. 1898,
Trudy Imp. S.-Peterburgsk. Bot. Sada, 14 (2): 299.
≡ E. colchica (Litv.) Geltman, 1995, in Czerepanov,
Vasc. Pl. Russia Adj. States: 224. — Lectotype
(Geltman, 2006c: 163): “Черноморский окр., нижнее
течение р. Псезуапсе, на каменистом склоне, 21
VI 1895, Д. Литвинов [Chernomorsky district, low-
er reaches of river Psezuapse, on rocky slope, 21 VI
1895, D. Litvinov]” (LE 01070907!, isolectotypes:
JE00002890— image!, LE 01070908!).
= E. cretophila Klokov, 1955, Fl. URSR, 7: 629. —
Holotype: “Сталинская обл., Славянский р-н,
горы Артема, правый берег р. Донца, на опушке
дубового леса, на меловых обнажениях, часто, 23 VI
1938, З. Сова [Stalino [Donetsk] region, Slavyansk dis-
trict, Artem hills, right bank of the Donets river, margin
of oak forest, on chalk outcrops, numerous, 23 VI 1938,
Z. Sova]” (KW 022209!).
= E. subhastifolia Klokov, 1977, Novosti Sist.
Vyssh. Nizsh. Rast. (Kiev), 1976: 99.— Holotype:
“Краснодарский край, хр. Маркотх, на каменистых
известняковых склонах, на высоте 500 м, 19 V 1951,
А. Колаковский (Список растений Гербария флоры
СССР, № 3688) [Krasnodar Territory, Markotkh range,
on rocky limestone slopes, 500 m, 19 V 1951, A. Kola-
kovsky]” (KW 022234!; isotypes: A00277321 — im-
age!, BM000751480!, G 00441493!, H 1298173!, K
000911992!, LE 01070951!, 01070952!).
Limestone rocks, stony steppes; 300–800 m.
WC (mainly the Skalisty range); CC: 4a; NWTC;
WTC: 7a, 7b.— NCS, TCS.— Submediterranean ele-
ment.
Galushko (1980: 200) records this species for WCC:
1b, but this record most probably belongs to E.glareosa.
40. E. erythrodon Boiss. et Heldr. 1853, Diagn.
Pl. Orient., ser. 1, 12 : 112. ≡Tithymalus erythrodon
(Boiss. et Heldr.) Klotzsch et Garcke, 1860, Abh. Kö-
nigl. Akad. Wiss. Berlin 1859: 87. — Lectotype
(I:Khan, 1964: 124; II: Geltman, 2005: 144): “in rupes-
tribus du Davros-Dagh, 5500′, 29 V 1845 [Heldreich],
№ 759” (G-BOISS!, isolectotypes: BM 000951547!,
G 00441424!, G-DC 00312951!, K 000911990!, LE
01071243!, 01071244!, P 00568286!, 00568287, W
1889-0159589, 0031049!).
= Euphorbia oschtenica Galushko, 1973, Novosti
Sist. Vyssh. Rast. 10: 325. ≡Tithymalus oschtenicus
(Galushko) Galushko, 1979, Fl. Severn. Kavkaza Vopr.
Ist. 3: 56.— Holotype: “Западный Кавказ, юго-
западный склон г. Оштена, 1 VIII 1963, А. Галушко
[West Caucasus, south-west slope of Mt. Oshten,
1VIII 1963, A. Galushko]” (LE 01070942!, isotype: LE
01070941!).
= Euphorbia kotovii Klokov, 1977, Novosti Sist.
Vyssh. Nizsh. Rast. (Kiev), 1976: 102. — Holo-
type: “Крымский государственный заповедник,
Бабуган-яйла, гора Роман-кош, осыпи известняков
на высоте 1400 м, 19 VII 1955, М. Котов, А. Евзеров,
В. Романов [Crimean state reserve, Babugan-yayla,
Mt. Roman-kosh, limestone screes, 1400 m, 19 VII
1955, M. Kotov, A. Evzerov, V. Romanov]” (KW!).
Montane limestone slopes and screes; (700)1200–
2500 m.— Submediterranean element.
WC: 3b (Mt. Oshten ); NWTC: 6a; WTC: 7a. —
NCS.
The montane race of E.petrophila probably deserv-
ing subspecies rank only.
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41. E.panjutinii Grossh. 1950, Bot. Mat. Gerb. Bot.
Inst. Komarova Akad. Nauk SSSR, 13: 18.— Holo-
type: “Предзыбский [Бзыбский] хребет, вершина
г. Напра [Накра], 2400 м, альпийский луг, 28 VII
1936, П. Панютин, № 1246 [Predbzybsky [Bzybsky]
range, top of Napra [Nakra] mountain, 2400 m, al-
pine meadow, 28 VII 1936, P.Panyutin, №1246]” (LE
01070943!, isotype: LE 01070944!).
Alpine meadows on limestone rocks; ca. 2400 m.
WTC: 7b.— TCS.— Submediterranean element.
An enigmatic species known only from the type
specimen. It was not found in the locus classicus during
a special search in 1991.
Sect. 6. Sclerocyathium (Prokh.) Prokh.
42. E. grossheimii Prokh. 1930, Izv. Glavn. Bot.
Sada SSSR, 29: 551. — Holotype: “Nachitsche-
van, 13 V 1929, A. Grossheim” (LE 01070918!).
= E. isthmia Täckh. 1932, Sv. Bot. Tidskr. 26: 374.
≡ Tithymalus isthmius (Täckh.) Soják, 1972, Čas. Nár.
Mus., Odd. Prír. 140: 173. — Holotype: “Sinai:
3–4 km S of Bir Lehfen S of el Arish, 21 III 1928, Täck-
holm” (S G-2570— image!).
= E. mariae Tamamsch. 1944, Dokl. Akad. Nauk Ar-
myanskoi SSR, 1 (1–2): 43.— Lectotype (Gelt-
man, 2015b: 130): “Prope Nachičevan, in steppa, 6 VI
1929, A.Shelkovnikov, E. Kara-Murza” (ERE 8663!).
= Euphorbia cheirolepioides Rech. f. 1955, Dansk
Bot. Ark. 15 (4): 48. ≡ Tithymalus cheirolepioides
(Rech.f.) Soják, 1972, Čas. Nár. Mus., Odd. Prír. 140:
171.— Holotype: “Buschir [Bushehr], 27 II 1937,
M. E. Køie, № 162” (C10011230 — image!, isotypes:
C10011229— image!, W1951-0011260!).
Dry stony slopes, gravelly river banks.
STC: 11a (near Kaghtsrashen), 11d.— TCS.— Ira-
no-Turanian element.
Sect. 7. Chylogala (Fourr.) Prokh.
43. E. heteradena Jaub. et Spach, 1845, Ill. Pl. Ori-
ent. 2: 42. ≡ Tithymalus heteradenus (Jaub. et Spach)
Soják, 1972, Čas. Nár. Mus., Odd. Prír. 140: 173. ≡ Eu-
phorbia megalantha Boiss. var. gracilis Boiss. 1862,
in DC., Prodr. 15, 2: 112. — Lectotype (Rech-
inger, Schiman-Czeika, 1964: 22): “Isphahan, Aucher
№ 5313” (P 00702303!, isolectotypes: W 0031063!,
1889-0077529!).
= E.ispahanica Boiss. 1846, Diagn. Pl. Orient. 7: 91.
≡ Tithymalus ispahanicus (Boiss.) Klotzsch et Garcke,
1860, Abh. Königl. Akad. Wiss. Berlin 1859: 89. —
Lectotype (Khan, 1964: 91): “Ispahan, Aucher,
№5287” (G-BOISS!).
= E. megalantha Boiss. 1846, Diagn. Pl. Orient.
7: 95. ≡ Tithymalus megalanthus (Boiss.) Klotzsch et
Garcke, 1860, Abh. Königl. Akad. Wiss. Berlin 1859:
91. — Lectotype (Geltman, 2013: 8): “in vir-
gultis prope ruinas u. Persepolis sparsim, 20 IV 1842,
Kotschy, № 270” (G-BOISS!, isolectotypes: BM
000951565!, E 00362390!, 00362391!, FI 011592 —
image!, FR 0031292 — image!, G 00441441!, G-DC
00311615!, H 1297808 — image!, HAL 0118586 —
image!, JE 00001691 — image!, K 000911928!, LE
01071217!, 01071219!, MA 205865!, US 00095363!,
W 0031043!).
= E. megalantha Boiss. var. denticulata Boiss. 1862,
in DC., Prodr. 15, 2: 112. — Lectotype (Gelt-
man, 2013: 8): “inter Feisabad et Dirachtanschan (in-
ter Chabbi et Kerman), 1 IV 1859 [Bunge] 24” (G-
BOISS!, isolectotype: LE 01071220!).
= E. megalantha Boiss. var. hirtifl ora Boiss. 1862,
in DC., Prodr. 15, 2: 112.— Holotype: “Assyria,
1836, Aucher, № 1823” (G-DC: G00311613!).
= E. coriacea K. Koch, 1849, Linnaea, 21: 730. —
Lectotype (Geltman, 2013: 8): “trans amnem
Araxen, 1837, Koch, № 867” (LE 01071327!).
= E. froedinii Rech. f. 1952, Symb. Bot. Upsal. 11
(5): 48.— Holotype: “Sarik Sifl a vid Vansjön, 28
VI 1939, Frödin, № 202” (UPS).
Montane rocky slopes, steppes, semideserts, river
banks, sometimes roadsides and fi eld margins; 1000–
2300 m.
STC: 11a, 11c, 11d, 11g.— TCS.— Irano-Turanian
element.
My previous records for SWTC: 10c and STC: 11f
(Geltman, 2012) were not confi rmed by herbarium
specimens, although the occurrence of this species there
is not impossible.
Sect. 8. Szovitsiae Geltman
44. E. szovitsii Fisch. et C. A. Mey. 1835, Index
Sem. Hort. Bot. Petropol. [1]: 27. ≡Tithymalus szovitsii
(Fisch. et C. A. Mey.) Klotzsch et Garcke, 1860, Abh.
Königl. Akad. Wiss. Berlin 1859: 65.— Lectotype
(Geltman, 2000: 102): “Nakitschevan, 1829, Szovits”
(LE 01070953!).
Open rocky and clayey slopes and outcrops, river
banks and beds; 400–2200 m.
EC; WTC: 7e; CTC: 8a, 8c; ETC: 9c; SWTC: 10a,
10c; STC; T.— NCS, TCS.— Irano-Turanian element.
Key to the varieties
1. Raylet leaves linear-oblanceolate, 1–3 mm wide, 3 or
more times longer than wide ......... E.szovitsii var. szovitsii.
D. V. Geltman
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+ Raylet leaves, at least upper ones, ovate-rhombic, 3–8 mm
wide, usually not more than 3 times longer than wide ........
........................................................ E.szovitsii var. kharputensis.
Var. szovitsii.
Open rocky and clayey slopes and outcrops, river
banks and beds; 400–2200 m.— Throughout the whole
area of the species.
Var. kharputensis Azn. ex M. S. Khan, 1964, Notes
Roy. Bot. Gard. Edinburgh, 25: 136.— Holotype:
“Harput to Khan Kezzin, 26 VII 1906, B. Post, № 349”
(G!).
Open rocky slopes.
T.— TCS.
The lectotype of E. szovitsii has two labels with in-
formation on locality: (1) “Prope Naktschevan, 1829,
Szovits” and (2) “in lapidosis montium ad latena saxo-
rum ad Seichaizi ditio Khoi prov. Aderbeidhan, 19 May
1828, Szovits, № 290” as well as the text of the original
species description. There are 6 individuals which pro-
visionally could be separated into 2 sets, but it is impos-
sible to determine to which locality they belong.
Ivanov (1997: 81) recorded E.szovitsii for ECC: 2a,
but this record is not supported by herbarium material.
Sect. 9. Patellares (Prokh.) Frajman
45. E.macroceras Fisch. et C. A. Mey. 1837 (publ.
1838), Index Seminum (LE), 4: 36. ≡ Tithymalus mac-
roceras (Fisch. et C. A. Mey.) Klotzsch et Garcke, 1860,
Abh. Königl. Akad. Wiss. Berlin 1859: 95.— Lecto-
type (Geltman, 1998: 199): “In locis altioribus um-
brosis Cartiliniae prope Malitzki, 10 V 1830, Szovits,
№ 40” (LE 01070936!, isolectotypes: LE 01070931–
01070935!).— Fig. 5: A.
= E.macroceras Fisch. et C. A. Mey. var. spectabilis
K. Koch, 1849, Linnaea, 21: 726.— Type unknown.
Forests, usually beech or with beech; 700–2150 m.
WC: 3b, 3c; CC: 4a, 4c; EC: 5a, 5b; WTC; CTC: 8a,
8c; ETC: 9a, 9c, 9d, 9e; SWTC: 10a, STC: 11a; T. —
NCS, TCS.— Euro-Siberian (Caucasian-Euxinian-Hy-
rcanian) element.
46. E. oblongifolia (K. Koch) K. Koch, 1849, Lin-
naea, 21: 726. ≡ E. amygdaloides L. var. oblongifolia
K.Koch, 1846, Linnaea, 19: 17. ≡ Tithymalus oblongifolius
(K. Koch) Klotzsch et Garcke, 1860, Abh. Königl. Akad.
Wiss. Berlin 1859: 96.— Type unknown.— Fig.5: B, C.
= Euphorbia rumicifolia Boiss. 1860, Cent. Eu-
phorb.: 39. — Lectotype (Geltman, 2002b: 21):
“supra Koeprubachu, V 1853, A. Huet de Pavillon” (G-
BOISS!; isolectotypes: BM001050481!, K 001080062!,
001080063!).
Subalpine meadows and krummholz, sometimes
near streams or on open slopes in forest belt, 1600–
2800 m.
WC: 3b, 3c; 3d; CC: 4c; WTC; CTC: 8a, 8c; ETC:
9a, 9d; SWTC: 10a; T.— NCS, TCS.— Euro-Siberian
(Caucasian-Euxinian-Hyrcanian) element.
47. E. glaberrima K. Koch, 1849, Linnaea, 21:
726. ≡ Tithymalus glaberrimus (K. Koch) Klotzsch et
Garcke, 1860, Abh. Königl. Akad. Wiss. Berlin 1859:
95.— Lectotype (Geltman, 2002a: 106): “In Lori
(parte provinciae Bambaki), 1837, Koch, №870, herb.
C. A.Meyer” (LE 01070916!).
= E. iteophylla Boiss. 1860, Cent. Euphorb.: 39.—
Lectotype (Geltman, 2002b: 22): “In Grusia am
Aragwi-fl üß bei [Gutgore], Sept., Hohenacker, №3811”
(LE 01070927!).
Subalpine meadows and krummholz, sometimes on
rocks, usually on calcareous substrate.
WC: 3b, 3c; CC: 4c; EC: 5c; WTC: 7a, 7b, 7c, 7d;
CTC; ETC: 9a, 9d; SWTC: 10b; STC: 11a, 11b, 11e.—
NCS, TCS.— Euro-Siberian (Caucasian-Euxinian) ele-
ment.
48. E.amygdaloides L. 1753, Sp. Pl.: 463.— Lec-
totype (Geltman, 2002a: 106): “Herb. Linn.
№630.71” (LINN!).
= E.sylvatica L. 1753, Sp. Pl.: 463. ≡ Tithymalus syl-
vaticus (L.) Hill, 1768, Hort. Kew.: 172.4. — Lecto-
type (Geltman, 2002a: 23): “Herb. Linn. № 630.72”
(LINN!).
Forests; 100–1000 m.
EC: 5c, 5d; NWTC: 6b (vicinity of Agoy); WTC;
T.— NCS, TCS.— Euro-Siberian element.
Record for CC: 4a (Galushko, 1980: 224) is likely er-
roneous.
Sect. 10. Herpetorrhizae (Prokh.) Prokh.
Subsect. Oppositifoliae Boiss.
49. E. aserbajdzhanica Bordz. 1928, Izv. Kievsk.
Bot. Sada, 7–8: 19.— Holotype: “Transcaucasia,
Aserbajdzhan, in decliviis lapidosis aridis haud procul
ab oppido Nachiczewan, 6 VII 1927, E. Bordzilowski”
(KW 022205!).
= Tithymalus pseudosororius Prokh. 1930, Izv.
Glavn. Bot. Sada SSSR, 29: 556. — Holotype:
“Нахичевань, 30 VI 1893, В. Липский [Nakhichevan,
30 VI 1893, V.Lipsky]” (LE 01070969!).
Open stony, gypsum or clayey places; 600–1000 m.
SWTC: 10c (Artik); STC: 11a, 11d.— TCS.— Ira-
no-Turanian element.
A synopsis of Euphorbia (Euphobiaceae) for the Caucasus
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63

Fig. 5. Caucasian species of Euphorbia section Patellares.
A— Euphorbia macroceras, habit; E. oblongifolia: B— habit, C— apical synfl orescence.
Sect. 11. Pachycladae (Boiss.) Tutin
50. E.terracina L. 1762, Sp. Pl., ed. 2: 654. ≡ Tithy-
malus terracinus (L.) Klotzsch et Garcke, 1860, Abh.
Königl. Akad. Wiss. Berlin 1859: 90.— Lectotype
(El Hadidi, Fayed, 1978: 50): “Herb. Linn. № 630.33”
(LINN).
— E. boissieriana auct. non (Woronow) Prokh.:
Kolakovsky, 1982, Fl. Abkhazii, ed. 2, 2: 192.
Sandy sea shore.
WTC: 7b (vicinity of Sukhumi).— TCS.— Macaro-
nesian-Mediterranean element.
The record for ETC: 9b (Baku Botanical Garden)
(Musaev, 1988: 69) is not supported by herbarium spe-
cimens.
E. terracina is likely an alien in the Caucasus, al-
though it may have the easternmost limit of its natural
distribution area there.
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Sect. 12. Exiguae (Geltman) Riina et Molero
51. E. exigua L. 1753, Sp. Pl.: 456. ≡ Tithymalus
exiguus (L.) Hill, 1768, Hort. Kew.: 172.3. — Lec-
totype (Jafri, El-Gadi, 1982: 39): “Herb. Linn.
№630.27” (LINN!).
Habitats unknown.
NWTC: 6a (Novorossiisk). — NCS. — European-
Pantethyan element.
E.exigua is known in the Caucasus from two speci-
mens collected in 1892 near Novorossiisk. It is likely
alien there, although may have the easternmost limit of
its natural distribution area there.
Sect. 13. Paralias Dumort.
52. E. paralias L. 1753, Sp. Pl.: 458. ≡ Tithymalus
paralias (L.) Hill, 1768, Hort. Kew.: 172.3. — Lec-
totype (Geltman, designated here): “Herb. Linn.
№199.15” (S 09-28594).— Fig.6.
Sandy sea shores.
NWTC; WTC.— NCS, TCS.— European-Pantet-
hyan element.
53. E. taurinensis All. 1785, Fl. Pedem. 1: 287; 3:
tab. 83, fi g. 2. ≡ Tithymalus taurinensis (All.) Klotzsch
et Garcke, 1860, Abh. Königl. Akad. Wiss. Berlin, 1859:
83.— Lectotype(Khan, 1964: 130): Herb. Allioni
(TO, photo!).
= E.graeca Boiss. et Spruner, 1844, in Boiss., Diagn.
Pl. Orient., sér. 1, 5: 53. ≡Tithymalus graecus (Boiss. et
Spruner) Klotzsch et Garcke, 1860, Abh. Königl. Akad.
Wiss. Berlin, 1859: 84. — Lectotype (Geltman,
2006: 163): “Hymetto, Spruner” (G-BOISS).
Open stony slopes, steppes, open forests (especially
Juniperus), roadsides.
TWTC: 6a. — NCS. — European-Pantethyan ele-
ment.
54. E. ledebourii Boiss. 1860, Cent. Euphorb.:
35. ≡ Tithymalus ledebourii (Boiss.) Prokh. 1949, Fl.
URSS, 14: 462, nom. altern.— Lectotype (Gelt-
man, 2000: 104): “In planitie territorii Elisabethopole-
os, fl ora Transcauc., 21 V 1844, Kolenati, № 1445” (LE
01071318!, isolectotypes: LE 01071319–01071321!).
Open stony places.
ETC: 9d (Ganja), 9f (Shusha); STC: 11f.— TCS.—
Submediterranean element.
Sect. 14. Tithymalus (Gaertn.) Roep.
55. E.peplus L. 1753, Sp. Pl.: 456. ≡ Tithymalus pep-
lus (L.) Hill, 1768, Hort. Kew.: 172.3.— Lectotype
(El Hadidi, Fayed, 1978: 46): “Herb. Linn. № 630.24”
(LINN!).
Rocks and screes, littoral sands, forest margins, as a
weed in settelments and roadsides.
WCC: 1b; CC: 4a; NWTC: 6a; WTC: 7a, 7b, 7c,
7e; ETC: 9b (Baku); T. — NCS, TCS. — European-
Pantethyan element.
Sect. 15. Arvales (Geltman) Geltman
56. E.arvalis Boiss. et Heldr. 1853, in Boiss., Diagn.
Pl. Orient., sér. 1, 12: 116. ≡Tithymalus arvalis (Boiss.
et Heldr.) Klotzsch et Garcke, 1860, Abh. Königl. Akad.
Wiss. Berlin 1859: 82.— Lectotype (Geltman, des-
ignated here): “Plaines d’Isbartha— a 2 lieuer d’Isbartha
sur la route d’Egidir, 31 V 1845 [Heldreich], № 769” (G!,
isolectotypes: BM 001050444!, GOET 03704— image!,
LE 01071140!, 01071141!, P 0552400!, US 01050201!).
= E.ruderalis Scheele, 1843, Linnaea, 17: 343, nom.
illeg., non Dumort. 1827.— Lectotype (Geltman,
Fig. 6. Euphorbia paralias.
A synopsis of Euphorbia (Euphobiaceae) for the Caucasus
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designated here): “In agris incultis et ruderatis prope
Gredück in ditione Swant Georg. Cauc. VI 1836, alt.
4000–5000′, R.F. Hohenacker” (K 000911915 — ima-
ge!; isolectotypes: LE 01071365–01071367!).
= E.parvula K. Koch, 1849, Linnaea, 21: 731, nom.
illeg., non Delile, 1813. ≡ Tithymalus parvulus Klotzsch
et Garcke, 1860, Abh. Königl. Akad. Wiss. Berlin 1859:
93. — Type: “Im russischen Armenien” (?B, de-
stroyed).
— E. punctata auct. non Delile: Ledeb. 1850, Fl.
Ross. 3, 2: 571.
Stony places, montane steppes; 1500–2000 m.
SWTC: 10c; STC: 11a, 11b, 11d; T.— TCS. — Ira-
no-Turanian element.
57. E. aulacosperma Boiss. 1853, Diagn. Pl. Ori-
ent., sér. 1, 12: 117. ≡ Tithymalus aulacospermus (Boiss.)
Klotzsch et Garcke, 1860, Abh. Königl. Akad. Wiss.
Berlin 1859: 82.— Holotype: “Palaestina, Eirusa-
lem, IV–V 1846, E. Boissier” (G-BOISS!).
= E. fossulata Boiss. et Gaill. 1859, in Boiss., Diagn.
Pl. Orient., sér. 2, 4: 87. ≡ E. aulacosperma Boiss. var.
fossulata (Boiss. et Gaill.) Boiss. 1862, in DC., Prodr.
15, 2: 140. — Lectotype (Geltman, designated
here): “Talus pierreax au jard Djebel Khailoun nord
du Damascus, 12 IV 1856, C. Gaillardot, № 2220” (G-
BOISS!, isolectotype: JE 00002431— image!).
Montane steppes and open forests, especially with
Juniperus; 400–800 m.
NWTC: 6a (vicinity of Novorossiisk). — NCS. —
Submediterranean element.
Sect. 16. Esula (Pers.) Dumort.
58. E. iberica Boiss. 1860, Cent. Euphorb.: 38. ≡
Tithymalus ibericus (Boiss.) Prokh. 1933, Sist. Obzor
Moloch. Sr. Azii: 183. — Lectotype (Geltman,
2002a: 110): “In demissis herbidis humidiusculis prope
Helenendorf, V 1835, R. F. Hohenacker” (G-BOISS!;
isolectotypes: G 00441417!, 00441418!, G-DC
00313261!, K 001096700!, 001080075!, LE 01070919–
01070923!).— Fig. 7: A, B.
= E. iberica Boiss. var. intermedia Boiss. 1862, in
DC., Prodr. 15, 2: 163. ≡E. intermedia (Boiss.) Fisch.
et C. A. Mey. ex Trautv. 1844, Trudy Imp. Peterb. Bot.
Sada, 9 (1): 159 (Increm. Fl. Fanerogam. Ross. 3). —
Lectotype (Geltman, designated here): “Caucasus,
Hohenacker” (LE 01070924!).
= E. sanasunitensis Hand.-Mazz. 1912, Ann. K. K.
Naturhist. Hofmus. (Wien), 26: 139. ≡ Tithymalus san-
asunitensis (Hand.-Mazz.) Soják, 1972, Čas. Nár. Mus.,
Odd. Prír. 140: 176. — Lectotype (Khan, 1964:
116): “Kurdistania media, Taurus Armenius, in monte
Meleto (Meretug) Dagh districtus Bitlis, in humo-
sis opimis copiose, 10–11 VIII 1911, Handel-Mazet-
ti, № 2789” (WU 0046584!, isolectotype: W 1913-
0015644!).
= E. kemulariae Ter-Chatsch. 1963, Zametki Sist.
Geogr. Rast. (Tbilisi), 23: 92.— Lectotype (Gelt-
man, designated here): “Мамисон, субнивальный
пояс, 9 VIII 1958, А. Харадзе, Л. Хинтибидзе
[Mamisson, subnival belt, 9 VIII 1958, A. Kharadze,
L. Chinthibidze]” (TBI 1025062!, isolectotype: LE
01070928!).
= E. vedica Ter-Chatsch. 1965, Zametki Sist. Ge-
ogr. Rast. (Tbilisi), 24: 24 (1965). — Holotype:
“АрмССР, Вединский р-н, Зинджириу × Биралу,
28 V 1960, А. Тахтаджян и др. [Armenian SSR,
Vedi district, between Zindzhiriu and Biralu, 28 V
1960, A. Takhtajan et al.]” (ERE 68547, isotype: TBI
1025066!).
Steppes, meadows, open forests, rarely a weed in ag-
ricultural fi elds, fallow lands and ruderal habitats; 10–
2500(3000) m.
WCC; ECC; WC; CC; EC; NWTC: 6a; WTC (al-
ien): 7c, 7d; CTC; ETC; SWTC; STC.— NCS, TCS.—
Euro-Siberian element.
Very common in most areas of the Caucasus. The
species is very variable, especially in the shape of stem
leaves. There are some individuals of E. iberica with
very scarce indumentum in WCC: 1a (Taman Peninsu-
la) and NWTC: 6a, which can be a result of introgres-
sive hybridization with E.dubovikiae.
59. E. dubovikiae Oudejans, 1969, Phytologia,
67 (1): 45. ≡ E. pinetorum Dubovik, 1977, Novosti
Sist. Vyssh. Nizsh. Rast. (Kiev) 1976: 105, nom. il-
leg., non (Small) G. L. Webster, 1967. — Holo-
type: “Краснодарский край, Геленджинский р-н,
Архипо-Осиповка, лес из Pinus pityusa Stev., 27 V
1975, О. Дубовик [Krasnodar Territory, Gelendzik dis-
trict, Arkhipo-Osipovka, forest of Pinus pityusa Stev.,
27 V 1975, O. Dubovik]” (KW 022223; isotype: KW
022224!).
Pine, deciduous and mixed forests.
NWTC: 6b.— TCS.— Euro-Siberian (Crimean-No-
vorossiisk) element.
E. dubovikiae is similar to E. iberica, but diff ers in
leaves with an indumentum and habitat preference: it
occurs in forests, while E.iberica is mainly confi ned to
meadows and steppes.
60. E.agraria M. Bieb. 1808, Fl. Taur.-Cauc. 1: 375.
≡ Tithymalus agrarius (M. Bieb.) Klotzsch et Garcke,
1860, Abh. Königl. Akad. Wiss. Berlin 1859: 89. —
Lectotype (I: Khan, 1964: 113; II: Geltman, desig-
D. V. Geltman
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Fig. 7. Caucasian species of Euphorbia section Esula.
Euphorbia iberica: A— habit, B— fragment of apical synfl orescence; C— E.virgata, habit.
A synopsis of Euphorbia (Euphobiaceae) for the Caucasus
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nated here): “Ex Tauria, 1794 [Marschall Bieberstein]”
(LE-Bieb.: LE 01070980!).
?CC: 4а (Zheleznovodsk — ?alien); ?WTC: 7b. —
?NCS, ?TCS.— Euro-Siberian element.
Steppes and meadows.
I do not know of any current reliable record of
E.agraria for the Caucasus. There are herbarium speci-
mens from the vicinity of Zheleznovodsk (“Zhelezno-
vodsk, 22 VI 1884, Akinfi ev” (KW!)), from Abkhasia
(“Чедым, альпийские луга, Н. Альбов [Chedym, al-
pine meadows, N. Albov] (TBI!)) and one without pre-
cise locality (“Caucasus, G. S. Karelin, Herbarium pro-
prium” (LE 01071329!)). These data are not confi rmed
by later gatherings, and the record for Abkhasia (in
habitat atypical for this species) could be the result of
mislabeling.
E.agraria is common in Crimea and can occur as na-
tive in WCC: 1a, especially on the Taman Peninsula, al-
though it has not been found there yet.
61. E. lucida Waldst. et Kit. 1802, Pl. Rar. Hung.
1: 54, tab. 54. ≡ Tithymalus lucidus (Waldst. et Kit.)
Klotzsch ex Garcke, 1858, Fl. N. Mitt.-Deutschland,
ed. 4: 292. — Lectotype (Chrtek, Skočdopolová,
1982: 212): “Waldstein” (PR 155814/781!).
Wetlands, especially in Colchis lowlands, often near
sea shore.
NTWC: 6a (Novorossiisk); WTC: 7b, 7c, 7d (Lake
Paleostomi).— NCS, TCS.— Euro-Siberian element.
Records for WC: 5c (Zernov, Onipchenko, 2011)
and CTC: 8a (Chinthibidze, Gvinianidze, 1983) belong
to E.iberica.
62. E. virgata Waldst. et Kit. 1803–1804, Pl. Rar.
Hung. 2: 176, tab. 162. ≡ Tithymalus waldsteinii Soják,
1972, Čas. Nár. Mus., Odd. Přír. 140, 3–4: 177. ≡ Eu-
phorbia waldsteinii (Soják) Czer. II 1981, Sosud. Rast.
SSSR: 216. ≡ E. waldsteinii (Soják) Radcl.-Sm. XI
1981, Kew Bull. 36 (2): 216, isonym.— Lectotype
(Chrtek, Skočdopolová, 1982: 224): “in Hungarn” (PR
155811/782, a!).— Fig. 7: C.
= E. repens K. Koch, 1849, Linnaea, 21: 728. —
Neotype (Geltman, 2011: 120): “Араратский
район, травянистые склоны в окрестностях оз.
Карахач, 1750–1950 м, северный склон, 18 VI 1977,
В. Манакян [Ararat district, grassy slopes in the vicin-
ity of Karakhar Lake, 1750–1950 m, northern slope, 18
VI 1977, B. Manakyan]” (LE 01070947!, isoneotype:
ERE!).
= E. virgata Waldst. et Kit. var. orientalis Boiss.
1862, in DC., Prodr. 15, 2: 160. ≡ E.virgata Waldst. et
Kit. subsp. orientalis (Boiss.) Velen. 1891, Fl. Bulg.:
507. ≡ Tithymalus boissierianus Woronow, 1931, Sched.
Herb. Fl. Cauc. 10, № 479. ≡ Euphorbia boissieriana
(Woronow) Prokh. 1949, Fl. URSS, 14: 445. ≡ E. esu-
la L. subsp. orientalis (Boiss.) Molero et Rovira, 1992
(publ. 1993), Collect. Bot. (Barcelona), 21: 163. —
Lectotype (Geltman, 2015b: 132): “Ad margines
hortorum circa Badalan distr. Khoi prov. Aderbeidzhan,
8 VI 1828, Szovits, № 399” (LE 01070958!).
= E. saratoi Ardoino, 1867, Fl. Anal. Alpes-Mar.:
335. ≡ E.esula L. var. saratoi (Ardoino) Fiori, 1901, in
A.Fiori et al., Fl. Italia, 2: 288. ≡ E.esula L. subsp. sara-
toi (Ardoino) P. Fourn. 1936, Quatre Fl. France: 274.
≡ E. waldsteinii (Soják) Czer. subsp. saratoi (Ardoino)
Oudejans, 1992 (publ. 1993), Collect. Bot. (Barcelo-
na), 21: 189.— Lectotype (Molero, Rovira, 1992:
163): “L’Ariane, sur rive droite dupaillon, VI 1864–65,
C.Sarato” (FI).
= Tithymalus hypoleucus Prokh. 1933, Sist. Ob-
zor Moloch. Sredn. Azii: 199. ≡Euphorbia hypoleuca
(Prokh.) Rech. f. 1948, Ann. Naturhist. Mus. Wien 56:
212. — Lectotype (Geltman, 2015b: 132): “Tur-
comania, ad fl . Dajne-ssu, 30 VII 1898, D. Litwinow,
№1985” (LE 01071117!, isolectotype: LE 01071118!).
= E. virgultosa Klokov, 1955, Fl. URSR, 7: 631.
≡ Tithymalus virgultosus (Klokov) Holub, 1973, Folia
Geobot. Phytotax. 8: 175. ≡ Tithymalus tommasinianus
(Bertol.) Soják subsp. virgultosus (Klokov) Soják, 1979
(publ. 1980), Čas. Nár. Mus., Odd. Prír. 148: 79. —
Holotype: “Kiovia, in decliviis ad Borysthemen
in horto urbica, 15 VI 1950, Klokov, Anfi lava” (KW
022239!).
Roadsides, human settelements, forest belts, weed
in agriculturals lands, fallow lands, steppes, slopes and
cliff s by sea shores and river banks; 0–1750 m.
WCC; ECC; WC: 3a, 3b, 3c; CC; EC; NWTC;
WTC: 7a; CTC: 8a, 8c; ETC; SWTC: 10c; STC:
11a, 11b, 11d, 11e, 11f, 11g; T.— NCS, TCS.— West-
Palearctic element.
E. virgata is one of the most problematic taxa of
the genus. In some publications it was accepted as a
subspecies of E. esula ([Radcliff e-]Smith, Tutin, 1968;
Molero, Rovira, 1992; etc.) or even synonymized with
it (e.g. Hämet-Ahti et al., 1984). It my opinion, there
are no grounds for such decision (Geltman, 1998b) and
this point of view was recently shared by Molero et al.
(2012) and Reichert et al. (2017).
However, E. virgata (described from Central Eu-
rope) itself is very variable, so there were several at-
tempts to split it into several more natural taxa. Bois-
sier described E. virgata var. orientalis, which diff ered
from the typical variety by branching stems and elon-
gated leaves. Klokov (1955) believed that true E. vir-
gata occurs in Central Europe and described the closely
related E.virgultosa from Kyiv. Recently Reichert et al.
D. V. Geltman
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68

(2017) proposed a similar approach based on karyologi-
cal and morphological data. According to their opin-
ion, the name E.virgata s. str. belongs to non-branched
plants with narrow leaves and 2n = 20 occurring in
Central Europe only; while for branched plants with
wider leaves and 2n=60 (presumably native to Eastern
Europe, but widely introduced, including North Ameri-
ca) the name E.saratoi should be applied.
Here I prefer to accept a very variable E. virgata
with several ploidy levels as a single species. It is quite
probable that the E. virgata s. l. complex was initially
represented by several geographical races of subspecies
or even species level. However, with the development
of agriculture a polyploid weedy form appeared; it was
widely introduced and now apparently has almost com-
pletely absorbed local races as a result of hybridization.
“Traces” of such races could be found with very caferul
analysis and good experience.
E. virgata defi nitely needs careful studies through-
out its whole distribution area.
Records of E.esula for ECC: 2a and CC: 4a (Tanfi ly-
ev, Kononov, 1987: 68; Ivanov, 1997: 81, Mikheev, 2009:
31) most likely belong to E.virgata.
63. E. pseudagraria P. Smirn. 1940, Byull. Mos-
kovsk. Obshch. Isp. Prir., Otd. Biol. 49, 2: 85. —
Holotype: “Верховья р. Голубой, мел, по
днищу балки, среди Juniperus sabina, близ хут.
Голубинского, 15 V 1938, П. Смирнов [Upper reach-
ers of the river Golubaya, chalk, bottom of ravine,
among Juniperus sabina, near Golubinsky, 15 V 1938,
P. Smirnov]” (MW 0593520!, isotypes: LE 01071084–
01071087!).
Steppes on calcareous substrate.
ECC: 2a. — NCS.— Euro-Siberian element.
64. E. sareptana Becker, 1858, Bull. Soc. Nat.
Moscou, 31 (1): 13. ≡ Tithymalus sareptanus (Becker)
Prokh. 1949, Fl. URSS, 14: 426, nom. altern.— Lec-
totype: (Geltman, 1998a: 39): “Sarepta, 27 V 1851,
A. Becker” (LE 01071089!).
= E. tanaitica Pacz. 1891, Mat. Fl. Step. Donskoi
Obl.: 78. ≡ Tithymalus tanaiticus (Pacz.) Galushko,
1979, Fl. Severn. Kavkaza Vopr. Ist. 3: 56. — Lec-
totype (Geltman, 2015b: 131): “In steppis pr.
stat. Kagalnickaja, Terra Cosacorum Tanaicens, 3 V
1889, I. Paczoski” (KW 022235!, isolectotype: LE
01071102!).
= E. chimaera Lipsky, 1899, Trudy Tifl issk. Bot.
Sada, 4: 444. ≡ Tithymalus chimaerus (Lipsky) Galush-
ko, 1979, Fl. Severn. Kavkaza Vopr. Ist. 3: 56.— Lec-
totype (Geltman, 1998a: 198): “Черноморский
округ, Новороссийск, [11] V 1892, Липский [Cherno-
morsky District, Novorossiisk, [11] V 1892, V. Lipsky”
(LE 01057270!).
= E. chimaera Lipsky var. kimmerica Lipsky, 1899,
Trudy Tifl . Bot. Sada, 4: 444. ≡ E. kimmerica (Lip-
sky) Grossh. 1932, Fl. Kavkaza, 3: 39. — Lecto-
type (Geltman, 2011: 120): “Черноморский округ,
Новороссийск, 11 V 1892, Липский [Chernomor-
sky District, Novorossiisk, 11 V 1892, V. Lipsky” (LE
01070905, isolectotype: LE 01070906!).
Steppes.
WCC; WC: 3a; NWTC: 6a.— NCS.—Euro-Siberi-
an element.
An enigmatic species occasionally found in steppes
of Eastern Europe and the Northern Caucasus, some-
times together with E.subtilis.
65. E. subtilis (Prokh.) Prokh. 1949, Fl. URSS,
14: 421.— E.gracilis M. Bieb. 1819, Fl. Taur.-Cauc. 3:
324, nom. illeg., non Loisel. 1807. ≡ Galarhoeus subti-
lis Prokh. 1941, Trudy Kuibyshevsk. Bot. Sada, 1: 48.
≡ Tithymalus subtilis (Prokh.) Prokh. 1949, l. c.: 421,
nom. altern.— Lectotype (Geltman, 1998a: 200):
“Kursk — Belgorod, 1812 [Marschall Bieberstein]”
(LE-Bieb.: LE 01071009!).— Fig. 8.
= E. baxanica Galushko, 1970, Novosti Sist.
Vyssh. Rast. 6: 216. ≡ Tithymalus baxanicus (Galushko)
Galushko, 1979, Fl. Severn. Kavkaza Vopr. Ist. 3: 56.—
Neotype (Geltman, 2015b: 127): “Кабардино-
Балкария, Баксан, р. Гижгит, в 4 км выше устья, на
скалах, 5 VI 1960, А. Галушко [Kabardino-Balkaria,
Baksan, Gizhgit River, 4 km above its mouth, on rocks,
5 VI 1960, A. Galushko]” (LE 01057265!).
= E. meyeriana Galushko, 1970, Novosti Sist.
Vyssh. Rast. 6: 216. ≡ Tithymalus meyerianus (Galush-
ko) Galushko, 1979, Fl. Severn. Kavkaza Vopr. Ist.
3: 56. — Holotype: “In rupestribus versus fl uv.
Kashaut, 5000–6000 ped., 3 VII [1834] [Meyer]” (LE
01070938!).
Steppes, steppifi cated meadows, rocks; 300–1600 m.
WCC; WC: 3c, 3d; CC: 4b; EC: 5b; NWTC; CTC:
8a (Borjomi).— NCS, TCS.— Euro-Siberian element.
This species is common in steppes of Eastern Europe
and the Northern Caucasus, but also penetrates into
arid montane areas.
66. E. daghestanica Geltman, 1997, Bot. Zhurn.
82 (3): 122. — Holotype: “Дагестанская АССР,
Левашинский р-н, окр. с. Цуда[к]хар, h — 1200 м,
в трещинах скал, 10 VII 1961, Н. Н. Цвелев и др.,
№3215 [Daghestan ASSR, Levashi district, vicinity of
Tsudakhar, h — 1200 m, in rock crevices, 10 VII 1961,
Tzvelev et al., № 3215]” (LE 01070912!).
Rocks and dry stony slopes; 900–3700 m.
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CC: 4c; EC: 5a, 5b, 5c; STC: 11e, 11g. — NCS,
TCS.— Euro-Siberian (Caucasian) element.
67. E.leptocaula Boiss. 1862, in DC., Prodr. 15, 2:
159. ≡ Tithymalus leptocaulus (Boiss.) Prokh. 1949, Fl.
URSS, 14 : 443, nom altern.— Lectotype (Gelt-
man, 2002a: 118): “Taur[ia], Steven” (G!, isolectotypes:
LE-Bieb.: LE 01071025!; LE 01071078!, 01071079!).
Steppes, dry stony slopes; 0–550 m.
WCC; ECC; WC: 3a; NWTC: 6a.— NCS.— Euro-
Siberian element.
The record for West Transcaucasia (Gouria) (Lede-
bour, 1850: 572) appeared by mistake (see Geltman,
2002a: 118). The record for WC: 3c (Zernov, Onipchen-
ko, 2011: 133) in fact refers to E.subtilis.
68. E. astrachanica C. A. Mey. ex Trautv. 1844,
Trudy Imp. Peterb. Bot. Sada, 9 (1): 156 (Increm. Fl.
Fanerogam. Ross. 3). ≡ E. leptocaula Boiss. var. prae-
cox Boiss. 1862, in DC., Prodr. 15, 2: 159.— E.praecox
(Boiss.) B.Fedtsch. et Flerow, 1909, Fl. Evrop. Rossii:
614. — Lectotype (Geltman, 2002a: 119): “Pl.
Wolga infer., locis salsis argillosis, Maio, A. Becker” (G!,
isolectotypes: LE 01071063!, 01071064!).
— E.praecox M. Bieb. ex Fisch. 1808, Cat. Jard. Pl.
Gorenk.: 120, nom. inval.
— E.astrachanica C. A. Mey. ex Claus, 1851, Beitr.
Pfl . Russ. Reich. 8: 254, nom. inval.
Steppes, usually on clayey substrates.
WCC.— NCS.— Irano-Turanian element.
Recorded for ECC: 2a, 2b (Galushko, 1980: 201;
Terekbaev, 1991: 94), but herbarium material is not
available. My previous record for EC: 5b (Geltman,
1996: 100) is erroneous and belongs to Е. daghestanica.
69. E. undulata M. Bieb. 1808, Fl. Taur.-Cauc. 1:
371. ≡ Tithymalus undulatus (M. Bieb.) Klotzsch et
Garcke, 1860, Abh. Akad. Wiss. Berlin 1859: 92. —
Lectotype (Geltman, 1998a: 201): “Ex dit. Wolgi-
cis: inter Astrachan et Zarizyn, 1800 [Marschall Bieber-
stein]” (LE-Bieb.: LE 01071058!).
Semideserts and dry steppes.
Can be found in ECC.— NCS— Euro-Siberian ele-
ment.
I know of a single specimen without indication of
the precise locality: “Caucasus, herb. Fischer” (LE
01071328!). Grossheim (1932: 37) mentioned this
species for “North Caucasus” and Tamamschyan (1962:
95, map 107) recorded on his map a single locality
in ECC: 2b. Galushko (1980: 201) believed that this
species occurs in several places of the central part of
Pre-Caucasia, but herbarium materials are not known.
Having in mind the distribution of this species and
Fig. 8. Euphorbia subtilis.
D. V. Geltman
Новости систематики высших растений | Том 51 | 2020
70

its habitat preferences, its occurrence in ECC is very
probable.
70. E.buschiana Grossh. 1940, Bot. Zhurn. 25 (4–
5): 330. ≡ Tithymalus buschianus (Grossh.) Soják, 1972,
Čas. Nar. Mus., Odd. Prír. 140: 170.— Holotype:
“р. Сукан, верховья, урочище Сукан-баши-цифи,
2400 м, лев. борт Сукана, 29 VII 1931, Е. и Н.Буш
[River Sukan, upper reaches, Sukan-bashi-tsifi terrain
feature, 2400 m, left bank of Sukan, 29 VII 1931, E. and
N. Busch]” (LE 01057266!, isotype: LE 01057267!).
Rocks in subalpine and alpine belt; 1400–2700 m.
CC: 4b, 4c; EC: 5b (Botlikh district, vicinity of
Alak).— NCS.— Euro-Siberian (Caucasian) element.
Subgen. 2. Chamaesyce Raf.
Sect. 1. Anisophyllum Roep.
71. E. nutans Lag. 1816, Gen. Sp. Pl.: 17.
≡ Chamaesyce nutans (Lag.) Small, 1903, Fl. S. E. U.S.:
712.— Lectotype (Wheeler, 1941: 144: “Habit. in
N[ova] H[ispania], J.D. Rodriguez” (MA 250449— im-
age!).
Roadsides, human settlements, agricultural fi elds,
pasturtes, sea shores and river banks, alien; 0–700 m.
WC: 3b; CC: 4a; EC: 5b; NWTC: 6a; WTC: 7a, 7b,
7c, 7d; CTC: 8a; ETC: 9a, 9b; STC: 11d.— NCS, TCS.
Grossheim (1949) mentioned E.indica Lam. for the
Apsheron Peninsula (ETC: 9b) and Talysh. Re-exam-
ination of corresponding herbarium material from the
Apsheron Peninsula (BAK) showed that in fact they
belong to E.nutans.
72. E. peplis L. 1753, Sp. Pl.: 455. ≡ Chamaesyce
peplis (L.) Prokh. 1933, Sist. Obzor Moloch. Sr. Azii:
15.— Type not designated.
Sea shores.
WCC: 1a; NWTC; WTC: 7a, 7b, 7c, 7d. — NCS,
TCS.— Macaronesian-Mediterranean element.
73. E.humifusa Willd. 1813, Enum. Pl. Horti Ber-
ol., suppl.: 27. ≡ Chamaesyce humifusa (Willd.) Prokh.
1927, Izv. Akad. Nauk SSSR, 3–4: 195. — Type not
designated.
= E. pseudochamaesyce C. A. Mey. 1842 (publ.
1843), Index Seminum (LE), 9: 73. ≡Chamaesyce hu-
mifusa (Willd.) Prokh. var. pseudochamaesyce (Fisch.
et C. A. Mey.) Hurus. 1954, J. Fac. Sci. Univ. Tokyo,
Sect. 3, Bot. 6: 288. ≡ Euphorbia humifusa Willd. var.
pseudochamaesyce (Fisch. et C. A. Mey.) Murata,
1962, Acta Phytotax. Geobot. 20: 198. — Lecto-
type (Baikov, 2007: 107): “In rupestribus inter Bu-
chtarminsk et Woronenskoi, 1840, Karelin et Kiriloff ”
(LE 01071324!).
River banks, pastures and fallow lands, roadsides,
agricultural lands; ?alien.
ECC; WC: 3c; CC: 4a, 4c; EC; WTC: 7b, 7c; CTC:
8a; ETC: 9e, 9f.— NCS, TCS.— South-Palearctic ele-
ment.
The lectotype selection of E.pseudochamaesyce pro-
posed by Baikov (2007) cannot be regarded as fully
suitable. One cannot be sure that this specimen was de-
fi netely seen by the author of the name, and, hence, may
not represent a part of the original material.
E.humifusa is a species of Asian origin, but was in-
troduced to Europe long ago. In the north and north-
eastern parts of the Caucasus it could be native, but in
other areas it is most probably alien.
74. E. chamaesyce L.1753, Sp. Pl.: 455.
≡ Chamaesyce vulgaris Prokh. 1941, Trudy Kuiby-
shevsk. Bot. Sada, 1: 8.— Lectotype (Khan, 1964:
152): “Löfl ing, №373, Herb. Linn. № 630.15” (LINN!).
= E.canescens L. 1762, Sp. Pl., ed. 2: 652. ≡ Chamae-
syce canescens (L.) Prokh. 1933, Sist. Obzor Moloch.
Sr. Azii: 19. ≡ E. chamaesyce L. subsp. canescens (L.)
Prokh. 1964, Novosti Sist. Vyssh. Rast. [1]: 237. —
Lectotype (Benedí González, Orell, 1993: 149):
“Alströmer, № 146a, Herb. Linn. № 630.16” (LINN!).
= E. massiliensis DC. 1815, in DC. et Lam., Fl. Fr.,
ed. 2, 5: 357. ≡ E.chamaesyce L. var. massiliensis (DC.)
Thell. 1917, in Ascherson u. Graebner, Syn. Mitteleur.
Fl. 7: 457. ≡Chamaesyce canescens (L.) Prokh. subsp.
massiliensis (DC.) Soják, 1972, Čas. Nár. Mus., Odd.
Prír. 140: 169. ≡ Chamaesyce massiliensis (DC.) Galush-
ko, 1974, Novosti Sist. Vyssh. Rast. 11: 299.
River banks and sea shores, steppes and semide-
sets, agricultural and fallow lands, human settlements;
5–550 m.
WCC; EC; NWTC: 6a; WTC; CTC: 8a; ETC: 9b,
9e; STC; T.— NCS, TCS.— European-Pantethyan ele-
ment.
E.chamaesyce varies in the presence and density of
the indumentum. Prokhanov (1949, 1964) treated gla-
brous and semigrabrous forms as E. chamaesyce s. str.
(or E.chamaesyce subsp. chamaesyce) and pilose forms
as E. canescens (or E. chamaesyce subsp. canescens).
There are some grounds for this point of view: e.g. in
the southeast of European Russia mainly glabrous forms
are found. However, in other areas both forms are usu-
ally present in the same local populations.
Here I follow Pahlevani, Riina (2011) and accept
E. chamaesyce in the broad sense. At the same time
careful revision of this group in all its vast distribution
area is absolutely necessary.
T
A synopsis of Euphorbia (Euphobiaceae) for the Caucasus
Novitates Systematicae Plantarum Vascularium | Volume 51 | 2020
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75. E.glyptosperma Engelm. 1858, in W. H. Emory,
Rep. U.S. Mex. Bound. 2(1): 186. ≡ Chamaesyce glypto-
sperma (Engelm.) Small, 1903, Fl. S.E. U.S.: 712. —
Lectotype (Wheeler, 1937: 496): “Fort Kearney
on the Platte, VII 1856, Engellmann” (MO 144635—
image!).
= E. glyptosperma Engelm. var. tenerrima Engelm.
1858, in W. H. Emory, Rep. U.S. Mex. Bound. 2 (1):
187. — Lectotype (Wheeler, 1941: 235): “[peb-
bly bars of the Nueces River, Texas, 25 V 1851]
C.Wright, № 1853” (MO 144673/2196682— image!;
isolectotypes: F 0056275F — image!, GH 00047685!,
00047686 — image!, K 001080336!, 001080340 — im-
age!, NY 00263120— image!).
= E. glyptosperma Engelm. var. pubescens Boiss.
1862, in DC., Prodr. 15, 2: 48. ≡Chamaesyce glypto-
sperma (Engelm.) Small var. pubescens (Boiss.) Millsp.
1913, Living Fl. W. Va, 5A (1): 294.— Lectotype
(Wheeler, 1941: 235): “Bords du Missisippi en Illinois,
VIII 1846, Riehl, № 472” (G!).
Steppes and semideserts, alien.
WCC: 1b; ECC: 2b.
This North American plant was collected for the
fi rst time in the Caucasus in 1987–1988, but it has only
recently been properly named (Geltman, Medvedeva,
2017). It could also be found in other steppe areas of the
Caucasus.
76. E. granulata Forssk. 1775, Fl. Aegypt.-Arab.:
94. ≡ Chamaesyce granulata (Forssk.) Soják, 1972, Čas.
Nár. Mus., Odd. Prír. 140: 169.— Type not designated.
= E. turcomanica Boiss. 1860, Cent. Euphorb.: 13.
≡ Chamaesyce turcomanica (Boiss.) Prokh. 1933, Sist.
Obzor Moloch. Sr. Azii: 21. ≡ Euphorbia granulata
Forssk. var. turcomanica (Boiss.) Hadidi, 1973, Bull.
Jard. Bot. Natl. Belg. 43: 93. — Lectotype:
(Rechin ger, Schiman-Czeika, 1964): “In littore orientali
maris Caspii, 1834, D. Karelin” (G-DC!).
Steppes and semideserts.
ETC: 9e (Mugan steppe); STC: 11d, 11f.— TCS.
I follow Radcliff e-Smith (1980) and Pahlevani, Rii-
na (2011) and treat E.turcomanica Boiss. as a synonym
of E.granulata Forssk.
77. E.maculata L. 1753, Sp. Pl.: 455. ≡ Chamaesyce
maculata (L.) Small, 1903, Fl. S.E. U.S.: 713.— Lec-
totype (Croizat, 1962: 191): “21. maculata; Herb.
Linn. №630.11” (LINN!).
Riverbanks and seashores, pastures, roadsides, hu-
man settlements; alien; 0–1500 m.
WCC: 1a; CC: 4a, 4c; NWTC: 6a; WTC; CTC:
8a; ETC: 9a, 9b, 9d; STC: 11a, 11f, 11g; T. — NCS,
TCS.
78. E.forskaolii J. Gay, 1847, in Webb et Berthelot,
Phytogr. Canar. 2 (3): 240. ≡Chamaesyce forsskaolii
(J.Gay) Prokh. 1949, Fl. URSS, 14: 493, “forsskalii”.—
Type not designated.
Agricultural lands; alien.
WTC: 7c; ETC: 9e; T.— TCS.
Sect. 2. Poinsettia (Graham) Baill.
79. E. davidii Subils, 1984, Kurtziana, 17: 125. —
Holotype: “Argentina: Córdoba: Depto. Río
Cuarto, Alpa Corral, Arroyo Las Moras, cerca de la
unión con Arr[oyo] el Talita, 19 Nov 1982, Subils &
Moscone 3115” (CORD).— Fig. 9.
Roadsides, human settlements, agricultural lands;
alien; 300–680 m.
WCC: 1a; ECC: 2b; EC: 3a, 3c; CC: 4a, 4c; EC: 5a;
WTC: 7a, 7b.— NCS.
First collected in 1968 in Pyatigorsk and named
E. dentata Michx. (Mikheev, 1971). The record of
E. geniculata Ortega for the same locality (Dorofeyev,
2011) refers to E.davidii as well.
Sect. 3. Alectoroctonum (Schltdl.) Baill.
80. E. marginata Pursh, 1814, Fl. Amer. Sept.
2: 607. ≡ Tithymalus marginatus (Pursh) Cockerell,
1909, Torreya, 9: 119. ≡ Agaloma marginata (Pursh)
Á.Löve et D. Löve, 1961, Bot. Not. 114: 40.— Type not
designated.
Fig. 9. Euphorbia davidii.
D. V. Geltman
Новости систематики высших растений | Том 51 | 2020
72

Cultivated as an ornamental plant. Known as
escaping from cultivation wild in CC: 4a, 4c, could be
found in other regions.
Discussion
The current Euphorbia treatment for the Caucasus
contains 80 species. From this number, 73 species
are highly likely indigenous and/or archaeophytes
(including E. agraria, E. platyphyllos and E. undulata
which could be found in the area although currently
there are no reliable records). There are also 7 alien
species, most of them (5) of New World origin.
The geographical analysis of the indigenous com-
ponent of Caucasian Euphorbia (see Table) shows that
the majority of species belong to Euro-Siberian, Irano-
Turanian and Submediterranean elements. The Euro-
Siberian geographical element evidently dominates
(41.10 %). However, if we apply a narrower approach
to the defi nition of geographical elements for the Cau-
casus (e.g. according to Portenier (2000) and Geltman
(2010)), one can see that 7 species of Euro-Siberian
element in the “broad sense” could be characterized as
Caucasian (E. ardonensis, E. aristata, E. czerepanovii,
E.daghestanica, E.normannii, E.procera, E.wittmannii),
3— Euxinian (E.djimilensis, E.eugeniae and E.scripta),
1— Caucasian-Euxinian (E.oblongifolia) and 2— Cau-
casian-Euxinian-Hyrcanian (E.macroceras and E.squa-
mosa). So, about 43% of Euro-Siberian Caucasian Eu-
phorbia species are restricted to three main fl oristic
provinces of the Caucasus (Caucasian, Euxinian and
Hyrcanian).
The Irano-Turanian element is the second largest
one by number of species (15, or 20.54 %); these spe-
cies occur mainly in Transcaucasia. An important num-
ber of species come from the Submediterranean element
(15.07 %) which includes (Geltman, 2016): (a) spe-
cies occurring in the Mediterranean region itself (usu-
ally at higher elevations) and also in neighboring areas
(E.myrsinites, E.rigida); (b) species found in the Medi-
terranean and/or at the western edge of the Irano-Tura-
nian region and at the same time in southern parts of
the Circumboreal region, where such plants are usually
restricted to “Mediterranean-like” habitats (E. aulaco-
sperma, E.condylocarpa, E.glareosa, E.ledebourii, E.mi-
crosphaera, E. petrophila, E. rhabdotosperma); (c)spe-
cies endemic to the mountain systems located at the
northern border of the Mediterranean, or local endem-
ics to the areas adjoining the Mediterranean (E.eryth-
rodon and E.panjutinii).
Eight species (10.96 %) belong to the European-
Pantethyan element. These plants are mostly weeds of
Mediterranean origin, but they also occur in Europe
and/or the Irano-Turanian region.
Mediterranean and Macaronesian-Mediterranean
species are poorly represented in the Caucasus and
include species restricted to sea shores (E. hirsuta,
E. peplis, E. terracina), and possible archaeophytes
(E. aleppica). The data on Euphorbia shows that there
are no true enclaves of Mediterranean fl ora in the
Caucasus.
Taxonomically most of Caucasian Euphorbia
species (70, or 87.5%) belong to subgen. Esula, which
is perfectly expected for temperate fl oras of Eurasia.
These species belong to 16 sections of the 21 currently
accepted sections in the subgenus (Riina et al., 2013).
By the number of species, section Helioscopia is the
largest (23 species) followed by sections Esula (13
species) and Pithyusa (11 species). The representatives
of these 3 sections comprise about 67% of the number
of species in the subgenus. Other sections contain 5
or less species, and 9 sections are represented by a
single species. Subgen. Chamaesyce is represented
by 10 species (of 3 sections) and only 3 of them are
indigenous.
Conclusion
The taxonomy and geography of Caucasian
Euphorbia species are of great interest because of high
biodiversity signifi cance of the area. The following
directions for future studies are the most notable.
1. Careful investigation of critical taxa, fi rst of
all E. virgata s. l. and E. chamaesyce s. l. Such studies
should include several techniques (including molecular
and karyological) and employ material from the entire
distribution area; data from the Caucasus are critical to
produce adequate results.
2. Although the distribution patterns of most native
Caucasian Euphorbia are more or less clear, some new
important data could be obtained. Special attention
Table. Geographical elements of indigenous fraction of
Caucasian Euphorbia
Geographical element Number
ofspecies
Percentage of total
number ofindigenous
species
Palearctic 1 1.37
West Paleartctic 2 2.74
Euro-Siberian 30 41.10
Pantethyan 1 1.37
European-Pantethyan 8 10.96
Mediterranean 2 2.74
Macaronesian-Mediterranean 3 4.11
Submediterranean 11 15.07
Irano-Turanian 15 20.54
TOTAL 73
A synopsis of Euphorbia (Euphobiaceae) for the Caucasus
Novitates Systematicae Plantarum Vascularium | Volume 51 | 2020
73

should be paid to lowland areas of Ciscaucasia,
especially still unknown but preserved spots of native
steppe vegetation; regarding Transcaucasia, there
are also some underinvestigated areas, especially in
Azerbaij an and southern and southwestern Georgia
(Ajara and Samtskhe-Javakheti).
3. It is important to clarify the current status of
some wetland species (E.hirsuta, E. lucida, E.palustris)
known from specimens that are 60 or more years old.
4. The distribution dynamic of alien species,
especially E.davidii and E.nutans, needs careful studies;
it would also be important to trace the current status of
E.forskaolii, known only from old gatherings. A special
search for possible new aliens is also very important.
5. According to my calculations at the moment at least
some molecular data are available for 59 Euphorbia spe-
cies occurring in the Caucasus and data for 21 species
were received using the material collected in the Cauca-
sus. So, additional new sample collecting and analyses are
necessary. Besides, molecular techniques could be used for
phylogeographic research which could clarify the history
of the distributions of some species (e.g. E.amygdaloides,
E.macroceras or E.squamosa), as well the affi nity of Cau-
casian species with those occurring in neighboring areas.
Ackowlegements
The work was carried out in the framework of the
institutional research project of the Komarov Botani-
cal Institute (№ AAAA-A19-1190312900521-1). I am
very grateful to Paul Berry for linguistic editing of the
manuscript and valuable suggestions which improved
the text. I also grateful to Irina Sokolova for her great
help in the preparation of illustrations and many use-
ful advices, and Ketevan Batsatsashvili for the transla-
tion of some texts from Georgian. Suggestions from two
anonymous reviewers are also very much appreciated.
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