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Feddes Repertorium 114 (2003) 5 – 6 , 350 – 357
DOI: 10.1002/fedr.200311003
Berlin, September 2003
Moscow State University M. V. Lomonossov, Botanical Garden, Moscow
M. G. PIMENOV & E. V. KLJUYKOV
What is Sium frigidum HAND.-MAZZ. (Umbelliferae)?
With 3 Figures
Summary
Zusammenfassung
A rare and enigmatic Chinese species, known as
Apium ventricosum H.BOISSIEU and Sium frigidum
HAND.-MAZZ. (the latter name is only used in current floristic literature), has been shown to be closer
related to Sinocarum H.WOLFF ex SHAN RENHWA &
PU FATING. This affinity, indicated recently by ITS
sequence investigation, was confirmed by morphological analysis. Attribution of the species to Apium,
Sium, or Chamaesium, was not confirmed by molecular and morphological data either. A new combination, Sinocarum ventricosum (H.BOISSIEU)
PIMENOV & KLJUYKOV, has been proposed. The
isolated position of S. ventricosum within the genus
Sinocarum is emphazised by separation of a new
monotypic section, Sinocarum sect. Apiopsis
PIMENOV & KLJUYKOV.
Was ist Sium frigidum HAND.-MAZZ. (Umbelliferae)?
Introduction
with other Chinese HANDEL-MAZZETTI’S Umbelliferae to H. WOLFF, but remained unpublished by the monographer, probably due to the
absence of ripe fruits and the uncertainty about
the genus to which the species could be referred. HANDEL-MAZZETTI believed the information sufficient to describe the species as a
new Sium species. The reason for the attribution of a new species to Sium was probably the
structure of leaves, being slightly similar to
pinnate leaves of various Sium species. But
even leaves in S. frigidum resemble Sium-type
very approximately, as the leaflets are dissected, sometimes deeply, and not serrate. The
unusual habit of the plant could be, however,
In 1933 the prominent Austrian explorer of
Chinese flora, Prof. H. F. HANDEL-MAZZETTI
described a rare and very unusual Sium from
the NW part of Yunnan province in SW China,
S. frigidum HAND.-MAZZ. In its habit this small
(up to 15, rarely 20 cm tall) high-mountainous
plant is in disharmony with all other known
species of the genus. It has 2 – 5-rayed umbels
without involucre or with one narrow-linear
bract; umbelets are 5 – 7-flowered, without involucel or with one bractlet. Calyx teeth absent.
Petals white, ovate, without claw, with short
incurved tip. The species was sent together
© 2003 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim
Eine seltene und rätselhafte chinesische Art, bekannt als Apium ventricosum H.BOISSIEU und Sium
frigidum HAND.-MAZZ. (letzterer Name wird in der
aktuellen floristischen Literatur allein benutzt),
scheint näher verwandt zu sein mit Sinocarum
H.WOLFF ex SHAN RENHWA & PU FATING. Diese
neuerlich durch IST-Sequenzanalyse festgestellte
Affinität wurde durch molekulare und morphologische Daten bestätigt. Der Einschluss der Art in die
Gattungen Apium, Sium oder Chamaesium konnte
nicht durch molekulare und morphologische Merkmale bestätigt werden. Es wird eine Neukombination: Sinocarum ventricosum (H.BOISSIEU) PIMENOV
& KLJUYKOV vorgeschlagen. Die isolierte Stellung
von S. ventricosum innerhalb der Gattung Sinocarum wird durch die Aufstellung einer monotypischen Sektion, Sinocarum sect. Apiopsis PIMENOV &
KLJUYKOV unterstrichen.
0014-8962/03/5-610-0350
M. G. PIMENOV & E. V. KLJUYKOV: What is Sium frigidum HAND.-MAZZ.?
connected with its growth under extreme conditions of high mountains. The plant life-form
also accords with ecology. As we could observe during our botanical trip to SW China, it
grows in wet Cobresia grasslands and could be
hardly extracted from Cobresia turf. So, in
herbaria S. frigidum is frequently represented
without its underground part. We tried to prepare its roots (slightly thickened) and vertical
rhizomes (with adventive roots in nodes), but it
remains are not clear, whether the species have
vegetative propagation. S. frigidum was included in standard Chinese flora (FRPS, 55,3 –
PU FATING et al. 1992), and pictured in it. The
picture corresponds to the type material, seen
by us in WU (Fig. 1), but the mericarp crosssection is rather incorrect (see below). The
species was cited also by WU ZHENG YI (1984).
Other hypotheses on generic attribution of
Sium frigidum
SHAN RENHWA ex PU FATING (1993) proposed
to transfer the species to the genus Chamaesium H.WOLFF under the name of Chamaesium
frigidum (HAND.-MAZZ.) SHAN RENHWA ex
PU FATING.
We have discovered that S. frigidum is not
the first legitimate name of the species, as it
was earlier described by H. BOISSIEU (1906)
under the name of Apium ventricosum
H.BOISSIEU. The good type material kept in P
(Fig. 2), being clearly identical with later described S. frigidum. The reason to attribute the
species to Apium was probably the same as in
the case of S. frigidum – leaf blade dissection.
Apium is the genus of 15 – 20 species of different continents and its monophyly is doubtful.
Leaf dissection is more variable than in Sium,
and some species have leaves distantly resembling those of A. ventricosum. A. ventricosum
was cited later only by H. WOLFF (1927) in his
treatment of a considerable part of DRUDE’S
Umbelliferae – Ammineae (correctly, Apieae)
for ENGLER’S, Das Pflanzenreich. The type
locality of A. ventricosum (Zogalougla) is
probably in the same region of Yunnan, or in
adjacent Sichuan. Modern data on distribution
show the area of the species limited by preTibetan regions of these two provinces of SW
China.
351
So, there are three viewpoints (hypotheses)
about the generic attribution of the species –
Apium, Sium, and Chamaesium, all these genera being distant one from other. This issue
was checked using modern molecular (sequencing) investigation of internal transcribed
spacers (ITS) of nrDNA (VALIEJO-ROMAN et
al. 2002), side by side with some other critical
taxa of Chinese Umbelliferae. The results
clearly did not confirm the hypotheses on the
attribution of the species to Apium or Sium.
Unfortunately, there were no data about ITS
sequences of Chamaesium, up to now not included in molecular Umbelliferae studies. This
last genus and S. frigidum are, however, morphologically not similar, and, in particular,
considerably differ in fruit anatomy (see below). Molecular analysis has shown a new
affinity for S. frigidum, in this case with the
genus Sinocarum H.WOLFF ex SHAN RENHWA
& PU FATING. This is the fourth hypothesis
about taxonomical relation of this enigmatic
species.
Carpological and other morphological
characters to check taxonomic position of
the species
Being a part of today’s molecular-phylogenetic
investigations, the taxa remain to be understudied morphologically, especially regarding
of the carpology. The fruit TS pictures in the
FRPS are not perfect, ignoring some essential
anatomical details. Sometimes, these pictures
are simply incorrect.
We tried to check all the four proposed
hypotheses on generic position of A. ventricosum to select the most probable, i.e. that confirmed by morphological data. Among morphological characters the carefully exolored
fruit structure could be regarded as the most
informative (although not always).
The fruits of A. ventricosum (Fig. 3a) are
compressed laterally, hardly dividing to mericarps, without carpophore. Stylopods small,
short-conical, styles very short, 0.5 – 0.7 mm
long, reflexed. Mericarps elliptic, 2.2 – 2.5 mm
long, ca 0.7 mm broad, in cross-section almost
round, ribs inconspicuous, commissure narrow.
Exocarp of 1 – 2 layers of comparatively large
thin-walled cells. Mesocarp of small parenchyma cells. Secretory ducts hardly visible,
© 2003 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim
352
Fig. 1
Type specimen of Sium frigidum HAND.-MAZZ. (WU)
© 2003 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim
Feddes Repert., Berlin 114 (2003) 5 – 6
M. G. PIMENOV & E. V. KLJUYKOV: What is Sium frigidum HAND.-MAZZ.?
353
Fig. 2
Type specimen of Apium ventricosum H.BOISSIEU (P)
© 2003 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim
354
Feddes Repert., Berlin 114 (2003) 5 – 6
Fig. 3
Transections of mericarps: a — Sinocarum ventricosum (H.BOISSIEU) PIMENOV & KLJUYKOV (Apium ventricosum H.BOISSIEU), origin: China, Sichuan, between Xinduqiao and Kanding, 30°04?N, 101°46?E.
21.09.1998, PIMENOV et al., 288 (MW); b — Apium graveolens L., origin: Kirghyzia, Osch prov., near
Suzak. 28.08.1975, PIMENOV et al., 1424 (MW); c — Sium latifolium L., origin: Russia, Volgograd prov.,
Ilovlja. PIMENOV et al., s.n.; d — Chamaesium paradoxum H.WOLFF, origin: China, Sichuan, between Danba
and Renmei, 30°31?N, 101°37?E. 18.09.1998, PIMENOV et al., 244 (MW); e — Sinocarum cruciatum
H.WOLFF, origin: China, Sichuan, route Hongyuan – Barkam, valley of Somohe, 32°01?N, 102°36?E.
16.09.1998, PIMENOV et al., 188 (MW)
© 2003 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim
M. G. PIMENOV & E. V. KLJUYKOV: What is Sium frigidum HAND.-MAZZ.?
one to three in valleculas, three on commissural side. Rib secretory ducts small. Endocarp of
comparatively large thin-walled cells with
slightly lignified walls. Spermoderma of small
cells. Endosperm plane on commissural side.
The Apium-, Sium-, and Chamaesium-hypotheses could be rejected on the basis of comparative carpological analysis; Apium and
Sium, as it was noted above – also according to
ITS sequencing results.
The fruits of Apium graveolens L. (Fig. 3b),
the type species of Apium, are slightly compressed laterally, dividing to mericarps, with
entire carpophore. Calyx teeth absent. Stylopods short-conical, styles ca. 0.5 mm long,
reflexed. Mericarps elliptic in outline, pentagonal in cross-section, 1.4 – 1.5 mm long, 0.9 –
11 mm broad, with equal short-winged ribs.
Exocarp of small cells with slightly thickened
outer walls. Commissure narrow. Mesocarp of
parenchymatous cells. Vittae 1 – 3 in valleculas,
large and small, 2 – 4 (?) on commissural side.
Rib vascular bundles lengthened in radial direction. Endocarp and spermoderma of small
cells. Endosperm plane on commissural side.
The similar description can be found in DRUDE
(1897 – 1898), and KLAN (1947).
The fruits of Sium latifolium L. (Fig. 3 c),
the type species of Sium, are slightly compressed laterally, dividing to mericarps, carpophore often reduced (some other species have
well developed carpophore). Mericarps elliptic
in outline, 2.8 – 3 mm long, 1.4 – 1.5 mm broad,
in cross-section pentagonal, with five keeled to
short winged, obtuse ribs, approximately equal.
Exocarp of one layer of small cells with
slightly thickened outer walls. Commissure
narrow. Mesocarp in ribs of comparatively
large round cells with thin, slightly lignified
pitted walls. Vascular bundles are solid, situated in distal parts of ribs, being accompanied
by several small secretory ducts. Vallecular
vittae 1 – 2, commissural 2 – 4. There are additional solitary vittae in proximal part of ribs.
Endocarp and spermoderma of small cells.
Endosperm plate on commissural side. The
similar histological structure can be found in
KLAN (1947); but the general picture with six
mericarp ribs is probably of terate fruit; in text
five ribs are indicated. In general, S. latifolium
fruits have a structure, rather usual in Apieae.
One of comparatively rare features is the posi-
355
tion of vascular bundles in distal parts of the
ribs. Similar fruit structure was described by
GOROVOY (1966) for S. suave WALTON, a
species from the same type section of Sium.
The fruits of Chamaesium paradoxum
H.WOLFF, the type species of Chamaesium
H.WOLFF (Fig. 3 d) are of two dividing mericarps with 2-cleft carpophore. Mericarps 2.7 –
2.8 mm long, 1.5 – 1.8 mm broad, ovate, convex at dorsal side, with groove at commissural
side. Calyx teeth connate, forming crown. Stylopods flat, styles ca. 0.8 mm long, reflexed.
Ribs 10, visible, approximately equal, terete. In
cross-section mericarp almost round. Exocarp
of large cells, slightly elongated in radial direction. Mesocarp in ribs of small parenchymous
cells, destroyed in maturity. Vascular bundles
10 (corresponding to ribs), thin. Vallecular
vittae solitary. There are small secretory ducts
in some ribs. Endocarp of large cells. Endosperm with broad deep groove at commissural side. Other species of the genus have a
similar structure. The peculiarity of Chamaesium fruit anatomy is the double number of
mericarp vascular bundles and ribs, what is
rare in the family. Are all these ribs primary,
appearing as a result of splitting of five primary ribs (usual structure in the majority of
Apioideae), or five ribs are primary and five
are secondary? This issue could be solved in
fruit development studies. In any case, fruit
structures in Chamaesium and A. ventricosum
are considerably different.
The attribution of the species under analysis
as a member of Sinocarum is more complicated. There is no clear idea about the volume
of this genus, its specific content and diagnostic characters for delimitation from some related genera. In this situation one should check
the nomenclatural type of Sinocarum, S. cruciatum H.WOLFF, and to compare it with our
critical species.
The fruits of S. cruciatum (Fig. 3 e) are
ovate, glabrous, dividing, with bifid carpophore. Stylopods short-conical, styles up to
0.5 mm long, reflexed. Mericarps 2.0 – 2.2 mm
long, 1.1 – 1.2 mm broad, slightly compressed
dorsally, in cross-section semicircular, with
thin pericarp; ribs subinconspicuous or thin
filiform. Exocarp unilayered, of small cell,
interrupting near column. Commissure narrow.
Mesocarp of small parenchymatous cells. Vit-
© 2003 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim
356
tae three per valleculas, four to six at commissural side. Rib secretory ducts small, visible
not in each rib. Endocarp unilayered, of narrow
cells, lengthened in tangential direction, with
slightly lignified walls. Spermoderma of comparatively large cells. Endosperm on commissural side with not deep double groove.
Of course, the fruit of A. ventricosum are
considerably more similar to the fruits of Sinocarum, than to those of Sium, Chamaesium, or
Apium. There are, however, some differences.
The fruits of A. ventricosum are compressed
laterally and have no carpophore (the last character can be an adaptation against a long dispersal of seeds). In S. cruciatum carpophore is
developed normally, and mericarp are compressed dorsally, although not strongly. There
are some particular traits in habit and vegetative part of A. ventricosum plants, which do not
correspond completely to those of Sinocarum.
Therefore, in Sinocarum the species occupies a
more or less isolated position. As the relationships the species within Sinocarum and with
presumably related taxa are obscure, we believe it would be best to separate the regarded
species into a monotypic section of Sinocarum.
Feddes Repert., Berlin 114 (2003) 5 – 6
Type:
China
“Zogalougla,
CH.I.A.BLANCHE,” (holo – P!)
marecages.
= Sium frigidum HAND.-MAZZ., 1933, Symb. sin. 7,
3: 719, tab. 11, fig. 2 – 4; WU ZHENGYI (ed.),
1984, Index Fl. Yunnan. 1: 928; PU FATING et al.,
1992, in Fl. Reip. Pop. Sin. 55, 3: 249, tab. 109,
fig. 5 – 9.
) Chamaesium frigidum (HAND.-MAZZ.) SHANRENHWA ex PU FATING, 1993, in WANG WENTSAI
(ed.), Vasc. Pl. Hengduan Mts.: 1290.
T y p e : China “Prov. Yunnan bor.-occid.: supra
vicum Anangu austro-orientalem pagi Dschundien
(“Chungtien”), in regione frigide temperatae limo
turfi Diolo. 16.08.1915. H. F. HANDEL-MAZZETTI,
7684” (holo – WU!; iso-US, PE – photo!)
D i s t r i b u t i o n : China (SW: Sichuan, Yunnan).
Acknowledgements
This work was supported by grants from the
Russian Foundation for Fundamental Investigations (RFFI) and the National Geographic
Society (USA). The curators of KUN, P, PE,
and WU are gratefully acknowledged for the
study facilities to examine collections. The
authors thank our French colleagues, M. JeanPierre R e d u r o n and Dr. Alain C h a n g y , for
preparation of electronic picture of Apium ventricosum.
Taxonomic implications
References
Sinocarum H.WOLFF ex SHAN RENHWA &
PU FATING sect. Apiopsis PIMENOV &
KLJUYKOV sect. nov.
Ab omnibus speciebus generis Sinocarum bene
differt fructibus lateraliter compressis, carpophoris expertibus, vittis mericarpiorum indistincte coherentibus, foliis pinnatis et partis
hypogaeeis radicibus tuberiforme incrassatis et
rhizomatis verticalibus nodis cum radiis adventivis prebentibus.
T y p e : S. ventricosum (H.BOISSIEU) PIMENOV &
KLJUYKOV (Apium ventricosum H.BOISSIEU)
Sinocarum ventricosum (H.BOISSIEU)
PIMENOV & KLJUYKOV comb. nova
) Apium ventricosum H.BOISSIEU, 1906, Bull. Soc.
Bot. Fr. 53: 425; H. WOLFF, 1927, in ENGLER
Pflanzenr. 90 (IV, 228): 46.
© 2003 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim
BOISSIEU, H. 1906: Note sur quelques Ombellifères
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genuinae: 1 – 398. – In: A. ENGLER (ed.), Das
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357
Address of the authors:
Professor Dr. M. G. P i m e n o v , Dr. E. V. K lju y k o v , Moscow State University M. V. Lomonossov,
Botanical Garden, R-119899 Moscow, Russia.
e-mail [email protected]
Manuscript received: March 15th, 2003.
© 2003 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim