2
AROIDEANA, Vol. 29
A Taxonomic Revision of Biarum (Araceae)
Peter C. Boyce
Malesiana Tropicals
Suite 4, Level 9, Tun Jugah Tower
No. 18 Jalan Abdul Rahman
93000 Kuching, Malaysia
ABSTRACT
A taxonomic revision of Biarum is presented. Hitherto species-ranked B. galianii is
reduced to a subspecies of B. tenuifolium.
Hitherto subspecific (to B. davisii) B. marmarisense is raised to full species rank.
The names Biarum abbreviatum, B. aleppicum, B. anguillare, B. arundanum, B.
bovei ssp. disparvar. purpureum, B. bovei
ssp. dispar var. zanonii, B. cupanianum,
B. platyspathum, B. spruneri, B. straussi,
B. tenuifolium var. latifolium, B. zelebori,
Cyllenium carduchorum, Ischarum crispulum, 1. jraasianum, 1. kotschyi and 1.
nobile are lectotypified. Ischarum pyrami
is epitypified. A key to all taxa is provided.
the Middle East, where 75% of the species
occur as endemics.
Biarum species have a strongly seasonal growth regime; the plants beginning
growth in late summer or early autumn
with the onset of winter rains and continuing into late spring when the plants become dormant at the start of summer heat
and drought. The majority of species blossom in autumn and early winter and this,
together with the often striking appearance of the inflorescence, had resulted in
a growing popularity of Biarum species
amongst alpine garden enthusiasts. Further aspects of this horticultural popularity
have been covered by Mayo (1983) and
Mathew (1987).
KEYWORDS
Biarum aleppicum, Biarum angustatum, Biarum auraniticum, Biarum bovei,
Biarum carduchorum, Biarum carratracense, Biarum crispulum, Biarum davisii,
Biarum dispar, Biarum ditschianum, Biarum eximium, Biarum jraasianum, Biarum kotschyi, Biarum marmarisense, Biarum mendax, Biarum olivieri, Biarum
pyrami, Biarum rhopalospadix, Biarum
straussii, Biarum syriacum, Biarum tenuifolium ssp. abbreviatum, Biarum tenuifolium ssp. arundanum, Biarum tenuifolium ssp. galianii, Biarum tenuifolium ssp. idomenaeum, Biarum tenuifolium ssp. tenuifolium, Biarum tenuifolium
ssp. zeleborii.
INTRODUCTION
Biarum comprises 21 species of dwarf
tuberous-stemmed herbs occurring in
semi-arid and seasonally dry areas of
southern Europe, North Africa, the Near
and Middle East. The centre of diversity is
HISTORY
Until the publication of Schott's articles
in the Wiener Zeitschrijt jur Kunst, Literatur, Theater und Mode (1829a, 1829b,
1829c, 1829d, 182ge, 1829f, 1829g, 1830a,
1830b, 1830c, 1830d, 1830e,) and his aroid
account in Meletamata Botanica (1832),
the genus Arum was ill-defined, containing numerous species bearing only superficial similarity to the type of the genus, A.
maculatum 1. Schott (1832) attempted to
bring a degree of homogeneity to Arum
by segregating new genera to account for
the anomalous taxa, proposing the genus
Biarum to account for two species of
Arum (sensu Linnaeus) with uniovulate
ovaries and a basal placenta, loosely arranged staminate flowers with anthers dehiscing by ventral longitudinal slits, scattered staminodes and pistillodes and connate spathe tube margins. The two species
included were: B. gramineum (Lam.)
Schott and B. tenuifolium (1.) Schott.
3
PETER C. BOYCE, 2006
Blume (1836) retained Schott's two species and described an additional three, B.
bovei, B. homeid and B. olivieri. Blume divided Biarum into two sections, placing
Schott's species in "Sectio 1" (i.e. sect. Biarum) and creating sect. Ischarum for the
three new ones. Blume distinguished sect.
Ischarum by the lack of staminodes, anther thecae dehiscing by apical pores and
more-or-Iess elongated style. By contrast
in sect. Biarum sensu Blume, both staminodes and pistillodes are present, the thecae dehisce by longitudinal slits and the
stigma is sessile. Blume emphasized the
different geographical distribution of the
sections as then known, with sect. Ischarum in the eastern Mediterranean region
and sect. Biarum in the western Mediterranean.
Boissier (1853) added two new species,
B. spruneri and B. alexandrinum, but did
not review any of the previous treatments.
He placed B. spruneri in sect. Biarum (as
sect. 'Eubiarum') with the note that B.
spruneri was apparently intermediate between sect. Biarum and sect. Ischarum.
Biarum alexandrinum was assigned to
sect. Ischarum.
Schott and Kotschy (1854) raised Blume's
sect. Ischarum to generic status with a single new species, I. eximium. No mention
was made, however, of the three species
previously described by Blume for the
sect. Ischarum and Schott (1856) eventually made the necessary new combinations
in the genus Ischarum.
In his Synopsis Aroidearum, Schott
(1856) published seven new combinations
and two new species for Ischarum. The
new combinations included the two taxa
described by Boissier (1853), Arum haenseleri published by Willkomm (1847) and
Biarum lehmannii Bunge (1851). The last
mentioned was later transferred to Eminium by Kuntze (1891). The new species
were I. kotschyi and I. dispar.
With the publication of Genera Aroidearum (1858) Schott proposed two new
genera, Cyllenium and Leptopetion, for B.
spruneri and B. alexandrinum respectively. The differences between the genera
concerned the presence or not of a style,
the means of thecae dehiscence, i.e. slits
as opposed to pores, and the shape and
distribution of the pistillodes.
Engler (1879) adopted what was essentially Schott's system except that he reduced all Schott's segregate genera to subgenera of Biarum and dispensed with Leptopetion altogether, referring it to subgen.
Ischarum. Engler also reduced many of
Schott's species to subspecific or varietal
status or to synonymy.
Boissier (1882) also chose not to recognize Schott's separate genera, and in
fact went a stage further than Engler
(1879) in distinguishing them at the rank
of section rather than subgenus. Boissier
followed Engler in not accepting Leptopetion at any rank, also referring it to section
Ischarum.
Engler's (1920) revision of Biarum for
Das Pjlanzenreich was the last comprehensive treatment of the genus. Although
little had changed since his 1879 classification, he published one new species, B.
straussii, and a number of subspecific
taxa.
Since the Pjlanzenreich account several
floristic examinations of Biarum have
been undertaken by various authors (e.g.
Riedl, 1963, 1985; Mouterde, 1966; Talavera, 1976; Mill, 1984; Koach & Feinbrun,
1986). Riedl (1980b) published a preliminary summary of the genus together with
a key to the species. However, none of
these accounts attempts a comprehensive
treatment and new discoveries and interpretations during the last fifty years have
made necessary this revision.
TYPIFICATION
During the course of this revision it has
been necessary to lectotypify a number of
names. The majority of these are names
published by Schott for which the types
were destroyed during the closing stages
of WW2. For lectotypes I have followed
the advice of Riedl & Riedl-Dorn (988) in
selecting illustrations that were commissioned by Schott and prepared from living
and herbarium specimens. These are
known collectively as the !cones & Reliq-
4
AROIDEANA, Vol. 29
uiae Aroideamm. It has also been necessary to lectotypify names based on types
destroyed in Berlin. In these cases it has
not been possible to trace any authentic
material, but in some instances drawings
of the types exist and have been chosen
to serve as the lectotype.
INFRAGENERIC CLASSIFICATION
I. Subgenus Biarum
[Biamm Sectio 1 Blume, Rumphia 1:112
(1836)]
[Biamm Subgen. Eubiamm Engler in A. &
C. DC., Monog. Phanerog. 2: 572
(1879) & in Engler, Das Pflanzenr. 73
(IV.23F): 134 (1920)]
[Biamm sect. Eubiamm (Engler) Boiss.,
Fl. Or. 5:31 (1882)]
Biamm subgenus Cyllenium (Schott) Engler in A. & c. DC., Monog. Phanerog.
2:574 (1879); Engler in Engler, Das
Pflanzenr. 73 (IV.23F):136 (1920).
Type: B. spmneri Boiss.
Cyllenium Schott, Gen. Aroid. t.9 (1858).
Type: C. spmneri (Boiss.) Schott
Biamm sect. Cyllenium (Schott) Boiss., Fl.
Or. 5:32 (1882).
Anthers with thecae dehiscing by ventral, longitudinal slits, connective beak-like
and extending beyond the anther surface.
Staminodes hooked, rarely peg-like or filamentous, mostly present above and below male flower zone, rarely present only
below the male flower zone and then
hooked.
Species:
1a.
lb.
Ie.
1d.
Ie.
If.
tenuifolium
tenuifolium
tenuifolium
tenuifolium
tenuifolium
tenuifolium
ssp.
ssp.
ssp.
ssp.
ssp.
ssp.
tenuifolium
zelebori
arundanum
galianii
abbreviatum
idomenaeum
Taxa of B. tenuf(olium are arranged by
relative similarity and thus supposed relationship.
2.
rhopalospadix
II. Subgenus Ischarum (Blume) Engler
in A. & c. DC. Monog. Phanerog. 2:
575 (1879). Type: I. bovei (Blume)
Schott (See Nicolson 1967).
Ischamm (Blume) Reichb., Deutsche Bot.
Bd.1 Herbuch Nom. 32 (1841) &
Schott & Kotschy, Oesterr. Bot. Wochen. 4: 81 (1854).
Biamm section Ischamm Blume, Rumphia 1: 112 (1836), Boiss., FI. Or. 5: 32
(1882).
Anthers with thecae dehiscing by apical
pores, connective barely prominent or
flush with the anther surface. Staminodes
absent above male flower zone.
Species:
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.
21.
aleppicum
angustatum
earduehorum
eximium
bovei
erispulum
dispar
olivieri
straussii
syriaeum
earratracense
kotsehyi
fraasianum
pyrami
mendax
auraniticum
ditsehianum
davisii
marmarisense
Biarum Schott nom. cons. in Schott & Endlicher, Melet. Bot. 17 (1832) & Syn.
Arokl. 6 (1856) & Gen. Aroid. t.7
(1858) & Prodr. Syst. Aroid. 60-64
(1860); Pfeiffer, Nomen. Bot. 1(1):
403-404 (1873); Engler in A. & c. DC.,
Monog. Phanerog. 2: 571-580 (1879)
& in Engler & Prantl., Die Natlirl.
Pflanzenfam. 149 (1889) & in Engler,
Pflanzenr. 73(IY.23F): 132-143 (1920);
Riedl in Aroideana 3(1): 24-31 (1980);
Mayo, Bogner & Boyce, The Genera
of Araceae, 266-268, PI.96 (1997).
Type: B. tenuifolium (1.) Schott. Homaid Adans., Fam. PI. 2:470 (1763) in
PETER C. BOYCE, 2006
syn. nom. rejic.; Pfeiffer, Nomen. Bot.
1(2): 1658 (1874). Type: H. tenuifolium (L.) Adans.
Ischarum Schott & Kotschy, Oesterr. Bot.
Wochen. 4: 81 (1854); Schott, Syn. Aroid. 6-8 (1856) & Gen. Aroid. t.1O
(1858) & Prod. Syst. Aroid. 65-70
(1860); Pfeiffer, Nomen. Bot. 1(2):
1764 (1874). Type: 1. bovei (Blume)
Schott (See Nicolson 1967).
Leptopetion Schott, Gen. Aroid. t.8 (1858)
& Prodr. Syst. Aroid. 64 (1860). Type:
L. alexandrinum (Boiss.) Schott.
Cyllenium Schott, Gen. Aroid. t.9 (1858) &
Prodr. Syst. Aroid. 64-65 (1860).
Type: C. spruneri (Boiss.) Schott.
Stenurus Salis., Gen. PI. 5 (1866). Type: S.
tenuifolium (L.) Salis.
Homaida Adans. emend O. Kuntze, Rev.
Gen. PI. 2: 742 (1891). Type: H. tenuifolium (L.) Adans. emend O. Kuntze.
Dwarf tuberous stemmed herbs. Tuber
dorso-ventrally compressed-discoid to ±
globose, encased basally by remains of the
previous season's tuber, growth point central, peripheral adventitious buds none to
many, usually giving rise to independent
tubers, tuber apex coated with moderate
to copious amounts of waxy farina and
bearing the remains of the previous season's cataphylls; roots simple, emitted in a
ring around the growth point, contractile
or feeding; contractile roots fusiform, usually thick, feeding roots slender, cylindrical. Foliage hysteranthous (emerging after flowering), occasionally synanthous
(emerging with inflorescence), rarely proteranthous (emerging before flowering),
petiolate, petiole sometimes ± subterranean, leaves erect or, rarely, reflexed; petioles enclosed proximally by several cataphylls, the inner papery and the outer fibrous, these often emerging above ground
and encasing the basal part of the aerial
shoot, petioles terete distally, canaliculate
and expanded into a membranous wing
proximally, petioles enclosing one another, forming a weakly defined aerial pseudostem in some species; leaf lamina linear,
lanceolate, elliptic-oblong or spathulate,
5
decurrent, ± cuneate, rounded or ± truncate apically, acute to obtuse basally, margins smooth to undulate, rarely crispulate,
mid-, light, bright or deep green, rarely
with green or black-purple bullae or silver-grey spotting. Inflorescence ± sessile
on the tuber or borne on a short to rather
long, subterranean peduncle, rarely peduncle emerging above ground, smelling
foetid or sweet. Spathe divisible into a tubular lower portion (spathe tube) and a ±
expanded upper portion (spathe limb);
spathe tube narrow to inflated, sometimes
greatly so, constricted distally or not,
emergent to partially buried, rarely entirely
subterranean, margins partially to fully
connate, sometimes convolute distally, exterior deep purple to dirty green or greenish purple above, ± white below, interior
off-white below, purple above, or wholly
purple or off-white with purple staining at
the base around the pistillate flowers;
spathe limb large to small, rarely ± absent,
linear, lanceolate or elliptic, erect, reflexed
or deflexed, flat to recurved and coiled or
strongly involute; exterior dirty green,
more rarely mid-green, dirty white, dull
yellow or pinkish brown sometimes with
purple spotting and staining; interior deep
purple-brown, yellow or pale greenish,
sometimes purple with a green apex, apex
subacute, acute to acuminate, margins
smooth to undulate, rarely crispulate. Spadix shorter than, equalling or exceeding
the spathe limb, divided into distinct
zones; a sterile terminal appendix, a zone
of staminodes (subgenus Biarum), a fertile
male zone, a stamino-pistillate interstice, a
further zone of staminodes (sometimes absent) and a fertile female zone; terminal
appendix sessile to stipitate, cylindrical to
fusiform, erect to flexuous, apex acute to
rounded, tapering, base rarely rounded or
sub-truncate, smooth, very rarely furnished with filamentous processes proximally, deep purple, brown-red or brown,
occasionally greenish, rarely dirty yellow.
Flowers: supra-stamina I staminodes present only in subgen. Biarum, arranged in
few to several irregular whorls, simple or
1-2-branched, hooked, peg-like or filamentous, partially expanded proximally,
6
AROIDEANA, Vol. 29
glossy, cream; staminate flowers arranged
in a cylindrical, globose or oblong zone,
each comprising two anthers, connective
short to :!: absent, rarely prolonged into a
:!: prominent beak, thecae dehiscing by
coalesced or separate apical pores Csubgen. Isbarum) or ventral longitudinal slits
Csubgen. Biarum), cream to purple; pollen
free or in strands, exine spinose or
smooth, interfloral staminodes usually
confined to the base of the stamino-pistillate interstice, more rarely arranged evenly
over the whole area or absent, slender-filiform to hooked, simple to 1-3-branched,
recurved, decurved or tangled, cream, offwhite or purple; pistillate flowers arranged
in a hemispherical cluster at the base of
the spadix, ovary oblong, sub-globose or
bottle-shaped, off-white to purple, uniovulate with a basal placenta, ovules orthotropous, style slender to rather stout or
absent, stigma capitate, pale greyish or
purple. Infructescence subterranean or
partially emergent, globose berries many,
globose to pyriform, white to lilac- or purple-stained, stigmatic remnants slightly
prominent or not; seed ovoid to globose,
large to small with a large elaiosome at the
hilum, testa leathery, :!: smooth to reticulate, pale to dark brown, endosperm copious, embryo straight; eophyll Ian ceolate
to spathulate.
Twenty-one species of dwarf tuberousstemmed strongly seasonal herbs occurring in semi-arid and seasonally dry areas
of southern Europe, North Africa, the Near
and Middle East.
KEY TO SPECmS
1. Staminodes occurring above and below
the staminate flower zone ........ .
. . . . . . . . . . . . . . . . . .. 1. tenuifolium
Staminodes absent above the staminate
flower zone; present or absent below
2
2. Staminodes hooked, stiff, thickened Thecae dehiscing by ventral longitudinal
slits, connective rostrate ......... .
................. 2. rhopalospadix
Staminodes filiform, flexuous, or absent,
never hooked and thickened Thecae dehiscing by apical pores, connective barely prominent or flush with the anther surface. . . . . . . . . . . . . . . . . . . . . . . . .. 3
3. Spadix appendix massively thickened
with reflexed basal 'hairs'; spathe limb
much reduced,. recurved ......... .
· . . . . . . . . . . . . . . .. 19. ditschianum
Spadix appendix with no ornamentation;
spathe limb well developed or, if reduced, then erect, cucullate ........ 4
4. Spathe limb usually much shorter than
the spathe tube, margins recurved,
spathe tube enclosing much of the spadix .......................... 5
Spathe limb exceeding the spathe tube in
length, flat or with the margins inrolled,
spathe tube enclOSing the base of the
spadix only .................... 6
5. Spathe 5-6 cm long, spadix appendix 33.5 cm long X 2 mm thick .. 20. davisii
Spathe 7-8 cm long, spadix appendix
3.5-5 cm long X 0.5 mm thick .....
· ............... 21. marmarisense
6. Spathe tube not inflated, the sides ± parallel . . . . . . . . . . . . . . . . . . . . . . . . .. 7
Spathe tube inflated, the sides ± gibbous
............................. 9
7. Spathe tube margins free ± to the base;
stigmas not borne on a stipe ..... .
· . . . . . . . . . . . . . . . . . .. 3. aleppicum
Spathe tube margins free for a quarter of
their length; stigmas borne on a short to
moderately long stipe ............ 8
8. Staminodes directed downwards; leaves
narrowly lanceolate-elliptic ....... .
· . . . . . . . . . . . . . . . . .. 4. angustatum
Staminodes directed upwards; leaves elliptic to broadly oblong-elliptic .....
· ................. 5. carduchorum
9. Spathe tube distinctly wider than the
spathe limb, the margins joined for their
entire length; spathe limb appearing linear due to inrolled margins; spadix appendix filiform . . . . . . . . .. 10. olivieri
Spathe tube as wide as or narrower than
the spathe limb, the margins free for at
least a quarter of their length ....... 10
10. Foliage proteranthous (emerging before
flowering) ..................... 11
Foliage hysteranthous (emerging after
flowering) or synanthous (emerging with
inflorescence . . . . . . . . . . . . . . . . . . . 12
11. Leaf lamina ovate-elliptic, oblong or linear; staminodes restricted to the bottom
quarter to third of the interstice ....
· . . . . . . . . . . . . . . . . . . .. 11. straussii
Leaf lamina in mature plants linear to linear-elliptic; staminodes distributed over
the basal half of the interstice (the axis
between the male and female flower
zones) . . . . . . . . . . . . . . . 12. syriacum
12. Staminodes distributed evenly over the
interstice separating the staminate and
pistillate flower zones .... 6. eximium
Staminodes arranged mostly above the
pistillate flower zone, intersticial stami-
PETER C. BOYCE, 2006
13.
14.
15.
16.
17.
18.
19.
20.
nodes adjacent to the staminate flower
zone much reduced or vestigial, or staminodes absent .................. 13
Spathe limb interior greenish white, spa18. auraniticum
dix appendix yellow
Spathe limb interior deep purple-brown;
spadix appendix similarly coloured ... 14
Spathe tube margins free for quarter to
half their length ................. 15
Spathe tube margins free for three-quarters their length ................. 16
Spathe tube margins free for half their
length, staminodes few, SW Spain ....
. . . . . . . . . . . . . . . . . 13. carratracense
Spathe tube margins free for a quarter of
their length, S Greece ........... .
. . . . . . . . . . . . . . . . .. 15. fraasianum
Spadix appendix ca. 2-4 mm in diam.,
slender-cylindric, to slender-fusiform, appearing ± consistent diameter ...... 17
Spadix appendix more than 4 mm in
diam., fusiform, widest below the middle
............................. 19
Spathe tube globose; interstice twice as
long as the staminate flower zone; staminodes very few or absent ... 9. dispar
Spathe tube oblong; interstice equalling
the staminate flower zone; staminodes
many ......................... 18
Foliage hysteranthous; spathe limb lanceolate, margins smooth ...... 7. bovel
Foliage synanthous; spathe limb linearlanceolate, margins crispulate ..... .
.. .. . . . . .. .. . . .. .. .. 8. crispulum
Spathe tube oblong, spathe limb narrowly lanceolate, acute, seldom exceeding
10 cm ............... 14. kotschyi
Spathe tube globose, spathe limb lanceolate, acuminate, exceeding 12 cm ... 20
Spathe tube margins fully fused; spathe
limb remaining erect during flowering
..................... 17. mendax
Spathe tube margins for 1A of their length;
spathe limb reflexing and curling during
flowering . . . . . . . . . . . . .. 16. pyrami
1. Biarum tenuifolium (L.) Schott in
Schott & EndI., Melet. Bot. 17 (1832)
& Syn. Aroid. 6 (1856) & Prodr. Syst.
Aroid. 60 (1860); Boiss., FI. Or. 5:31
(1882); Engler in A. & c. DC., Monog.
Phanerog. 2:573 (1879) & in Engler,
Das Pflanzenr. 73 (IV. 23F): 134
(920); Polunin, FI. Europe t.183
nO.1820 (1969); Riedl in Aroideana
3(1): 26 (1980).
Arum tenuifolium L., Sp. PI. ed.1 967
(1763). Type: Konig 77 (syntypes:
Herb. Linn. 1079.13, 1079.14).
Arum gramineum Lam., Encyc. 3:10
7
(1789). Type: no data (P). Biarum
gramineum (Lam.) Schott in Schott &
EndI., Melet. Bot. 17 (1832).
Biarum constrictum C.Koch, Ind. Sem.
Hort. BeroI. App. 2 (1853). Type: ITALY. Plantam in regno Neapolitano
sponte crescentum ex horto Societ.
reg. bot. Ratisbonensis ante decenium
accepimus. In horto sub diu cultum
sero auturnno floret. (holotype B destroyed; isotype K (tracing of Koch's
drawing of the type). B. spruneri
Schott, Gen. Aroid. t.7 (1858) non
Boiss. (1853). Type: Greece, Spruner
s.n. (holotype B).
Biarum anguillare Schott, Prodr. Syst. Aroid. 62 (1860). Type: YUGOSLAVIA,
Dalmatia (holotype W destroyed; lectotype W chosen here (Schott's Reliquiae Aroideae no. 360)). This pencil
illustration is chosen in preference to
the coloured illustrations (Schott's
/cones Aroideae nos. 1493, 1494) and
other pencil illustrations (leones 1495,
1496 & 1497) since it combines inflorescence, infructescence and foliage
in one plate.
Biarum tenuifolium (L.) Schott var. typicum Engler in Engler, Das Pflanzenr.
73 (IV. 23F): 134 (1920). Type: as for
B. tenuifolium (L.) Schott.
Biarum tenuifolium (L.) Schott var. typicum Engler subvar. constrictum (C.
Koch) Engler in Engler, Das Pflanzenr. 73 (IV. 23F): 136 (1920).
Tuber dorso-ventrally compressed discoid, offsetting freely, 2-6 cm X 1.5-2.5
em, mid-brown. Leaves 4-20, hysteranthous, distinctly to rather obscurely long
petiolate, bases encased by 3-many, 2-8
X 0.5-2 cm oblong-Ianceolate sub-fleshy,
later papery, cataphylls, these pale green
drying off-white to pale straw-yellow; petiole 1-8 cm X 2-5 mm, adaxial surface
channeled distally, expanded proximally
into a membranous wing, mid-green; leaf
lamina oblong-Ianceolate, linear-Ianceolate, spathulate or linear-oblong, 2.5-49
cm X 2-21 mm, apex acute to obtuse or
rounded, base long decurrent to cuneate,
margins smooth, gently undulate or
8
AROIDEANA, Vol. 29
strongly undulate-crispulate, ca. 5-9 primary lateral veins per side, mid- to dark
green. Inflorescence appearing in late
summer to mid-autumn, occasionally in
spring, usually strongly foetid of cattle
dung, Italian populations of ssp. abbreviatum reported to smell goat-like (Paglia,
1909); peduncle 6-10 cm X 3-5 mm,
white, clothed with few to several oblonglanceolate sub-fleshy, later papery, cataphylls, 4-12 X 1.5-2 cm, these pale green
drying off-white to pale straw-yellow.
Spathe 3-30 cm long; spathe limb 2-25 X
0.5-6 cm, apex acute to acuminate exterior
mid-green heavily stained deep brownpurple, sometimes paler to green apically,
interior deep brown-purple, green towards the apex in ssp. abbreviatum; lower
spathe cylindric to oblong cylindric, usually constricted above the pistillate flowers, sometimes further constricted ca.
along its length, margins fused for their
whole length, 2-6 X 1-1.25 cm, exterior
white, stained purple towards the apex, interior white. Spadix shorter than to greatly exceeding the spathe limb, 4-40 cm
long; spadix appendix cylindrical to stoutly fusiform, 3-41 cm X 1.5-9 mm, deep
brown-purple, often somewhat paler than
the spathe limb, rarely olive green or dirty
yellow; upper staminodes in a zone 317 mm long, in 2-10 regular to rather irregular whorls, hooked, peg-like or filamentous, cream to ivory; staminate flowers in a zone 3-20 X 1-6 mm diam.,
cream to ivory; interstice usually :!:: absent, occasionally up to 15 mm long above
the pistillate flowers or below the staminate flowers, cream; lower staminodes
in a zone 2-23 X 1-4 mm, in 2-13 regular
whorls, hooked, peg-like or almost filamentous, cream to ivory; pistillate flowers in a hemispherical to slightly cylindrichemispherical zone 3-7 X 2-5 mm; ovary
oblong, 0.5-2 X 0.25-2 mm, cream; stigma
sessile, capitate, 0.25-0.33 mm in diam.,
cream. Infructescence globose, 1.5-4 cm
in diam., consisting of 15-45 berries; berry
oblong to oblong-globose, 3.5-9 X 2.5-5.5
mm, white when ripe; seed ovoid, 2.5-4.5
X 3-6 mm, pale brown, barely reticulate.
*
KEY TO THE SUBSPECIES OF BIARUM
TENUIFOLIUM
1. Staminodes hooked ............. "
2.
3.
4.
5.
2
Staminodes not hooked . . . . . . . . . . .. 5
Staminodes simple, very rarely branched
............................... 3
Staminodes always 2-3-branched ..... 4
Leaf lamina 15-25 cm X 11-15 mm; spadix appendix 10--41 cm X 2-3 mm; leaves
in mature individuals oblong-lanceolate
early in the season, linear-lanceolate later
in the season . . . .. la. ssp. tenuifolium
Leaf lamina 20--40 cm X 16-21 mm; spadix appendix 10-12 X 4-9 mm; leaves in
mature individuals always oblong-lanceolate . . . . . . . . . . . . . .. lb. ssp. zelebori
Leaf lamina oblanceolate to linear-lanceolate; staminodes 2-branched, arranged in
ca. 8 regular whorls. Plants of limestonederived red heavy clay soils ....... .
· . . . . . . . . . . . . .. Ie. ssp. arundanum
Leaf lamina linear; staminodes 2-3branched, arranged in ca. 7 irregular
whorls. Plants of loose sandy soils .....
· . . . . . . . . . . . . . . . . . . Id. ssp. gaIianii
Staminodes peg-like; leaf lamina spathulate, erect, margins gently undulate ....
· . . . . . . . . . . . . . . Ie. ssp. abbreviatum
Staminodes filamentous; leaf lamina linear-oblong, usually adpressed to the
ground, margins strongly undulate-crispulate ......... If. ssp. idomenaeum
a. ssp. tenuifolium
Leaves in mature individuals oblonglanceolate early in the season, linear-Ianceolate later in the season, lamina 15-25
cm X 11-15 mm,. Spadix appendix 1041 cm X 2-3 mm. Staminodes hooked,
simple. 2n = 26 (Marchant 1972), 16, 20,
26 (Monti & Gabari 1974).
Distribution-S Italy, Sicily, Yugoslavia,
Serbia, Bosnia-Hercegovina, FYRO Macedonia, Albania, Greece.
Ecology-Limestone-derived red clays
in garigue, open maquis, grazed hillsides,
olive groves, shallow-ploughed fields,
along margins of deep-ploughed fields.
Alt. 25-1220 m.
Etymology-From the Latin tenuo (slender) and folium (leaf), referring to the
slender foliage.
b. ssp. arundanum (Boiss. & Reuter)
Nyman, Consp. Fl. Europ. 755 (1882).
PETER C. BOYCE, 2006
Biarum arundanum Boiss. & Reuter,
Pug. PI. Nov. Afr. Bor. 110 (1852); Talavera in Lagascalia 6(2): 586-8, t.1,
A,Al (1976) & Talavera, Valdes & Galiano, Fl. Vasco Andal. Occ. 3: 209
(1987). Type: SPAIN, Grazelema, June
1849, Boissier & Reuter s.n. (lectotype
G-BOIS! chosen here). Boissier and
Reuter (loc. cit.) cite two specimens
in the protologue, the other being
'circa Ronda, Reuters.n.'. I have been
unable to locate the whereabouts of
Reuter's herbarium and, in the absence of the second specimen, hereby select the Boissier & Reuter specimen in the Boissier Herbarium, Geneva to serve as the lectotype.
Biarum bovei Blume ssp. dispar (Schott)
Engler var. discolor Maire in M.e. 640
(1930). Type: not cited.
Leqf lamina oblanceolate to linear-Ianceolate. Staminodes hooked, 2-branched,
arranged in ca. 8 regular whorls. Plants of
heavy red clay soils. 2n = 22 (Marchant
1972 as B. carratracense; Talavera 1976;
Elena & Gallego 1984).
Distribution-SW Spain, Gibraltar, S
Portugal, N Morocco.
Ecology-Restricted to red clay soils derived from the decomposition of limestone, usually in open situations, particularly along field margins or in long-fallow
fields. Alt. 50-1200 m.
Etymology-The specific epithet is derived from the Roman name for Ronda, a
town in southwestern Spain and the type
locality of this species.
c. ssp. galianii (Talavera) P.C. Boyce
comb. et stat. nov.
Biarum galianii Talavera in Lagascalia
6(2): 289, t.1 B, Bl (1976); Talavera,
Valdes & Galiano, Fl. Vasco Andal.
Occ. 3:209 (1987). Type: SPAIN, Huelva, entre San Bartolome de la Torre y
Alosno, 1 June 1976, Talavera s.n.
(holotype SEV 24330).
Leaf lamina linear. Staminodes 2-3branched, arranged in ca. 7 irregular
9
whorls. Plants of loose sandy soils. 2n =
26 (Talavera 1976; Elena & Gallego 1984).
Distribution-Badajoz and Huelva districts, SW Spain.
Ecology-Biarum galianii occurs in
loose sandy soils on open hill slopes, alt.
ca.600m.
Etymology-Named for the Spanish
botanist Emilio Fernandez Galiano.
d. ssp. zelebori (Schott) P.e. Boyce in R.
Govaerts & D.G. Frodin, World
Checklist Bibliogr. Araceae 245
(2002). Type: TURKEY, prope Smyrnam (Izmir). Zelebor s.n. (holotype W
destroyed; lectotype W (chosen here
(Schott's leones Aroideae no. 1532)).
The plate chosen as the lectotype is
annotated Zelebor 56. The plates present in Vienna are /cones nos 15281534, Reliquiae no 362 and an unnumbered /cone depicting germination and subsequent development of
the seedlings. Biarum zelebori Schott
in Oesterr. Bot. Wochenbl. 7:245
(1857). Biarum tenuifolium (1.)
Schott var. zelebori (Schott) Engler in
A. & e. DC., Monog. Phanerog. 2: 574
(1879) ["zeleborin & in Engler, Das
Pflanzenr. 73 (IY.23F): 136 (1920)
["zeleborif'l; Mill, FI of Turkey. 8:56
t.2 no.12 (1984) ["zeleborif'l.
Leaves in mature individuals always
oblong-Ianceolate, lamina 20-40 cm X 1621 mm. Spadix appendiX 10-12 X 4-9
mm. 2n not recorded.
Distribution-Crete, Rhodes, Cos, SW
Turkey (provinces of Aydin, Izmir and
Mugla).
Ecology-As for the typical variety. Alt.
300-1350 m.
Etymology-Named for the collector of
the type material.
e. ssp. abbreviatum (Schott) K. Richt.,
PI. Eur. 1: 174 (1890). Biarum abbreviatum Schott, Prodr. Syst. Aroid.
62 (1860). Type: GREECE, Heldreich
AROIDEANA, Vol. 29
10
s.n. (holotype B destroyed; lectotype
W chosen here (Schott's leones Aroideae no. 1491».
The plate selected is the most complete present in Vienna. Other plates
present are leones no. 1492, Reliquiae
no. 361 and an un-numbered leone
depicting germinating seeds.
[Arum eupanianum Guss., Fl. Sic. Syn.
2(2): 598 (1844) nom. nud.]
[Biarum tenuifolium (1.) Schott var. eupanianum Nicotra in Malpighia 22:
541 (1908), nom. nud. ['eupaniana']]
Biarum eupanianum Guss. ex Paglia in
Riv. Ital. Sci. Nat. 29: 24 (1909). Type:
Ie. 284 in Barralier, Plantae Galliam,
Hispaniam et ltaliam observertae
(1714) (lectotype chosen here).
Gussone (1844) cites the illustrations in Clusius' Rariorum Plantarum
Historia (1601), an unspecified volume by Dodoens, and Barralier's
Plantae Galliam, Hispaniam et ltaliam observertae (1714). Of the available plates, no 284 in Barralier's
work, with the caption Arum angustifolium maii, is the best candidate as
the lectotype since it closely matches
the spring flowering Biarum present
in southern Italy. Although it is not
stated that the plate was drawn from
Italian material I feel that it can be assumed that this is the case since Biarum is absent from France and there
are no spring blossoming Biarum
species in Spain.
Biarum tenuifolium (1.) Schott var. abbreviatum (Schott) Engler in A. & c.
DC., Monog. Phanerog. 2: 574 (1879).
Biarum tenuifolium (1.) Schott var. eupanianum (Guss. ex Paglia) Nicotra
ex Fiori, Nuova Fl. Anal. ltal. 1: 210
(1923).
Leaves erect, 10 cm X 14 mm, lamina
consistently oblanceolate to spathulate,
margins smooth to gently undulate. Staminodes peg-like. 2n = 26 (Marchant 1972
as var. abbreviatum).
Distribution-S Italy, Sicily, former Yugoslavia, W Greece.
Ecology-As for the typical variety. Alt.
120-1200 m.
Etymology-From the Latin abbreviatus
(shortened), referring to the smaller inflorescences compared with the type.
f. ssp. idomenaeum P.C. Boyce & Athanasiou in Flor Med. 1: 6 (1991). Type:
GREECE, Crete, Rethyrnnou, Mt. Psiloritis, above the village of Vizari,
650-750 m, Atbanasiou & Anagnostopoulos 566, (holotype UPA!, isotype
K! (photo)).
[B. tenuifolium (L.) Schott var. zelebori
auet. Crete. non (Schott) Engler
(1879)]
Leaves adpressed or parallel to the
ground, lamina linear to linear-oblong,
margins strongly undulate. Staminodes
filamentous. 2n = 26 (Athanasiou, unpublished).
Distribution-Crete.
Ecology-Grazed maquis on stony limestone-derived red clay. Alt. 100-300 m.
Etymology-Named in honour of the
mythical Cretan King Idomeneus.
As here defined Biarum tenuifolium
comprises six subspecies: viz. tenuifol-
ium, arundanum, galianii, abbreviatum,
zelebori and idomenaeum, separated on
the basis of leaf shape and size, spathe
size, spathe limb/spadix length ratio, degree of staminode development and phytogeography.
The typical subspecies has a long-exserted slender spadix appendix and
densely arranged, well-developed, simple,
curving staminodes. Early in the growing
season leaf blades are elliptic-Ianceolate,
later emerging leaves are linear-Ianceolate.
In immature plants the leaf blade is always
elliptic-Ianceolate. The typical subspecies
occurs from southern Italy to the southern
Balkans and is the common species in
mainland Greece.
The westernmost element is ssp. arundanum. It has a spadix appendix that does
not or only barely exceeds the spathe limb
and branching staminodes. Biarum arun-
PETER C. BOYCE, 2006
danum is one of two common Biarum in
southern and western Spain (the other is
B. carratracense) where it grows on heavy
red clay soils.
The easternmost subspecies, ssp. zelebOri, is distinguished by large bulky inflorescences, with the spathe limb averaging
20 X 3 cm, a robust, moderately exserted
spadix appendix and rather sparse but
substantial staminodes. The leaves do not
display the marked heteromorphy found
in the typical variety and the leaf blade is
spathulate-Ianceolate, often with gently
undulate margins. Subspecies zelebori is
restricted to southwest Turkey, Rhodes,
Cos and a few scattered sites on Crete.
Subspecies abbreviatum has erect, short
spathulate leaves, usually with undulate to
rarely somewhat crispulate margins. The
spathe limb averages 9 X 1.5 cm and is
notable for its bicoloured interior, deep
purple brown below with a striking green
apical portion. The spadix appendix is
generally only slightly longer than the
spathe limb and, compared to its length,
rather stout. The staminodes are rather
poorly developed and peg-like. Subspecies abbreviatum occurs in Italy (where it
has been called B. cupanianum Guss. ex
Paglia), Yugoslavia, FYRO Macedonia,
northern mainland Greece and on Corfu.
It has yet to be recorded from Albania
where the typical subspecies occurs, but
the presence of ssp. abbreviatum to the
south of Lake Ochrid, close to the Albanian frontier, suggests that it does occur in
Albania. Italian populations of B. tenuifolium have been referred to as B. cupanianum Guss. ex Paglia. Work by Monti &
Gabari (974) concluded that B. cupanianum could not be maintained at specific
rank and they reduced it to synonymy in
B. tenuifolium. However, the Italian populations are somewhat different to typical
ssp. tenuifolium, and require further scrutiny. Paglia (909) separated B. cupanianum from B. tenuifolium on differences in
the leaf emergence (hysteranthous vs. synanthous), leaf lamina shape Oanceolate-elliptic vs. lanceolate to linear), leaf length
(7-8 cm vs. 20 cm or more), spathe limb
size and colour (limb small, short, violet
11
or dark purple vs. limb large, long,
brown), features of the spadix appendix
(Y.! longer than the spathe limb thin, cylindrical, erect, dark purple with a dull grey
bloom vs. 3-4 times longer than the spathe
limb, thick, procumbent, reddish) and
phenology (spring flowering, inflorescence odourless or smelling of goats vs.
autumn flowering, inflorescence smelling
of dung). Comparison of these data with
the description below shows the majority
of characters used by Paglia fall within the
variation found in B. tenuifolium in the
eastern Mediterranean. The morphological
and phenological characters listed indicate
that B. cupanianum agrees closely with
ssp. abbreviatum as here defined. The
one disparity concerns Paglia's description
of a goat-like odour produced by B. cupanianum in flower. According to my
own observations Italian plants of ssp. tenuifolium smell almost identical to plants
from Greece, producing a powerful smell
of horse-dung. Biarum plants referable to
B. cupanianum have a sharper, more
urine-like, odour when in flower. A range
of diploid chromosome numbers has been
recorded for B. tenuifolium, e.g. 16, 20, 26
(Petersen 1989) and thus the count of 2n
= 16 recorded for B. cupanianum. (Del
Caldo 1971) does not exclude its amalgamation in B. tenuifolium. Given the
number of similarities between the taxa I
feel it best to regard B. cupanianum as a
synonym of B. tenuifolium ssp. abbrev-
iatum.
The remaining subspecies of B. tenuifolium are of more limited distribution.
Subspecies idomenaeum, from Crete, is
notable for its strongly undulate-crispulate
leaves that are closely adpressed to the
ground. The spathes are generally of similar size to those of ssp. abbreviatum, but
the staminodes are densely arranged and
slender. Most authors, e.g. Prime & Webb
(980); Barclay (986) and Greuter (973),
have referred these Cretan populations to
ssp. zelebori but this is incorrect. Although
ssp. zelebori occurs on Crete it is quite different in appearance. Mill (984) suggested that the smaller plants might be referable to ssp. abbreviatum. Morphologically
12
ssp. idomenaeum is closest to ssp. abbreviatum but readily separable by the filamentous staminodes. Other data including cytology (see Boyce & Athanasiou,
1991) also support this.
Two subspecies occur in the Iberian
peninsula: ssp. arundanum and ssp. galianii. While clearly defined by their ecological requirements-ssp. arundanum a
plant of red clay soils derived from limestone while ssp. galianii is restricted to
loose sandy soils-they are difficult to separate morphologically in the absence of
ecological data. Talavera (1976) cites differing cytology, staminode branching and
leaf lamina reduction as distinguishing
characters of the then species-ranked B.
arundanum and B. galianii, but the extent of disparity in the two taxa is rather
insignificant. However, one feature not
mentioned by Talavera, that of the arrangement of the staminodes, does appear
to be characteristic. In all the material of
ssp. arundanum and ssp. galianii examined the staminodes are always strictly
whorled in arundanum and irregularly
scattered in galianii. Quantitatively there
are sufficient grounds to maintain these
plants at a formal taxonomic rank.
Subspecies arundanum is widespread
in southwestern Spain, occurring in the regions of Cadiz, C6rdoba, Granada, Malaga
and Seville, and is often extremely abundant, forming extensive colonies alongside
cultivated land and beside paths, it is also
found in southern Portugal, Gibraltar and
northern Morocco. The freshly opened
spathe emits a particularly offensive odour
similar to cattle dung.
Subspecies galianii occurs in the regions of Badajoz and Huelva near the Portuguese border where it is restricted to
loose sandy soils.
2. Biarum rhopalospadix C. Koch,
Ind. Sem. Hort. BeroI. App. 2 (1853).
Type: 'Greece' (holotype B destroyed;
isotype K (tracing of Koch's specimen).
Epitype selected here:
GREECE, Attica, Mt Hymettus, 16 May
1856, Heldreich 512 (C, FI, G, L).
Biarum spruneri Boiss., Diagn. 13:5
AROIDEANA, Vol. 29
(1853); Engler in A. & C. DC., Monog.
Phanerog. 2: 574 (1879); Boiss., FI.
Or. 5: 32 (1882); Engler in Engler, Das
Pflanzenr. 73 (IV. 23F): 136 (1920).
Type: GREECE, in collibus apricis Atticae ad Phalerum portum ubi duce
amiciss, Spruner florere incipientem
Maii initio legi, (lectotype selected
here, G-BOIS). There are several
sheets collected from Phalire in the
Boissier herbarium. The sheet chosen
is the only Spruner collection and
thus is the logical lectotype. Ischarum
spruneri (Boiss.) Schott, Syn. Aroid. 7
(1856). Cyllenium spruneri (Boiss.)
Schott, Gen. Aroid. t.9 (1858).
Tuber dorso-ventrally compressed-discoid, 1.5-3 x 1-1.5 cm, offsetting sparsely,
mid-brown. Leaves 3-5, hysteranthous,
short-petiolate, bases encased by 3 or 4,
5-8 cm X 5-10 mm lanceolate, sub-fleshy,
later papery, cataphylls, these mid-green
drying to very pale green or creamy-white;
petiole 3-5 cm X 2-3 mm, adaxial surface
channeled distally, expanded proximally
into a membranous wing, mid to rather
dark green; leaf lamina oblong to oblongspathulate, 5-11 X 1.5-2.5 cm, apex obtuse to sub-acute, base decurrent to cuneate, 5-6 primary lateral veins per side,
margins smooth to slightly undulate, lamina mid-green. Inflorescence appearing
in spring, moderately foetid and smelling
of cattle dung, peduncle 3-9 cm X 2-4
mm, encased by 2 to 4, 4-11 X 1-1.5 cm
lanceolate, sub-fleshy, later papery, cataphylls, these mid-green drying to pale
straw-yellow. Spathe 7-24 cm long;
spathe limb oblong-lingulate to elliptic-oblong, 5-14 X 1-4 cm, acute to shortly acuminate, exterior green with dense purplebrown staining, especially towards the
margins and apex, interior concolorous
purple-brown, occasionally very slightly
greenish distally; spathe tube narrowly cylindrical, slightly inflated, 3-5.5 cm X 7-9
mm, margins connate for 3,4 of their length,
exterior off-white below, purple-brown
flushed above, interior white, slightly purple stained distally. Spadix sub-equal to
shorter than the spathe limb, 8.5-14.5 cm
13
PETER C. BOYCE, 2006
long; appendix fusiform, ± sessile, though
tapering in some specimens, 6-10.5 cm X
2-6 mm, deep purple. Staminate flowers in an oblong-cylindric zone 6-12 X
ca. 4 mm diam., cream. Interstice 11-25
X 3-4 mm, cream. Staminodes covering
the proximal half of the interstice, recurved, falcate, pointed, 1.5-3 mm long,
mostly simple but occasionally bifid,
cream. PistiUateflowers in a hemispherical cluster ca. 5 mm wide, 3 mm high;
ovary globose, slightly dorso-ventrally flattened, 0.25 mm wide and tall, cream; stigma. ca. 0.25 mm in diam., borne on a
0.25--0.5 mm long style, stigma greyish,
style purple. I7ifructescence not seen.
Chromosome number not recorded.
Distribution-S Greece, (Attica, Peloponnese).
Ecology-Limestone-derived red clay in
grazed fields, open hillslopes, abandoned
olives groves and field margins. Alt. 150450 m.
Etymology-From the Greek, rhopaloa club or cudgel-and spadix, in allusion
to the club-like spadix.
Biarum spruneri is superficially similar
to B. tenuijolium, especially to ssp. abbreviatum. It may be readily distinguished
by the lack of staminodes, the stigma
borne on a short style, the narrow, parallel-sided spathe limb, and the considerably
stouter spadix appendix. The spring flowering syndrome is also useful in distinguishing B. spruneri from the majority of
the Greek mainland populations of B. ten-
uijolium.
Boissier published B. spruneri in sect.
Biarum based on anther dehiscence via
ventral longitudinal slits and hook-like
staminodes, but noted the lack of staminodes above the staminate flower zone
and the well-developed style and suggested that it was intermediate between sect.
Biarum and sect. Ischarum. Schott (1858)
took this observation further by creating a
new genus, Cyllenium, for B. spruneri citing the rostrate anther connective, the
style and absence of upper staminodes as
the main diagnostic features of his new genus.
The hitherto obscure name B. rhopalospadix must now be used for Biarum spru-
neri.
3. Biarum aleppicum Thiebaut in Bull.
Soc. Bot. Fr. 95: 21 (1948); Mouterde,
Nouv. FI. Liban et Syrie 1: 193 (1966);
Riedl in Aroideana 3: 28 (1980). Type:
SYRIA, Alep, Fr. Louis s.n. (lectotype
P!, (chosen here)). Thiebaut cited two
separate collections in the protologue. There are four collections of B.
aleppicum present in Paris, including
both syntypes. The specimen chosen
as the lectotype is the most complete
of the cited specimens, consisting of
flowering, vegetative and fruiting material. [B. bovei Blume var. aleppicum
(Thiebaut) Gombault, in sched. nom.
nud.l
Tuber slightly dorso-ventrally compressed-globose, 3-4 X 1.5-2.5 cm, apparently not offsetting, mid-brown.
Leaves 10-35, hysteranthous, long but
obscurely petiolate, bases encased by 5-7,
2-15 cm X 10-13 mm, narrowly lanceolate
cataphylls, inner cataphylls sub-fleshy, later papery, pale greenish white drying pale
straw-yellow, outer cataphylls fibrous,
dark brown; petiole 3-7 cm X 1-3 mm,
abaxial surface slightly channeled distally,
expanded into a membranous wing proximally, dull green, expanded portion
tinged purple basally; leaf lamina linearlanceolate to oblanceolate-elliptic, 7-13
cm X 3-10 mm, apex obtuse to subacute,
base long-decurrent, ca. 5 primary lateral
veins per side, margins crispulate, rarely
smooth, lamina mid-green. Inflorescence
appearing in mid-autumn; peduncle 4-17
cm X 3-5 mm, whitish, clothed with several 2-16 cm X 10-13 mm fibrous brown
outer and papery, pale straw-yellow inner
cataphylls. Spathe 14-16.5 cm long;
spathe limb oblong-lanceolate, 12-13.5 X
4-5 cm, apex sub-acute, exterior pale
green ± heavily spotted deep purple, interior deep purple, occasionally with pale
green mottling and spots; spathe tube
14
AROIDEANA, Vol. 29
slender, 2.5-3 X ca. 1.5 cm, margins free
to the base, exterior white below ground,
deep purple above, interior white. Spadix
sub-equal to the spathe limb, 11-12 cm
long; spadix appendix sessile, fusiform, 88.5 cm X 2-6 mm, deep purple. Staminate flowers in a zone ca. 12 mm X 1.5
mm diam., anthers cream. Interstice ca.
16 X 2 mm, pale cream. Staminodes
clothing the basal half of the interstice,
densely arranged, slender, 4.5-5 mm long,
purple, often a few 1-1.5 mm staminodes
present high up on the interstice. Pistillate flowers in a hemispherical cluster ca.
2.5 X 7-9 mm; ovaries oblong, 1.5 mm X
0.5 mm, pale cream, stigma sessile, 0.250.33 mm, capitate, deep purple. Itifructescence globose, ca. 2 cm diam. when
semi-mature, consisting of ca. 40 berries;
berries 4 X 5 mm, pale lilac when ripe;
seed ovoid, 5 mm X 5-6 mm, testa slightly
reticulate, mid-brown. Chromosome number not recorded.
Distribution-NW Syria.
Ecology-Bare fields and plains on
limestone-derived red clay soils. Alt. 250450 m.
Etymology-The specific epithet comes
from Aleppo, a major town in northwestern Syria and the type locality of the species.
Biarum aleppicum is related to B. carducborum and B. angustatum. From either species B. aleppicum is readily distinguished by having the spathe tube margins
free almost to the base and by the sessile
stigmas. Biarum aleppicum has many
more leaves than either B. angustatum or
B. carducborum, although the leaf shape
approaches that of B. angustatum.
4. Biarum
angustatum
(Hook.f.)
N.E.Br. in Journ. Linn. Soc. 18: 255
(1881); Engler in Engler, Das Pflanzenr. 73 (N.23F): 142 (1920); Koach
& Feinbrun in Feinbrun, FI. Palaestina
4: 338 (1986); Koach in Rotem 28 t.19,
20 (1988). Type: ISRAEL, Tiberias,
September-October 1860, Hooker &
Hanbury s.n. (holotype K!). Iscbarum
angustatum Hook.f. in Bot. Mag. 104,
t.6355 (1878).
Tuber dorso-ventrally compressed-discoid, 2-5(-7) X 1.5 cm, sparsely offsetting,
mid-brown. Leaves 3-5, hysteranthous,
long-petiolate, bases encased by 3-6, 4.58.9 cm X 7.5-15 mm, lanceoiate-elliptic
cataphylls, inner cataphylls subfleshy, later
papery, dirty white drying straw-yellow,
outer cataphylls fibrous, brown, drying
slightly darker; petiole 13-17 cm X 2.5-3
mm, adaxial surface slightly channeled
distally, expanded into a narrow wing
proximally, mid-green; leaf lamina narrowly lanceolate-elliptic, 13.5 X 1.5 cm,
apex acute, base long-decurrent, 4-5 primary lateral veins per side, margins
smooth, mid-green. Inflorescence appearing in the autumn, strongly foetid of
horse dung and urine; peduncle 5.5-9 cm
X 2.5 mm, encased by several 3-10 cm X
8-15 mm sub-fleshy, later papery, cataphylls, these very pale greenish white,
pale creamy white on drying. Spathe 1924cm long; spathe limb narrowly lanceolate, 12-18 X 3-5 cm, apex acute, exterior
mid-green spotted and stained brownish
purple, interior deep brownish purple,
sometimes fading to green apically; spathe
tube narrowly cylindric, 3-3.5 X ca. 1 cm,
margins connate for ca. :JA of their length,
exterior whitish, stained deep purple towards the apex, interior white, slightly
purple stained near the apex. Spadix
slightly shorter than the spathe limb, 11.517 cm long; spadix appendix slender-cylindric, 13-14.5 X ca. 1.5 mm, deep
brownish purple. Staminate flowers in
zone 13-15 X 33.5 mm, anthers off-white.
Interstice 12-15 X 1-2 mm, off-white.
Staminodes arranged mostly at the base
of the interstice though with scattered rudimentary filaments higher up; filaments
slender, 2.5-4 mm long, directed downwards, white. PistiUate flowers in a
hemispherical cluster, ca. 8 mm X 4 mm
high; ovaries bottle-shaped 2-2.5 X 1-1.25
mm, purple, style 1.5-2 mm X ca. 0.2 mm
wide, purple, stigma sub-capitate, ca. 0.3
mm in diam, white. Infructescence
slightly compressed-globose, consisting of
15
PETER C. BOYCE, 2006
ca. 40 berries; berries ca. 5 X 6 mm, dirty
white, seeds globose, ca. 5 mm diam., testa brown, slightly reticulate. Chromosome
number not recorded.
Distribution-Syria, Israel. Its presence
in Lebanon has yet to be confirmed.
Ecology-Limestone-derived red clay
soils in open, grazed, sometimes almost
completely bare fields, undisturbed lake
side fields. Alt. 75-350 m.
Etymology-The specific epithet comes
from the Latin angustus, narrow, in allusion to the narrow leaves in comparison
to the remainder of Ischarnm, the genus
into which the name was first published.
A large-flowered species that although
fairly common in the wild has been seldom collected. Biarnm angustatum is outwardly very similar to B. carduchornm
when in flower but is easily separated by
the downward directed staminodes and
narrower leaves with only 4 to 5 primary
lateral veins per side. Another point of
separation concerns the distribution of the
species. Biarnm angustatum is essentially
a 'coastal' species, restricted to Syria and
Israel. Biarnm carduchornm is an inland
species, distributed from southeastern Turkey and northwestern Syria through Iraq
and into Iran.
5. Biarum carduchorum (Schott) Engler in A. & C. DC., Monog. Phanerog.
2: 575 (1879) & in Engler, Das Pflanzenr. 73 (IY.23F): 137 (1920); Mill in
Davis, Fl. Turkey 8: 57 t.2 no.14
(1984); Riedl in Townsend, Fl. Iraq 8:
194 (1985). Cyllenium carduchornm
Schott, Prodr. Syst. Aroid. 65 (1860).
Type: TURKEY, Kurdistania, Schirwan
(S ,irvan) , Kotschy s.n. (holotype W
destroyed; lectotype W, chosen here,
Schott's leones no. 1825». There are
four illustrations present in Vienna
(leones nos. 1824-1827), the one selected is annotated 'Kurdistan Schirwan bei Kariig'.
Biarnm platyspathum Bornm. in Feddes
Rep. Nov. Sp. 5: 57 (1908). Type:
IRAN, Persiae austro-occidentalis
provincia Farsistan, ad Bascht et Fahliun, 12 November 1905, Herzfeld s.n.
(holotype B destroyed; lectotype B
(selected here (drawing of the holotype made by Bornmtiller». B. carduchornm (Schott) Engler var. platyspathum (Bornm.) Engler in Engler,
Pflanzenr. 73 (N.23F): 137 (1920).
B. platyspathum (Bornm.) Engler var.
bakhtyarnm Parsa in Kew Bull. 4: 36
(1949). Type: IRAN, Fars, Abe Bariq
(Abibarik, about 6 miles from E Assupas), 1 September 1885, Stapf s.n.
(holotype K).
[B. angustatum (Hook.f.) N.E.Br. var. kurdistanicum Zohary, in sched. nom.
nud.l
[B. bakhtyarnm Stapf, in sched. nom.
nud.l
Tuber dorso-ventrally compressed-discoid, 4-7 cm X 1.5-2.5 cm, sparsely offsetting. Leaves 3-5, hysteranthous, longpetiolate, bases encased by 3 to 5, 6-14(24) X 1-2 cm sub-fleshy, later papery, cataphylls, these pale whitish green, pale
straw-yellow on drying; petiole slender, 59(-26) X 2-4 mm wide, expanded proximally into a membranous wing, dull
green, wing paler; leaf lamina elliptic to
spathulate-elliptic, 5-12 X 2.5-5 cm, apex
subacute to rather obtuse, base briefly decurrent, 5-7 primary lateral veins per side,
margins smooth, dull mid-green. Inflorescence appearing in late autumn; peduncle
4-13(-24) cm X 2-4 mm, dirty white,
clothed with several 3-11(-23) X 1-2 cm
sub-fleshy, later papery, cataphylls, these
whitish green, pale straw-yellow on drying. Spathe 12-18(-31) cm long; spathe
limb lanceolate to Ian ceolate-elliptic, 814(-25) X 2-3(-4.5) cm wide, apex acute
to attenuate, exterior pale green to whitish
yellow usually ± speckled with dull purple, interior deep brown purple, becoming
paler and eventually green distally; spathe
tube slender, 4-6 X 0.75-1.5(-2.5) cm
wide, margins connate for ca.
of their
length, exterior dirty white where buried,
purple where exposed, interior white,
stained purple especially basally and towards the opening. Spadix sub-equal to
*
16
but rarely exceeding the spathe limb, 1318(-32) cm long, spadix appendix slender
cylindric, 9-12(-28) cm X 2-4 mm, deep
purple. Staminate flowers in a zone 1525 X 2-3.5 mm, anthers cream ± stained
deep purple. Interstice 2-3 cm X 2-3
mm, deep purple, occasionally somewhat
paler than the appendix. Staminodes
densely arranged at the base of the interstice directly above the pistillate flowers
and usually extending ca. half way up the
interstice; filaments directed upwards, 3-7
mm long, those higher up the interstice
shorter than those lower down, purple.
Pistillate flowers in a hemispherical
cluster ca. 7 X 7 mm; ovaries squatly bottle-shaped, 1.5-2 X 0.75 mm, pale cream,
style ca. 0.33 mm long, purple, stigma subcapitate, ca. 0.25 mm in diam, purple. Infructescence not seen. 2n = 24 (Marchant 1972 as B. platyspathum).
Distribution-S and SE Turkey, Syria,
Iraq, W Iran ..
Ecology-Bare limestone-derived red
clay hill slopes, in open situations, field
margins, long-follow fields. Alt. 290-2,750
m.
Etymology-After the name of an ancient tribe, the Carduchi, that inhabited the
region of southeastern Turkey where the
type was gathered.
Biarum carduchorum is fairly widespread, occurring from southern and
southeastern Turkey to southern Iran.
There is some variation in the populations
and names have been published to account for this. However, any variation present is generally of little or no taxonomic
significance. The type of the plant described by Parsa (1949) as var. baktaryanum on the basis of a narrow spathe with
a whitish yellow exterior, is almost identical to the illustration in Vienna that serves
as the type of B. carduchorum. Biarum
platyspathum is a particularly vigorous
form of B. carduchorum.
Biarum carduchorum is most readily
separated from B. angustatum by the upward-directed staminodes and the consid-.
erably wider leaves. As pointed out above,
AROIDEANA, Vol. 29
B. carduchorum is found further inland
than B. angustatum.
6. Biarum eximium (Schott & Kotschy)
Engler in A. & C. DC., Monog. Phanerog. 2: 576 (1879) & in Engler, Das
Pflanzenr. 73(IV.23F): 139 (1920); Mill
in Davis, FI. Turkey 8: 57 (1984); Mathew, The Smaller Bulbs 16 (1987). Ischarum eximium Schott & Kotschy in
Oesterr. Bot. Wochenbl. 4: 81 (1854).
Type: TURKEY, Taurus, [prope Adana, in via romana versus Miaret Chan,
60 m, 28 September 1853, Kotschy
3431 (holotype W destroyed; isotypes
G-BOIS, K, M, P).
Tuber dorso-ventrally compressed-discoid, 3-7 X 2-3 cm, rarely offsetting.
Leaves hysteranthous, long-petiolate, bases encased by several 5-7 cm X 7.5-12
mm lanceolate-elliptic cataphylls, these
pale greenish white, occasionally purplespotted apically, drying dark straw-yellow;
petiole 5-11 cm X 4-6 mm, mid-green,
adaxial surface broadly channeled distally,
expanded into a narrow membranous
wing proximally; leaf lamina elliptic, 1012.5 X 3-3.5 cm, apex sub-acute, base decurrent, ca. 9 primary lateral veins per
side, margins smooth, mid-green. Inflorescence appearing in autumn, strongly
foetid; peduncle 1-3 cm X 3-5 mm, off
white, encased by several 2-4 cm X 5-9
mm wide smooth pale straw-yellow cataphylls. Spathe 9-13 cm long; spathe tube
cylindrical, 4-4.5 cm long, ca. 2 cm wide,
moderately inflated, margins connate for
ca. Ih their length, exterior green with
much purple staining and spotting distally;
interior white, stained purple proximally;
spathe limb broadly elliptic, 8-11 X 3.755 cm wide, rounded distally, exterior
green with numerous irregular purple
spots, interior deep brown-purple. Spadix slightly shorter than the spathe limb,
9-10.5 cm long; spadix appendix fusiform,
7-7.5 cm X 5-7 mm, purple, sometimes
slightly paler than the spathe limb. Staminate flowers in a zone 12-14 mm X
4-5 mm wide, purple; Interstice 25 mm
long, ca. 4 mm wide, purple. Staminodes
17
PETER C. BOYCE, 2006
distributed ± evenly over the entire length
of the interstice, filaments ca. 5 mm long,
purple. Pistillate flowers in a hemispherical cluster, ca. 10 mm X 3 mm tall;
ovaries oblong, ca. 2 mm X 0.7 mm,
cream below, purple above, style ca. 1-1.5
mm long, purple, stigma capitate, purplish
grey. Infructescence a globose cluster of
ca. 40 pyriform berries 2.5-3 cm in diam.;
berries ca. 5 X 8 mm, dull white ± stained
purple; seeds ovoid, ca. 4-5 mm in diam.,
mid-brown, testa reticulate. 2n = 16 (Marchant 1972).
Distribution-S Turkey, Jordan.
Ecology-limestone-derived red clays
in open habitats. In Jordan it occurs in
stony loamy soils in semi-desert. Alt. ca.
200 m.
Etymology-Either from the Latin eximius (unexpected) or eximie (exceptionally). The exact derivation is unclear.
Biarum eximium differs from all other
species of subgenus Ischarum in that the
staminodes are evenly distributed over the
entire length of the interstice between the
male and female flower zones. Examination of a wide range of material of other
species, particularly B. carduchorum and
B. angustatum, revealed that although the
main area of staminode distribution was
directly above the pistillate flowers, in
many individuals staminodes were present
on the upper portion of the interstice, although usually in a depauperate condition.
Biarum eximium has been little collected and were it not for the large number of
isotypes, it would be poorly known. The
other collections in European herbaria are
Siehe 22 (dry sterile material with a single
spirit-preserved inflorescence in B) and
Dinsmore 11725 from Jordan. This latter
collection is most interesting since, to date,
B. eximium has not been collected in the
area between the type locality in southern
Turkey and this Jordanian site. El-Eisawi
(1981) sites another collection (Thab'a
(Dab'a) Reserve, ca. 50 km south of Amman, along the road to Aqaba, AI-Eisa wi
8861) which is deposited in the University
of Jordan herbarium (AMM). Although I
have not seen this specimen, I have examined living plants collected by Salmon
and Lovell very close to the same site in
1988. Further collections of B. eximium,
particularly from Turkey, would be most
desirable.
7. Biarum bovei Blume, Rumphia 1: 114
(1836); Engler in A.& C. DC., Monog.
Phanerog. 2: 577 (1879) & in Engler,
Das Pflanzenr. 73(IV.23F): 140 (1920);
Mill in Davis, Fl. Turkey 8: 58(984);
Riedl in Townsend, Fl. Iraq 8: 195
(1985); Koach & Feinbrun in Feinbrun, FI. Palaestina 4: 337 (986);
Koach in Rotem 26 t.17,18 (988).
Type: LEBANON, Mt liban, 1832,
Bove 379 (holotype L; isotypes G, K,
P).
The material in Leiden has been annotated as the isotype by Nicolson. Blume
(1836) states that the material that he
based the description of B. bovei on was
in both the Leiden and Paris herbaria. Both
sets of material have been annotated by
Blume and furthermore Blume worked out
of Leiden. Since the Leiden specimens are
in reasonably good condition while those
in Paris are rather poor, it is preferable that
the Leiden material is regarded as the holotype.
Caladium bovei (Blume) Steud., Nomen.
Bot. ed.2, 1: 249 (1840).
Ischarum bovei (Blume) Schott, Syn. Aroid. 7 (1856).
Biarum bovei Blume var. blumei Engler in
A. & c. DC., Monog. Phanerog. 2: 577
(1879). Type: as for B. bovei Blume.
Biarum homeid Blume, Rumphia 1: 115
(1836). Type: SYRIA, Aleppo, Rauwolff s.n. (holotype not traced).
Ischarum homeid (Blume) Schott, Syn. Aroid. 8 (1856) ["homaid'J.
Tuber globose, slightly compressed
dorsally, 1.5-3.5 cm in diam., offsetting
sparsely. Leaves 3-12, hysteranthous, bases encased by several 4-11 X 1-1.5 cm
sub-fleshy, later papery, cataphylls, these
pale green to dirty white, drying to off-
18
white; petiole 5-9 cm X 3-5 mm, adaxial
surface slightly to rather strongly channeled distally, expanded proximally into a
wide membranous wing, mid-green, wing
paler; leaf lamina lanceolate-oblong to
ovate-Ianceolate with 6-8 primary lateral
veins per side, 5-10 X 1.5-4 wide, margins
smooth to undulate, more rarely crispulate, mid-green. InjIorescence appearing in
late autumn, 7-19 cm long, strongly malodorous of horse dung; peduncle 4-7 cm
long, encased by numerous 5-9 X 1-1.5
cm sub-fleshy, later papery, cataphylls,
these pale green, dull straw-yellow on drying. Spathe limb lanceolate to linear-Ianceolate, 5-14.5 X 1.75-3 cm, margins flat
and ± smooth, exterior dull green with
some purple staining or spotting, especially towards the margin, interior deep
purple-brown; spathe tube globose-cylindric to globose, margins connate for ca. 1,4
of their length, 2.5-4 X ca. 1.5 cm in
diam., exterior off-white, dull green with
slight purple staining where exposed, interior pale green, heavily stained purple at
the base. Spadix 9-14 cm long; spadix
appendix narrowly fuSiform, 6-10 cm X
2.5-3 mm, deep purple. Staminate flowers in a zone 8-12 X 3-4 mm diam.,
cream. Interstice 1.5-2 cm X 2-3 mm.,
cream, stained purple in the lower quarter.
Staminodes few to many, clustered in a
zone ca. 1 cm long directly above the pistillate flowers; bristles filiform, 5-15 mm
long cream, tinged purple basally. Pistillate flowers in a hemispherical cluster ca.
9 X 6 mm, ovaries pale cream, style 1.5 X
0.25 mm, purple, stigma capitate, purple.
Infructescence a globose cluster of ca. 40
pyriform-globose berries 2.5-3 cm in
diam.; berries ca. 7 X 4 mm, pale lilac-grey
with purple tinges when ripe; seeds globose, ca. 4-4.5 mm in diam., pale brown,
testa reticulate. Chromosome number not
recorded.
Distribution-Turkey, Syria, Lebanon,
Israel, Iran and Iraq.
Ecology-Limestone-derived red clays
in open situations, hill slopes, grazed pasture, field margins. Alt. 800-1,750 m.
AROIDEANA, Vol. 29
Etymology-Named for the collector of
the type material.
Biarnm bovei has been confused with
B. kotsehyi and B. pyrami both in the field
and in herbaria. Part of the trouble appears to stem from the scarcity of true B.
bovei in herbaria coupled with the fact that
the most of the type specimens are
pressed in such a way that comparative
analysis is rather difficult. In addition, confusion with the Afro-Iberian B. dispar has
led to the belief that B. bovei is consistently variable throughout its range when,
in fact, the variation has a clear geographical basis. Schott understood this and published names to account for the various
populations; e.g. Iseharnm erispulum and
I. earsaami (Kunth) Schott. Biarnm erispulum is morphologically constant in the
field and restricted to a small region: the
provinces of Adana and Hatay in southern
Turkey and in northern Syria. There is,
however, a western extension to Cappadocia in the Siehe collection Berlin, discussed below. The differences that distinguish B. erispulum from B. bovei are rather minor. I have reduced it to varietal rank
within B. bovei.
The supposed presence of B. bovei in
Sardinia (Monti & Gabari, 1974) is problematic and, on the basis of material flowered in cultivation, would appear to be the
result of a misidentification. Monti & Gabari (974) only briefly discussed the Sardinian populations of B. bovei. Recently I
have been sent living material of B. bovei
collected in Sardinia by Josef Bogner. On
flowering it was clearly referable to B. dis-
par.
8. Biarum crispulum (Schott) Engl. in
Bot. Jahrb. 5: 334 (1884). Type:
'IRAQ, ad Arcem Semiramidis' (but
see note below), Kotschy, cult.
Schoenbrunn (holotype W destroyed;
lectotype G-BOIS, chosen here: TURKEY: Adana, Kassan Oghlu (Hasanoglu) Gorumse (Giiriimze) valley, 21
May 1859, Kotschy 442).
The type locality stated by Schott for Ischarnm crispulum is possibly in error.
PETER C. BOYCE, 2006
Several of Schott's Ischarum protologues
state the type locality to be Semiramis but,
as pointed out by Mill (1984) when discussing B. pyrami, Semiramis is in Iraq,
whereas the annotations on the type material of B. pyrami state that is was collected near the Ceyhan River, Adana, in
southern Turkey. Given the paucity of authenticated Iraqi B. bovei (with which B.
crispulum is much confused) collections,
it seems likely that I. crispulum originated
from either northwestern Syria or south
eastern Turkey. This is further supported
by more recent records of this taxon from
these areas but not, as yet, from Iraq. The
illustration in Vienna (W) of living material
of the lectotype (Schott, /cones Aroideae
no. 2141), is annotated in Schott's hand as
I. crispulum with the data 'Kassan Oghlu,
Gorumse, Kotschy 1859', Le., from Gilrilmze, Adana, southern Turkey. Ischarum
crispulum Schott, Prodr. Syst. Aroid. 68
(1860).
Calla orientalis 1., Sp. PI. ed.2: 1373
(1763). Type: 'Arum carsamf Rauw
it. 115. Halepi in montosis (L), nom.
rejic.
Arum carsaami Kunth, Enum. PI. 3: 25
(1841), nom. illeg. Type as for Calla
orientalis.
Eminium carsaamii (Kunth) Schott, Syn.
Aroid. 17 (1856), nom. illeg.
Ischarum carsaamii (Kunth) Schott,
Prodr. Syst. Aroid. 67 (1860), nom. illeg.
Biarum bovei Blume 13 carsaami (Kunth)
Boiss., FI. Or. 5: 34 (1882) ['karsaamit'].
Biarum orientale (L.) Druce in Bot. Exc.
Club Brit. Isles 3(5): 415 (1914)
[Ischarum christmannii Siehe in sched.
Berol. nom. nud.l A flowering specimen collected by Siehe (Siehe s.n.)
preserved in alcohol in Berlin (B) is
annotated Ischarum (Biarum) christmannii Siehe. I have been unable to
trace any publication place for the
name. The specimen is clearly referable to B. crispulum.
Tuber globose, slightly compressed
19
dorsally, 1.5-3.5 cm in diam., offsetting
sparsely. Leaves 3-12, synanthous, bases
encased by several 4-11 XI-I. 5 cm subfleshy, later papery, cataphylls, these pale
green to dirty white, drying to off-white;
petiole 5-9 cm X 3-5 mm, adaxial surface
slightly to rather strongly channeled distally, expanded proximally into a wide
membranous wing, mid-green, wing paler;
leaf lamina lanceolate-oblong to ovate-lanceolate with 6--8 primary lateral veins per
side, 5-10 X 1.5-4 wide, margins smooth
to undulate, more rarely crispulate, midgreen. Itiflorescence appearing in late
autumn to early winter, weakly odorous of
sour milk, 7-11 cm long; peduncle 4-7 cm
long, encased by numerous 5-9 X 1-1.5
cm sub-fleshy, later papery, cataphylls,
these pale green, dull straw-yellow on drying. Spathe limb linear-triangular, 5-14.5
X 1-1.5 cm, margins incurved and strongly crispulate, exterior dull green with some
purple staining or spotting, especially towards the margin, interior dark greenish
purple; spathe tube globose-cylindric to
globose, margins connate for ca. % of their
length, 2.5-4 X ca. 1.5 cm in diam., exterior off-white, dull green with slight purple
staining where exposed, interior pale
green, heavily stained purple at the base.
Spadix 4-10 cm long; spadix appendix
narrowly fusiform, 3-8.5 cm X 2.5-3 mm,
deep greenish purple. Staminate flowers in a zone 8--12 X 3-4 mm, cream, thecae tipped deep purple. Interstice 1.5-2
cm X 2-3 mm., cream, stained purple in
the lower quarter. Staminodes few to
many, clustered in a zone ca. 1 cm long
directly above the pistillate flowers; bristles filiform, 5-15 mm long cream, tinged
purple basally. Pistillate flowers in a
hemispherical cluster ca. 9 X 6 mm, ovaries pale cream, style 1 X 0.5 mm, purple,
stigma capitate, purple. Infructescence a
globose cluster of ca. 40 pyriform-globose
berries 2.5-3 cm in diam.; berries ca. 7 X
4 mm, pale lilac-grey with purple tinges
when ripe; seeds globose, ca. 4-4.5 mm in
diam., pale brown, testa reticulate. Chromosome number not recorded.
Distribution-S Turkey (provinces of
Adana, Hatay and Konya), NW Syria.
20
Ecology-Limestone-derived red clays
in open situations, hill slopes, grazed pasture, field margins. Alt. 650-900 m.
Etymology-The epithet crispulum refers to the crispulate spathe limb margins.
Since publication B. crispulum has been
universally treated as a synonym of B. bovei although B. crispulum is readily separable by its synantherous leaves, the narrow, incurved and heavily crispulate
spathe margins, and a spadix appendix
smelling of sour milk (dung in B. bovez).
Biarnm crispulum is the commonest
Biarnm in NW Syria, forming extensive
colonies in bare red soil to the north of
Aleppo.
9. Biarum dispar (Schott) Talavera in
Lagascalia 6(2): 293 t.l, D, D1 (1976);
Talavera, Valdes & Galiano, F!. Vase.
de Anda!. Occ. 3: 210 (1987). (1976).
Ischarnm dispar Schott, Syn. Aroid. 7
(1856). Type: ALGERIA, Constantine,
mountains, October 1838, Bove s.n.
(holotype W destroyed; isotypes FI,
G, OXF, P).
Biarnm numidicum Par!., F!. Ita!' 2: 243
(1857) ("1852") nom. superfl. Type:
as for B. bovei Blume.
B. macroglossum Pomel, Nouv. Mat. Fl.
Adant. 2: 390 (1874). Type: ALGERIA,
Valle du CheJif, Tell, terrains argileux
(holotype not traced).
B. longifolium Pomel, Nouv. Mat. F!. Atlant. 2: 391 (1874). Type: ALGERIA,
Nador de Tiaret, au pied des rochers
(holotype not traced).
B. rnpestre Pomel, Nouv. Mat. Fl. Adant. 2:
391 (1874). Type: ALGERIA, Miliana,
Boghar, cavites des roc hers calcaires
(holotype not traced).
B. bovei Blume ssp. dispar(Schott) Engler
in A. & c. DC., Monog. Phanerog. 2:
587 (1879).
B. bovei Blume ssp. dispar (Schott) Engler
var. viride Battandier in Bull. Soc. Bot.
Fr. 28: 269 (1881) ["viridis"]. Type: not
designated (AL?).
B. bovei Blume ssp. dispar (Schott) Engler
var. rnpestre (Pomel) Battandier &
AROIDEANA, Vo!' 29
Trabut in Trabut, Fl. d'Aiger 17 (1884)
["rnpestris' 'l.
B. bovei Blume ssp. dispar (Schott) Engler
var. zanonii Pamp. in Nuov. Giorn.
Bot. Ital. 24: 124 (1917). Type: LIBYA,
Raaba, steppe, 2 December 1916,
Pampanini 216 (lectotype Fl, chosen
here). Pampanini cites two specimens
in the protologue, the other (Guarcia,
27 December 1916, Pampanini 189)
consists of leaves and fruit, while that
chosen is in flower and thus the better choice as lectotype.
B. bovei Blume ssp. dispar (Schott) Engler
var. purpureum Engler in Engler,
Pflanzenr. 73 (IV.23F): 141 (1920).
Type: ALGERIA, Thikilmouth, Constantine, in pasquis argilloso petrosis
summi montis Mansourah, 10 November 1868, Paris 293 (lectotype B, chosen here; isolectotypes FI, G, K, P).
The collection selected is the best of
those cited by Engler. I have chosen
the Berlin specimen as the lectotype
since it was undoubtedly seen by Engler.
B. bovei Blume ssp. dispar (Schott) Engler
var. macroglossum (Pome!) Maire &
We iller, Fl. de l'Afr. Nord 4: 247
(1957). Type: Not cited.
B. bovei Blume ssp. dispar (Schott) Engler
var. macroglossum (Pome!) Maire &
Weiller f.
longifolium (Pome!) Maire & Weiller, Fl.
de I'Afr. Nord. 4: 247 (1957). Type:
Not cited.
Tuber globose-discoid, 2.5-4 X 2-3.5
cm, mid-brown. Leaves 4-10, hysteranthous, long petiolate, bases encased by 3
to 5, 8-14(-16) cm X 3-10(-15) mm lanceolate sub-fleshy, later papery cataphylls,
these pale greenish white at first, drying
pale straw-yellow; petiole 10-17 cm long,
2-3 mm wide, adaxial surface strongly
channeled distally, expanded into a wide
membranous wing proximally, mid-green;
leaf lamina oblong-elliptic, 6--8 X 3-3.5
cm, apex cuneate to rounded, base obtuse
to sub-acute, ca. 3--6 veins per side, margins smooth, lamina mid-green, very rarely
bullate. Inflorescence appearing in late
PETER C. BOYCE, 2006
summer to autumn, smelling moderately
of cattle dung and carrion; peduncle 3--9
cm X 3--5 mm, clothed by few to many 29 cm X 5-12 mm papery, pale creamy
white cataphylls. Spathe 8-10(-14) cm
long; spathe limb lanceolate, 6-8(-12) cm
X 4-12 mm, apex acute to acuminate, exterior green ± heavily blotched and
stained purple-brown, rarely unstained, interior deep purple-brown, paler distally,
rarely entirely dull green; spathe tube oblong-globose, strongly inflated, 2-3 X 1.52.5 cm wide, margins fused for 1,4 of their
length, exterior pale green, occasionally
stained purple-brown towards the opening, interior off-white distally, deep purple
proximally. Spadix sub-equal to just exceeding the spathe limb, 8-12 cm long;
spadix appendix slender fusiform, 6.5-11
cm X 3-5 mm, deep purple-brown. Staminate flowers in a zone 9-13 X 3-6
mm, deep purple. Interstice 12-20 X 36 mm, deep purple. Staminodes situated
at the base of the interstice, few to many,
thickened-filiform, 2-10 mm long, deep
purple. Pistillate flowers in a hemispherical cluster 2.5-4 X 3-7 mm wide;
ovary 2-3 mm long, cream; style 1-1.5 X
0.25 mm, purple, stigma capitate, ca. 0.5
mm in diam., grey-purple. Infructescence globose, 1.6-3 cm in diam., consisting of ca. 35 berries; berries 4-6 X 4-5
mm, pale whitish lilac when ripe; seed
globose, ca. 5 mm in diam, testa reticulate,
pale brown. 2n = 74 (Chiappini & Scrugli
1972, Talavera 1976).
Distribution-SW Spain, N Morocco, N
Algeria, N Tunisia, N Libya, Sardinia.
Ecology-Open stony fields, rocky hill
slopes, crevices and chimneys in limestone rocks, field margins, track sides, disused olive groves. Alt. 25-250 m.
Etymology-From the Latin dispar, unlike, unequal, but in which context is not
known.
This interesting species has been much
confused with the closely allied B. bovei
from the eastern Mediterranean. It can be
distinguished readily by the interstice
which is approximately twice as long as
21
the staminate flower zone, the presence of
fewer and more scattered staminodes, and
the narrower mature leaf blade. The overall size of the inflorescences has been used
previously as a diagnostic feature, B. dispar having a smaller inflorescence than B.
bovei. While this appears to be true for the
North African populations of B. dispar, it
does not necessarily hold true for Spanish
populations, which are often as large as, if
not larger, than typical B. bovei. However,
part of the apparent size overlap between
the species appears to be due to a previously overlooked taxon, B. mendax,
which displays dimensions in excess of
both B. dispar and B. bovei and is readily
separable from either. References to B.
bovei in the Flora of Libya (El Gadi 1977)
are referable to B. dispar.
10. Biarum olivieri Blume, Rumphia 1:
115 (1836); Engler in A. & c. DC.,
Monog. Phanerog. 2: 580 (1879) & in
Engler, Pflanzenr. 73 (IY.23F): 142
(1920); Koach & Feinbrun in Feinbrun, Fl. Palaestina 4: 339 (1984);
Koach in Rotem 26 t.21,22 (1988).
Type: EGYPT, Olivier & Bruguiere
s.n. (holotype P; isotypes K, L). Ischarum olivieri (Blume) Schott, Syn.
Aroid. 8 (1856) & in Miq., Ann. Mus.
Bot. Lugd.-Bat. 1: 278 (1864).
Biarum alexandrinum Boiss., Diag. 13: 6
(1853). Type: Egypt, near Alexandria,
Cadet de Fonteney s.n. (holotype not
traced). Ischarum alexandrinum
(Boiss.) Schott, Syn. Aroid. 8 (1856).
Leptopetion alexandrinum (Boiss.)
Schott, Gen. Aroid. t.8 (1858).
Tuber globose to somewhat sub-cylindric, 7.5-20 X 5-25 mm. Leaves 3-8, hysteranthous to synanthous, long-petiolate,
bases encased by 2 to 4, 5-14 cm X 5-13
mm, lanceolate, sub-fleshy, later papery,
cataphylls, these green at first, drying
greenish white; petiole 4-11(-16) cm X 12 mm, adaxial surface channeled distally,
expanded proximally into a membranous
wing, pale green; leaf lamina linear to linear-lanceolate, rarely lanceolate, 7-14 cm
X 2.5-10 mm, apex acute to sub-acumi-
AROIDEANA, Vol. 29
22
nate, base decurrent to rounded or subtruncate, ca. 3-5 veins per side, margins
undulate to crispulate, lamina pale to midgreen. Inflorescence partially encased by
2-3 cataphylls, the spathe tube often partially to completely buried. Spathe ｾ＠
cm
long; spathe limb linear, the margins incurved, 1-6 cm X 1.5-3 mm, exterior midgreen; interior dull purple, rarely olive
green; spathe tube ovoid, strongly inflated,
1-2.5 X 0.75-1.5 cm, margins connate for
their whole length, exterior pale to midgreen, sometimes flushed purple towards
the opening, interior deep purple. Spadix
subequal to the spathe limb, 2.5-8 cm
long; spadix appendix filiform, 2-6.5 cm
X 0.5-1 mm, deep purple. Staminate
flowers in a zone 3-4 mm X ca. 1 mm
diam., anthers scattered to rather dense,
purple. Interstice 4-5 X ca. 0.75 mm,
deep purple. Staminodes few, often absent, filiform, situated at the base of the
interstice, 2-3 mm long, purple. Pistillate
flowers in a hemispherical cluster 2-4 X
3-4 mm; ovaries bottle-shaped, ca. 1-5 X
0.75 mm, cream, style ca. 1 X 0.15 mm,
purple, stigma capitate, ca. 0.33 mm in
diam., purple-grey. Infructescence globose, ca. 1.1-2 cm in diam., often enclosed in the persistent spathe tube, head
consisting of ca. 25-30 berries; berries globose, ca. 5 mm in diam., pale whitish-purple when ripe; seed spherical, ca. 3 mm in
diam., testa ± smooth, elaiosome barely
developed, pale brown. Chromosome
number not recorded.
Distribution-N Egypt, S Jordan, S Israel.
Ecology-In consolidated sand in open
situations, often close to the coast. Alt. 5500 m.
Etymology-Named after Guillaume
Antoine Olivieri, a French naturalist who
travelled extensively in Asia Minor during
the late eighteenth century.
Biarum olivieri is a curious species that
is allied to B. bovei, B. dispar and B. crispulum but is separable by the linear to linear-Ianceolate leaves, the much thinner, almost papery, spathe texture, the fully con-
nate spathe tube margins and the filamentous spadix appendix. The floral odour
produced by B. olivieri is also distinctive.
Both B. dispar and B. bovei produce a
dung-like odour which, although unpleasant, is not nauseating; Biarum olivieri produces a disgusting smell of soured milk
when in blossom, similar to that produced
by B. crispulum. The habitat favoured by
B. oliVieri, consolidated sand, is quite different from the heavy red clays favoured
by B. bovei, B. dispar and B. crlspulum.
To date B. olivieri is known from three
locations. The type locality is an area of
sub-coastal sands on the Egyptian coast
where it forms extensive colonies in association with Arisarum vulgare Targ.Tozz. and Eminium spiculatum (Blume)
Schott. A second, more recently discovered, site is in the Negev Desert in southern Israel, while in 1995 Chris Lovell collected B. olivieri in Jordan on the road
from Ibria to Rauble.
11. Biarum straussii Engler in Engler,
Das Pflanzenr. 73(IY.23F): 142(920);
Riedl in Townsend, Fl. Iraq. 8: 196-7
t.51 (1985). Type: IRAN, tal des Drehemschur bei dem Dorf Meikham auf
Brachen, 25 May 1906, Strauss 590
(lectotype B, chosen here). All of the
syntypes listed by Engler are present
in Berlin. The specimen selected is
the most complete of these.
Tuber slightly compressed-globose, 24 X 1.5-3 cm, not offsetting (?), pale
brown. Leaves 4-18, proteranthous, long
petiolate, bases encased by 3 to 4,6-14 X
1-1.5 cm rather fleshy, later papery, cataphylls, these pale greenish white at first,
pale brown on drying; petiole 1-3 mm
thick, adaxial surface barely channeled
distally, the outer-most petioles expanded
into membranous wing proximally, the innermost ± the same width along their
length, petiole mid-green, wing paler; leaf
lamina ovate-elliptic to oblong to linear, 514 cm X 4-20 mm, apex obtuse to subacute, base decurrent to almost truncate,
ca. 6-8 veins per side, margins smooth to
undulate or slightly crispulate, lamina dull
PETER C. BOYCE, 2006
green. Inflorescence emerging from the
middle of the leaf rosette; peduncle 4-13
cm X 2-2.5 mm, subterranean or slightly
emergent, whitish to pale green where visible. Spathe 8-16 cm long; spathe limb
lanceolate, 6-12 cm X 5-10 mm, acute,
exterior green, interior deep purple;
spathe tube oblong, inflated, 3-5 X l.752.3 cm, margins fully connate, exterior
green, interior deep purple, occasionally
paler than the spathe limb interior. Spadix sub-equal to slightly exceeding the
spathe, 8-16 cm long; spadix appendix cylindric-fusiform, 6-13.5 cm X 2-3 mm,
deep purple. Staminate Jlowers in a
zone 12.5-15 mm long and 2-4 mm diam.,
anthers cream. Interstice l.5-3 cm X 23 mm, cream. Staminodes restricted to a
5-10 mm zone above the pistillate flower
zone, few to many, 4-6 mm long, mostly
pointing upwards, cream. PistillateJlowers in a hemispherical cluster 3-3.5 X 56 mm; ovaries oblong, 1-l.5 X ca. 1 mm,
cream with a purple apex, style ca. 1 X
0.25 mm, purple, stigma capitate, ca. 0.33
mm in diam, purple. Infructescence globose, 2.5-3 cm in diam., consisting of 4050 berries; berries 7-10 X 4.5-6 mm pale
purple-white when ripe; seed ca. 4 mm in
diam., ovoid, testa slightly reticulate. Chromosome number not recorded.
Distribution-Northern Iraq, northern
Iran.
Ecology-Limestone-derived red clay
soils or occasionally on shale outcrops on
rocky and stony limestone hills, in open
situations, amongst low Berberis scrub or
on grazed hillslopes. Alt. 300-550 m.
Etymology-Named for the collector of
the type material.
The distinctive appearance of B. straussii in flower-the inflorescence emerging
from the middle of a mature leaf rosetteprovides little chance of confusing it with
any other species of Biarum, except perhaps B. syriacum from which it differs by
the much broader leaf lamina.
Confusion might occur with species of
Eminium, especially with the entireleaved forms of E. intortum. The entirely
23
free spathe margins and different arrangement of the staminodes in Eminium
should readily separate them. In the sterile
state, however, B. straussii is quite similar
to B. bovei, and they are often confused in
herbaria.
12. Biarum syriacum (Spreng.) H.Riedl
in Aroideana 3(1): 19 (1980). Type:
(SYRIA) prope Aleppo, Russell s.n.
(holotype BM). Arum syriacum
Spreng., Syst. Veg. 3: 768 (1826).
B. gramineum Banks & Sol. in Russell,
Nat. Hist. Aleppo ed.2, 2: 264 (1794),
nom. illeg. Type as for B. syriacum
(Spreng.) H.Riedi. B. gramineum
(Banks & Sol.) Eig in ]ourn. Bot.
Lond. 75: 189 (1937), nom. illeg. et superjl.
B. russellianum Schott, Prodr. Syst. Aroid.
63 (1860), nom. illeg et superjl.
Tuber dorso-ventrally compressed-discoid, 2-3.5 X 1.5-2 cm, not offsetting (?),
dark brown. Leaves 12-25, proteranthous, long-petiolate, the bases encased
by 3 to 6 ligulate, 6-14 X 1.5-2 cm, subfleshy, later papery, whitish cataphylls and
2 to 3 ligulate 9-13 X 1-1.5 cm, fibrous
brown cataphylls; petiole 6-15 cm X 1-3
mm, abaxial surface channeled distally,
expanded into a membranous wing proximally, mid to dark green; leaf lamina linear to linear-elliptic, the first few leaves
emerging spathulate, 6-14 cm X 2.5-4(14) mm, apex acute, base long-decurrent
to cuneate, 3-5 primary lateral veins per
side, margins smooth to undulate, lamina
mid- to dark green, rarely somewhat glaucous abaxially. Inflorescence appearing
in spring; peduncle 3.5-6 cm X 2-3 mm,
dirty white, emerging from amidst the foliage. Spathe 1l.5-18 cm long; spathe
limb elliptic to lanceolate-elliptic, 9-15 X
l.8-3.5 cm, exterior green, heavily stained
deep purple, interior deep purple; spathe
tube oblong, moderately inflated, 4-5.5 X
l.5-2.5 cm, margins fully connate, exterior
dirty white, stained purple around the upper margins, interior white, stained midpurple distally. Spadix sub-equal to just
exceeding the spathe limb, 15.5-18 cm in
24
AROIDEANA, Vol. 29
total length; spadix appendix fusiform,
sub-sessile to shortly stipitate, 8-15 cm X
6-9 mm, deep purple. Staminate jlowers in a zone 9-14 mm long, 5-6 mm
diam., anthers cream. Interstice 2-3.2 cm
.x 2-3 mm, dark cream. Staminodes arranged on the lower half of the interstice,
moderately dense; bases barely to not
swollen, dark cream, filaments filiform,
spreading to erect, 3-6 mm long, the longest nearest to the pistillate flowers,
cream. PistiUatejlowers in a hemispherical cluster 1-2 X 4-5 mm; ovaries oblong,
ca. 1-2 X 0.5 mm, dark cream, stigma subsessile, capitate, ca. 0.3 mm in diam. In.fructescence known only from immature
material, ca. 10 X 7 mm; berries ca. 3 X
1.5 mm, whitish; mature seed not seen.
Chromosome number not recorded.
Distribution-NW Syria.
Ecology-Bare earth plains in limestone-derived red clays. Alt. 150-300 m.
Etymology-From Syria, alluding to the
country of origin of the type and all other
material.
The confused nomenclatural history of
this species was discussed by Riedl
(1980a). When Banks and Solander published the name Arum gramineum (Banks
& Solander, 1794), based on a Russell collection in the British Museum, they appear
to have been unaware of Lamarck's Arum
gramineum (1789). Schott (1860) proposed B. russellianum as a nomen novum
for the Banks & Solander species, since
the epithet gramineum was unavailable
under either Arum or Biarum. However,
Schott overlooked that Sprengel (1826)
had proposed the name A. syriacum for
the Russell material and that he should
have adopted Sprengel's earlier name in
Biarum.
Very little material of B. syriacum has
been collected. The five sheets of material
in Paris are all from the Aleppo area of
northern Syria, one being the 'type' of
Schott's B. russellianum. The only other
material I have been able to trace is Russell's gathering, the type of Arum syriacum, in the British Museum and four con-
/
temporary gatherings in Geneva. According to the notes on the various sheets it
appears that B. syriacum is quite abundant, especially in the north of Syria.
If the unusual flowering period is ignored, B. syriacum would appear, on the
basis of floral morphology, to be related
to B. bovei and B. kotschyi. The rather oblong spathe tube and the arrangement of
the staminodes are similar to those of B.
bovei. However, the foliage of B. syriacum
is quite different and, in fact, no other species of Subgen. Ischarum has similar
leaves.
Reidl (1980) stated that the spathe tube
margins in B. syriacum were free for more
than half their length. I suspect that this
mistake arose due to a tracing of the "holotype" of B. russellianum that is present
in the Herbarium at Kew and which depicts the spathe tube with free margins.
Examination of the specimen in Paris that
was used for the tracing reveals that the
"free" margins are actually the result of
creasing of the inflated spathe tube during
drying; the margins are fully connate.
13. Biarum carratracense (Haenseler)
Font Quer in Bull. Inst. Catal. Hist.
Nat. 26: 53 (1926); Talavera in Lagascalia 6(2): 290-93 t.1, C, C1 (1976);
Talavera, Valdes & Galiano, Fl. Vasco
Andal. Occ. 3: 210 (1987). Arum, carratracense Haenseler in Bot. Zeit. 4:
313 (1846). Type: SPAIN, in agris cultis montuosisque ad Carratraca, jam
Sierra de Agua dictis, 18 November
1839, Haenseler s.n. (holotype G).
Biarum haenseleriWillk., Bot. Zeit. 5:
49 (1847), nom. illeg. et superfl. Type:
based on the same type as B. carratracense (Haenseler) Font Quer. Ischarum haenseleri (Willk.) Schott,
Syn. Aroid. 8 (1856), nom. illeg. B.
bovei Blume ssp. haenseleri (Willk.)
Engler in A. & C. DC., Monog. Phanerog. 2: 578 (1879), nom. illeg.
B. intermedium Amo, Fl. Fan. Penin. Iber.
1: 394 (1847). Type: SPAIN, crece en
las Alpujarras, cerca Orgiva y tembien
en Sierra Elvira, provincia de Granada, Amo s.n. (holotype not traced).
25
PETER C. BOYCE, 2006
B. tenuifolium (L.) Schott var. latifolium
Lange, Pugillus 81 (1860) [" latifolia"]'
Type: SPAIN, Sierra Elvira prope Granada, Lange 147 (lectotype C, selected here; isolectotypes C, G, P). There
are five sheets of this taxon present in
the Copenhagen herbarium, representing many individual plants. The
best preserved and most complete
sheet has been chosen as the lectotype.
Tuber slightly dorso-ventrally compressed discoid to slightly compressed
globose, 1.5-3 X 1-2 cm, mid-brown.
Leaves 3-7, hysteranthous, very rarely
synanthous, bases encased by three, 4.712 X 1-2 cm, lanceolate, sub-fleshy, later
papery, cataphylls, these pale greenish
white to dull green, turning pale greyish
yellow on drying; petiole 8-15 cm X 2--4
mm, adaxial surface channeled with slightly winged margins distally, expanded basally into a papery wing; leaf lamina elliptic, elliptic-ovate or oblanceolate, 5-12(15) X 2-3 cm, ca. 6 primary lateral veins
per side, margins smooth, lamina rather
dull mid-green. Inflorescence appearing
in late autumn, 11-17.5 cm long, strongly
foetid of horse dung and carrion; peduncle 3-7 cm X 2-3 mm. Spathe 10-17.5 cm
long; spathe limb lanceolate, 8-13.5 X 12 cm, margins smooth, exterior :±: green
with much rich purple staining, especially
towards the margin, becoming paler towards the base near soil level; spathe tube
elliptic-cylindric, slightly inflated, 2--4 X
1.5-2 cm, margins connate for ca. 1,2 their
length, exterior off-white beneath soil level, exposed portion rich purple, interior
white, stained purple at the base and towards the opening. Spadix 10-17 cm
long; spadix appendix slender-fusiform,
8-13 cm X 1.4-3.5 mm, deep brown-purple. Staminate flowers in a zone 10-15
X 3-5 mm diam., anthers cream, stained
purple in the region of the connective. Interstice 1-1.5 cm X 2-2.5 mm, purple.
Staminodes few, scattered over the lower
1 cm of the interstice and directly above
the pistillate flowers. PistiUateflowers in
a hemispherical cluster ca. 5-7 X 6 mm,
ovaries ovoid, 0.5-1 mm in diam., purple,
style ca. 0.5 mm X 0.25, purple, stigma
sub-capitate, ca. 0.33 mm in diam., pale
purple. Inj'ructescence a gJobose cluster
of ca. 50 berries 3-3.5 cm in diam.; berries
globose, ca. 8-9 mm in diam., pale purple;
seeds globose, ca. 5-5.5 X 5 mm, testa
strongly reticulate, mid-brown. 2n = 98
(Fernandez Piqueras & Ruiz Rejon 1976 &
Palomeque Messia & Ruiz Rejon 1976), ca.
96 (Talavera 1976), 36 (Fernandez Casas et
aI., 1978).
Distribution-SW Spain.
Ecology-Dry mountain pastures, field
margins, track sides in limestone-derived
red clay soils. Alt. 300-1,100 m.
Etymology-Originating from Carratraca
in southwestern Spain, the town closest to
the type locality.
Biarum carratracense has been associated with B. bovei and B. tenuifolium but
is quite clearly distinguishable from either.
The oblong, slightly inflated spathe tube
with the margins connate for over half
their length and the fusiform spadix appendix suggest an affinity to B. kotschyi
and B. jraasianum, although B. carratracense is geographically isolated from either species. Vegetatively B. carratracense
would appear to be closest to B. kotschyi,
which has a similar lanceolate-elliptic leaf
lamina. However, B. kotschyi has the petioles free to the ground whereas in B. carratracense the petiole bases are often imbricated to form a weak pseudostem. Further, the staminodes are far fewer in B.
carratracense than in B. kotschyi; some
material of the Spanish taxon almost lacks
pistillodes except for a couple of vestigial
bristles on the upper part of the interstice.
14. Biarum kotschyi (Schott) B.Mathew
ex H.Riedl in Aroideana 3(1): 28
(1980) ("kotshcyf'). Ischarum kotschyi Schott, Syn. Aroid. 7 (1856).
Type: (TURKEY?) Eastern Lebanon,
Kotschy s.n. (holotype W destroyed;
lectotype W, selected here). Schott's
leones Aroideae no. 2151. There are
four leones present in Vienna (nos
26
AROIDEANA, Vol. 29
2150-2154) and a single reliquiae (no.
366). The leone selected as the lectotype is by far the most informative
plate.
Tuber slightly dorso-ventrally compressed-discoid to ± globose-discoid, 2-5
cm in diam., 1-1.2 cm thick, pale brown.
Leaves 5-15, hysteranthous, distinctly
petiolate, bases encased by 3 to 6, 6-9 X
1.5-2 cm elliptic lanceolate, sub-fleshy, later papery, cataphylls, these pale greenish
white at first, pale straw-yellow on drying;
petiole 5-9(-22) cm X 4-5 mm, adaxial
surface channeled and winged distally,
slightly expanded into a wing proximally;
leaf lamina elliptic to lanceolate-elliptic,
rarely oblanceolate, 6-9.5 X 2.5-4 cm,
apex sub-acute, base short-decurrent, ca.
5-6 primary lateral veins on each side,
margins smooth, lamina dull mid-green.
Inflorescence appearing in late autumn,
smelling pungently of horse dung; peduncle 3-6(-17.5) cm X 4-6 mm, pale green
to dirty white. Spathe 8.5-10(-15) cm
long; spathe limb lanceolate, 5.5-11 X 23.5 cm, apex acute, exterior pale dirty
green with numerous small pale purple
speckles and some purple staining, especially towards the margin, interior deep
purple; spathe tube oblong, slightly inflated, 3-4.5 X 1.5-2.5 cm, margins connate
for ca. 14 of their length, exterior white
with some purple staining basally and apically, interior white stained purple at the
base and apex. Spadix ca. 3,4 as long as,
to just exceeding the spathe limb, 8-13 cm
long; appendix ± sessile, slender-fusiform, 5-9 cm X 2.5-4.5 mm wide, deep
purple. Staminate flowers in a zone 912 X 3-4.5 mm wide, anthers cream,
tinged purple apically. Interstice 1.75-2.5
cm X 2 mm wide, purple. Staminodes arranged at the base of the interstice, directly
above the pistillate flowers, filaments 3-6
mm long, purple. Pistillate flowers in a
hemispherical cluster 12-15 X 7-9 mm;
ovaries bottle-shaped, 1-1.5 X 0.75-1 mm,
cream, stained purple, style ca. 0.5 X 0.15
mm, purple, stigma capitate, ca. 0.33 mm
in diam., purple. Infructescence globose,
ca. 3-4.5 cm in diam., consisting of ca. 40
berries; berries oblong-ovoid, ca. 5- X 45 mm wide, pale purple with darker staining when ripe; seed globose, ca. 4 mm in
diam., strongly reticulate. Chromosome
number not recorded.
Distribution-SE Turkey, provinces of
Bitlis, Diyarbakir, Urfa and Gaziantep and
Maras.
Ecology-Dry clay-loam hillslopes,
stony plateaux, long-fallow fields. Alt.
600-2000 m.
Etymology-Named after the Austrian
botanist Theodor Kotschy.
Due to Engler's misidentification of the
type material as B. bovei, B. kotschyi has
only recently been accepted as being a
distinct species. Biarum kotschyi could be
regarded as intermediate between B. bovei
and B. pyrami on the basis of the shape
of the spathe tube, spadix appendix, staminodes and foliage.
Biarum kotschyi and B. carratracense
are also similar. The rather slender spathe
limb, fusiform spadix appendix and the arrangement of the staminodes is similar in
both species. They can be readily distinguished by the degree of connation of the
spathe tube margins (three quarters free in
B. kotschyi, half free in B. carratracense)
and the distinct geographical distributions.
Biarum kotschyi is a common species in
parts of southeastern Turkey and a search
of a dry hillside will usually reveal this
species. In light of this it might appear
strange that Mill (1984) omitted B. kotschyi
from his Flora of Turkey account were it
not for the fact that almost all herbarium
material of this species has hitherto been
annotated as B. bovei.
Once again, Schott's citation of the type
locality must be regarded as suspect. Biarum kotschyi has never been found in Lebanon or Syria and it appears to be restricted to a few provinces in Turkey. It is most
likely that the type of B. kotschyi originated in southeastern Turkey.
15. Biarum fraasianum (Schott) Nyman, Syl!. F!. Europ. supp!.: 68 (1865);
Brown in J. Linn. Soc. 18: 254 (1881);
27
PETER C. BOYCE, 2006
Engler in Das Pflanzenr. 73 (IV.23F):
139 (1920).
Ischarum fraasianum Schott in Oesterr.
Bot. Zeit. 9: 98 (1859). Type: GREECE,
in campis Thebaicis, Fraas s.n. (holotype B destroyed; lectotype W, selected here Schott's leones Aroideae
no. 2147) There are two illustrations
of this species in Vienna (Reliquiae
367 & leones 2147). They are essentially the same illustration arranged in
different ways. Nevertheless, the leones 2147 is annotated 'Arum 4 in
camp is Thebaicis (Gr.) comm. & coli.
Fraas! Herb. Sartorii'
Biarum fraasianum (Schott) N.E. Br. in
Joum. Linn. Soc. 18: 254 (1881),
comb. superfl.
Tuber compressed discoid 4 X 2 cm,
offsetting little, pale brown, older tubers
with conspicuous, concentric brown rings.
Leaves 13-20 cm, hysteranthous, longpetiolate, petioles clasping basally, enclosed by few, ca. 3 cm X 2-4 mm lineartriangular to lanceolate pale brownishcream papery cataphylls; petiole 7-9 cm X
ca. 2 mm; leaf lamina oblong to oblanceolate, 6-9 X 1.5-4 cm, apex obtuse to subacute, base short to long decurrent, ca. 6
primary lateral veins per side, margins
smooth, lamina mid-green. Inflorescence
appearing in spring, smelling strongly of
overripe fruit; peduncle ca. 4-6 cm X 5-6
mm, swollen apically in fruit to ca. 1 cm,
clothed, together with the base of the
spathe, by several 6-10 X 1.5-2 cm broadly linear cataphylls. Spathe 13-24 cm
long; spathe limb ovate-Ianceolate to oblong, 10-20 X 4-6 cm, shortly cuspidate,
exterior greenish purple, interior dark
brownish purple, upper part of spathe
tinged green; spathe tube subcylindric, 34 X 1.5-2.5 cm, margins fused for their
length, exterior whitish, tinged apically
green and purplish brown, interior white
above, deep purple below. Spadix approximately half as long as the spathe
limb, 7.5-12 cm long; appendix briefly
stipitate, fusiform, 4.5-10 cm X 4-7 mm,
greenish purple-brown, stipe and base of
*
appendix paler. Staminate flowers in a
zone ca. 15 mm long and 4 mm diam.,
cream, slight stained purple towards the
tips. Interstice 16-25 mm long and 2-3
mm diam., white. Staminodes sparse, arranged mostly directly above the pistillate
flowers, but with one or two scattered on
the upper portion of the interstice; filaments slender, ca. 4 mm long, white. Pistillate flowers in a hemispherical cluster
10-13 X 3-6 mm; ovaries bottle-shaped,
ca. 1.5 X 0.5 mm wide, green, style ca. 0.5
X 0.15 mm, flushed purple, stigma globose, ca. 0.25 mm in diam. Infructescence hemispherical, 2.5 X 1 cm, consisting of ca. 30 fruits, berries pyriform-globose, ca. 4 X 4 mm, pale silvery purple,
stylar portion purple, stigmatic remains
prominent, ca. 1 X 0.5 mm; seeds globose,
ca. 3 mm in diam., pale brown, somewhat
reticulate. 2n = 36 (Popova & Ceschmedjiev 1978) but see note below.
Distribution-Long thought to be restricted to the type locality, where extensive urban and agricultural development
has almost certainly extinguished most
populations, B. fraasianum has recently
been recollected in the Pelopponense by
amateur bulb enthusiast Mike Salmon.
Ecology-Stony red clay soils derived
from limestone. Alt. 50-260 m.
Etymology-The species is named in
honour of Carl Nicholaus Fraas, collector
of the material used to describe the species.
Until recently Biarum fraasianum was
a species for which very little material was
available for study. The Fraas' type collection was at Berlin and a search of the herbarium did not locate the material and it
must be assumed it was among the material destroyed in WW2. Only two other
contemporary herbarium collections of B.
fraasianum exist. One of these, held at
the Natural History Museum (BM), consists
of a tuber, leaves and a semi-mature infructescence. The other, in Patras University Herbarium (UPA), is a flowering specimen. The type material was used to pre-
AROIDEANA, Vol. 29
28
pare two illustrations held in the Naturhistorisches Museum, Vienna (W).
The recent recollection of B. fraasianum has revealed numerous hitherto unknown characteristics, including spring
flowering and a sweet smell at anthesis.
The latter is particularly uncommon in
Biarum, shared only with B. davisii and
B. marmarisense.
The report of a chromosome count for
= 32 (Popova &
Ceschmedjiev 1978) must be regarded as
probably based on a misidentified plant.
Nevertheless the Popova and Ceschmedjiev count is intriguing in that it is different
from any species that might conceivably
be misidentified as B. fraasianum.
Biarum fraasianum is most similar to B.
bovei, B. kotschyi and B. carratracense,
particularly in the degree of spathe tube
inflation and the fusiform spadix appendix. The arrangement of the staminodes in
B.fraasianum is closer to that found in B.
disparthan to that of B. carratracenseand
B. kotschyi but its geographical distribution and the greater overall similarity to B.
kotschyi leads me suspect that the closest
relationship lies with the latter species.
tion, leones Aroideae no. 2161 in Vienna
is ideal since it is annotated with the same
data as the isotypic fragments in Geneva.
As pointed out by Mill (1984) the locality
cited in Schott's Prodromus Systematis
Aroidearum (1860) is almost certainly
wrong. The fragmentary isotype specimens in Geneva are annotated 'Messis et
Scheih Meran' and do not carry the data
cited by Schott in the protologue.
B. fraasianum of 2n
16. Biarum pyrami (Schott) Engler in A.
& c. DC., Monog. Phanerog. 2: 576
(1879) & in Engler, Pflanzenr.
73CIY.23F): 139 (1920); Koach & Feinbrun in Feinbrun, FI. Palaestina 4: 337
(1984); Koach in Rotem 26 t.l5,16
(1988). Type: Juxta arcem Semeramidis [plantae ad Pyramum (Ceyhan river) in monte Nur lectae: inter Messis
(Misis) et castellum Scheih Meran
(Yilankale), 60 mJ, Kotschy s.n. (holotype W destroyed, isotypes G (sterile fragments».
Since isotypes exist, albeit fragmentary,
it is not possible to select a lectotype for
Ischarum pyrami other than from this material (Art. 7.4, ICBN Code). This, due to
the condition of the specimens, is a pointless exercise. Nevertheless, it is useful to
have a fixed point on which to base an
interpretation of the name pyrami and for
this purpose I epitypify the Schott Illustra-
Ischarum pyrami Schott, Prodr. Syst. Aroid. 66 (1860).
l. nobile Schott, Prodr. Syst. Aroid. 66
(1860). Type (Turkey) juxta arcem
Semiramidis, cult. Schoenbrunn,
Kotschy s.n. (holotype W destroyed; lectotype W (chosen here)
(Schott's leones Aroideae no.
2157». Of the plates present in Vienna, the one chosen bears the
same data as given in the protologue and is thus the logical choice
for the lectotype. The same confusion with localities detailed under
B. pyrami apply to l. nobile.
Tuber globose-discoid 2-4 X 2-2.5 cm,
mid-brown. Leaves hysteranthous (var.
pyramt) or synanthous (var. serotinum),
long-petiolate, encased basally by numerous (2-)7-13 X 0.5-2 cm elliptic-lanceolate sub-fleshy, later papery, cataphylls,
these pale greenish, occasionally with faint
purple spots externally at first, later pale
straw-yellow; petiole 8-16 cm X 2.5-4.5
mm, margins slightly winged, adaxial surface channeled distally, expanded proximally into broad membranous wing, midgreen; leaf lamina oblong-elliptic to ovateelliptic, 6-18 X 3-8 cm, apex subacute to
slightly rounded, base briefly decurrent to
rounded, occasionally sub-truncate, ca. 910 primary lateral veins per side, margins
flat, occasionally crispulate, leaf lamina
deep green, with black-purple or green
bullae (var. pyramt) or smooth (var. serotinum). InJ10rescence appearing in late
autumn (var. pyramt) or early winter (var.
serotinum), smelling strongly foetid of cattle dung and carrion; peduncle 4-7 cm X
ｾ＠
mm, off white. Spathe 10-25 cm long;
29
PETER C. BOYCE, 2006
spathe limb lanceolate, (7-) 15-18.5 X 23.5 cm, long-acuminate, erect at first but
soon reflexing and coiling, exterior midgreen stained and spotted purple-brown,
especially towards the margins (var. pyramt) or unspotted (var. serotinum), interior
deep purple brown, sometimes with paler
speckling along the middle and towards
the apex; spathe tube globose, ventricose,
3-4 X 2.5-3.5 cm, margins connate for 14
of their length, exterior whitish below
ground, purple-brown above, interior dark
purple. Spadix equalling to slightly exceeding the spathe limb, ｾ＠
cm long; spadix appendix cylindric-fusiform 5.7-22 cm
X (3-) 5-9 mm, deep purple-brown. Staminatejlowers in a zone 10-15 mm long
X 5-6 mm diam., anthers purple. Interstice 14-16 X 2-3.5 mm, deep purple,
rarely dull creamy yellow. Pistillodes in
a zone 5-10 mm long at the base of the
interstice, sparse; bases slightly swollen,
purple; bristles filiform, flattened distally,
5-7 mm long, cream, sometimes flushed
deep purple. Pistillate jlowers in a
hemispherical cluster ca. 10 X 3-5 mm
high, ovaries bottle-shaped, 1.5-2 X 11.25 mm, deep purple, paler below, style
ca. 1.5 X 0.33 mm, purple, stigma subcapitate, ca. 0.50 mm in diam., white. Infructescence globose, 2-3 cm in diam.,
consisting of ca. 50 berries; berries oblongpyriform, 7-10 X 5-6 mm, silvery-lilac,
stained purple proximally, seed sub-turbinate, 5-6 X 4-5 mm wide, testa reticulate,
dark brown.
a. var. pyrami
Leaves hysteranthous, bullate. lriflorescence appearing in late autumn.
Spathe limb exterior mid-green, stained
and spotted purple-brown. Chromosome
number not recorded.
Distribution-SW and central S Turkey,
Syria, Lebanon, Jordan and Israel.
Ecology-Garigue or low Maquis on red
clay soils derived from limestone, limestone hill slopes, amongst loose stones or
in disused fields. Alt. 30-450 m.
Etymology-Named after Pyramus, the
ancient name for the Ceyhan River in
southern Turkey, the type locality.
h. var. serotinum Koach
& Feinbrun in
Feinbrun, Fl. Palaestina 4: 398 (1986).
Type: ISRAEL, Golan, near Katsrin,
hills, basalt rocks and soil, 22 December 1979, Heiman 79 (holotype HUJ).
Leaves synanthous, lacking bullae. Inflorescence appearing in late winter.
Spathe limb exterior concolorous midgreen. Chromosome number not recorded.
Distribution-Israel.
Ecology-Hillslopes in soils derived
from basalt rocks. Alt. 60 m.
Etymology-From the Latin serus (late),
in allusion to the winter flowering of this
variety.
This large, showy species is perhaps the
most readily distinguishable of the autumn
flowering Biarum species due to the globose, strongly inflated, spathe tube, large
spathe and greatly attenuated spadix appendix. The bullate leaves of the typical
variety are also unusual in the genus and
provide a ready means of identifying B.
pyrami in Turkey, where no other species
shares this character. In Turkey, B. pyrami,
B. bovei and B. kotschyi, form an apparently closely related group of species but
they are all readily distinguishable from
one another.
Variety serotinum is maintained here
with some reluctance. There is no denying
that some Israeli material of B. pyrami is
rather distinct from that of Turkish and
Syrian origin. The lack of leaf bullae, used
as one of the distinguishing characters for
var. serotinum, is generally a stable character, although the Davis 3844 collection
from Israel cited above lacks leaf bullae
although phenologically and ecologically
it is clearly referable to var. pyrami, and
the two varieties are apparently ecologically different. The floral characteristics
used to distinguish var. serotinum seem
less reliable, especially with regard to the
flowering period on which Koach and
30
AROIDEANA, Vol. 29
Feinbrun laid great emphasis. Elsewhere
in Biarnm, phenology has proved to be
variable, with some species (e.g. B. straussii and B. syriaeum) displaying a rigid
flowering cycle while others (e.g. B. tenuifolium and B. arnndanum) have a wide
degree of variability even in single populations.
The plant described as Isebarnm nobile
by Schott (1860) has been suggested by
some authors (e.g. Mill, 1984) as possibly
representing a distinct species. Although
the type material is no longer extant, the
painting in Vienna (Schott's /cones Aroideae no. 2155) that serves as the type of
l. nobile suggests that it is conspecific with
B. pyrami. The illustrations of l. nobile and
I. pyrami (Schott's leones Aroideae no.
2161) are very similar. Apart from the
slightly less inflated spathe tube and rather
sparser pistillodes, the two illustrations almost certainly depict the same species.
17. Biarum mendax P.C. Boyce, Aroideana 22: 90 (1999). Type: SPAIN.
Badajoz: Between Hobr6n and Solana
des los Barros, 15 October 1976, Cabezudo et al. 2201/76 (holotype SEV
25005; isotypes G, K).
Tuber globose-discoid, 4 X 3.5 cm,
mid-brown. Leaves not seen. Inflorescence appearing in early autumn smelling
very strongly foetid of cattle dung and carrion; peduncle 5-15 cm X 3-4 mm, off
white, clothed by few to many, 5-13 cm
X 7-15 mm papery, pale yellowish white
cataphylls. Spathe 16-24 cm long; spathe
limb lanceolate, 18-20 cm X 15-18 mm,
apex long acuminate, exterior green ±
heavily blotched and stained purplebrown, interior deep purple-brown;
spathe tube globose, strongly inflated, 45 X ca. 3 cm wide, margins fused for their
entire length, exterior pale green stained
purple-brown especially towards the
opening, interior off-white distally, deep
purple proximally. Spadix sub-equal to
exceeding the spathe limb, 16-21 cm long;
spadix appendix slender fusiform-cylindric, 14-16 cm X 6-8 mm, deep purplebrown. Staminate flowers in a zone 11-
13 mm long and 6-9 mm diam., deep purple. Interstice 23-25 X 5-6 mm, deep
purple. Staminodes situated at the base
of the interstice, few, filiform, 5-14 mm
long, deep purple. PistiUate flowers in
a hemispherical cluster 4-5 X 7-13 mm
wide; ovary 2-3 mm long, cream; style 1
X 0.33 mm, purple, stigma capitate, ca. 0.5
mm in diam., purple.ltifructescence not
seen. Chromosome number not recorded.
Distribution-SW Spain.
Ecology-Rocky hill slopes on limestone-derived red clays. Alt. 50-75 m.
Etymology-From the Latin mendax,
deceitful, in allusion to the similarity in the
dried state between the new species, B.
bovei and B. dispar which has resulted in
it being hitherto overlooked.
In herbaria, B. mendax has been assigned to B. bovei Blume, or B. dispar
(Schott) Talavera, on the basis of its overall
similarity to them. However, B. mendax is
readily separable by its greater size, exceeding that attained by B. pyrami. Indeed, B. mendax bears the largest inflorescence yet recorded in Biarnm. From all
three species it can be distinguished by the
completely fused spathe tube. Biarnm
mendax belongs to a group of species defined by spadices bearing sterile flowers
only between the male and female flower
zones, and in the inflated lower spathe.
18. Biarum auraniticum Mouterde,
Nouv. FI. du Liban et Syrie 1: 193
(1966). Type: SYRIA, tel pres du Zraikiye, au nord de Sanamein, December 1954, Pabot 326 (holotype G!; isotypes G!, K! (photograph».
[Biarnm luteum Pabot, in sebed. nom.
nudJ
Tuber slightly dorso-ventrally compressed discoid, 2-3 X 1.5-2 cm, rnidbrown. Leaves 2-3, hysteranthous, shortto rather long-petiolate, bases encased by
2 to 3, 6-12 cm X 7.5-20 mm, lanceolate,
papery, cataphylls, these pale straw-yellow on drying; petiole 7-12 cm X 2-10
mm, adaxial surface slightly channeled
distally, outer petioles expanded proxi-
PETER C. BOYCE, 2006
mally into a broad membranaceous wing,
inner :!:: the same width for their whole
length, mid-green, paler below; leaf lamina elliptic, 4-8 cm long, 2-4.5 cm wide,
apex rounded to sub-acute, base decurrent, 6-8 primary lateral veins per side,
margins smooth, lamina mid-green. Inflorescence emerging in winter; peduncle 25 cm X 3-4 mm diam., off-white, intensely
foul smelling of horse dung. Spathe 8.517 cm long; spathe limb lanceolate to narrowly lanceolate, 9-13 X 1-3 cm, apex
acute to rather acuminate, exterior greenish white, interior paler; spathe tube narrowly cylindric, wider at the mouth, slightly inflated in the mid-region, 2-3 cm X 8-13 mm, margins connate for ca. ¥.J their
length, exterior pale green, interior pale
green distally, deep purple proximally.
Spadix sub-equal to shorter than the
spathe limb, 7.5-14 cm long; spadix appendix slender fusiform to fusiform, 5.510.2 cm X 2-7 mm, sub-sessile to briefly
stipitate, yellow. Staminate flowers in a
zone 18--22mm long and 3-3.25 mm diam,
cream. Interstice 8--11 X ca. 1.5 mm in
wide, cream. Staminodes absent. Pistillate flowers in a hemispherical cluster 35 X 6-8 mm; ovaries globose, 1.5 X 1.5
mm, cream, style ca. 1 X 1.25 mm, purple,
stigma ca. 0.5 mm in diam., purple. InJructescence known only from immature
material, consisting of ca. 35 globose berries in a globose cluster; berries sub-globose, 2-3 X 1.5-3 mm, dull purple. Seed
(immature) spherical, ca. 2 mm in diam.,
testa reticulate, mid-brown. Chromosome
number not recorded.
Distribution-SW Syria, known only
from the type locality.
Ecology-Open hillslopes and fields in
stiff clay-like volcanic soils.
Etymology-Named after the ancient region of Aurantis in southern Syria. The
town of Sanamein, close to the type locality, being in this area.
Biarum auraniticum is unique in possessing a white spathe limb. There is no
doubt that this is a most singular species,
not only from the inflorescence coloration
31
but also because of the remarkably wide
bottle-shaped ovaries. The odd inflorescence coloration and unusual appearance
of the ovaries make it difficult to envisage
a close relationship. The lack of pistillodes, the ovate-elliptic, long-petiolate
leaves and slightly inflated spathe tube
support a link to B. hovei or B. kotschyi.
19. Biarum ditschianum Bogner &
Boyce in Willdenowia 18(2): 409
(1989). Type: TURKEY, Antalya, Xanthos hill; in holes and crevices in
limestone, 30 m, 24 April 1988, Koenen Bonn 22592 (holotype K; isotypes B, BONN, K, M).
Tuber depressed-globular to sub-globular, 2.5-3 X 1. 5-2 cm, light brown. Roots
1.4-2 mm in diam. Leaves 2-3, rarely up
to 5, hysteranthous, long petiolate, bases
partly encased by several 5-10 cm long,
membranaceous, whitish cataphylls; petiole 8--20 cm X 1.5-3 mm diam., channelled, laterally compressed, mid-green,
sometimes reddish tinged distally; leaf
lamina oblanceolate early in the season,
subsequent leaves linear, narrowly-elliptic
or lanceolate, 6-15(-20) X 0.6-3(-3.5) cm,
base cuneate, decurrent, apex acute to obtuse, 4-7 primary lateral veins on each
side, leaf lamina mid-green, veins paler.
Inflorescence appearing in spring, smelling strongly foetid, base enclosed by 5--6
cataphylls, these at first membranaceous,
whitish, soon withering to become papery
and brown, 1.5--6(-7) X ca. 1.5 cm, the
longest equalling the spathe; peduncle
subterranean, 2-5 cm X 3.5-5 mm diam.,
whitish. Spathe 4-5 cm long, spathe limb
much reduced, 1.8--2 X ca. 2 cm diam.,
sub-triangular, terminating in a ca. 2 mm
mucro, exterior greenish to light green,
sometimes with a reddish tinge, interior
purplish-red, spathe tube ca. 3 X 1.8--2
cm, margins connate proximally for ca. 2.5
cm, exterior whitish, sometimes slightly
reddish tinged, interior reddish purple.
Spadix exceeding the spathe, 7-8 cm
long; spadix appendix elongate-conoid to
somewhat sub-cylindric, shortly stipitate,
4-4.5 cm X 7-11 mm, apex obtuse, base
32
AROIDEANA, Vol. 29
rounded, the basal 7-10 mm furnished
with reflexed, filiform, acuminate, 1-2.5 X
0.1-0.25 mm, transparent white 'hairs', appendix dark yellow except for the reddish
purple basal 7-10 mm; stipe 5-6 X 3-4
mm, cream. Staminate flowers arranged
in an oblong zone 5-6 ,x 7-9mm, stamens
sub-sessile, yellow proximally, purple-red
distally, occasionally entirely yellow. Pollen extruded in strands. Interstice 1.3-3.5
cm X 2.5-3.5(-4) mm diam., light purple,
fading to creamy white apically purple.
Staminodes absent. PistiUate flowers
arranged in a 2-2.5 mm high hemispherical cluster; ovary bottle-shaped, ca. 1 mm
in diam., purplish red, occasionally pale
cream, styles and stigmas curved outwards, style 1.2-1.3 X ca. 0.4 m, purplish,
colour intensifying towards the ovary, occasionally cream, stigma sub-capitate, 0.50.6 mm in diam., yellow. Infructescence
depressed-globular, 3-3.5 X ca. 2 cm, consisting of ca. 50 berries; berries obovoid,
6-10 X 4-7 mm, whitish, sometimes with
a very slight reddish tinge proximally, seed
obovoid, 5.5-7.5 X 4-5.5 mm; testa with
the upper part very slightly irregular-reticulate, lower part smooth. 2n = 26 (Petersen 1989).
Distribution-SW Turkey.
Ecology-Low to middle-high garigue,
partly loam-filled chimneys or crevices in
limestone. Alt. 30-120 m.
Etymology-Named in honour of Friedrich Ditsch, the original discoverer of the
species.
Biarum ditscbianum was first collected
in spring 1987 in southwestern Turkey by
Friedrich Ditsch, a student at Bonn University, Germany. A single tuber was
grown on at Bonn by him and flowered in
May that year. Unfortunately the plant died
soon after flowering and no voucher specimen was preserved. However, colour
photographs of the plant were made. Attempts to recollect the species in October
1987 failed but in April 1988 Manfred Koenen of the Bonn Botanical Garden succeeded in recollecting it. Plants flowered
at Bonn in May 1988. Herbarium and spirit
specimens were made and used to prepare a description of the plant (Bogner &
Boyce, 1989).
Biarum ditscbianum has an extraordinary appearance in flower compared with
most other Biarum species. The spathe
limb is reduced to a narrow rim on the
spathe tube and the most notable feature
is the relatively massive, dark yellow spadix appendix. Two other Biarum species
have unusual inflorescences. Biarum davisii (Turrill, 1938; Boyce, 1987) has pinkish
white spathes, a purple spadix appendix
and produces a sweet, not foul, odour at
anthesis. Biarum auraniticum (Mouterde,
1966) has a greenish white spathe and a
yellow spadix; the odour produced by the
inflorescence is unknown. Both species
lack a zone of sterile flowers (staminodes)
on the interstice separating the male and
female flower zones, a feature also seen in
B. ditscbianum.
Perhaps the most unusual feature of B.
ditscbianum is the presence of hair-like
processes on the base of the spadix appendix. Such structures are otherwise unknown in the genus and are uncommon
in the Araceae. Their function is not clear,
although it is possible that they playa role
in pollination.
The inflorescence of B. ditscbianum is
also notable for being exceptionally foul
smelling and produces a powerful odour
of carrion and excrement which attracts
carrion flies. Experiments undertaken in
Bonn have shown that the spadix appendix absorbs UV light, suggesting that it
contrasts well against the surrounding
limestone in habitat. Carrion flies are
known to perceive mainly UV light and it
is possible that the appendix is the most
visible part of the inflorescence to this
type of fly.
20. Biarum davisii Turrill in The Gard.
Chron. ser.3, 104:437 (938); Rix, The
Bulb Book 185 (981). Type: CRETE,
above and north-east of Sphakia,
amid rocks and amongst semi-scree,
23 April 1938, Davis 114 (holotype K;
isotype E).
PETER C. BOYCE, 2006
Tuber globose to rather hemispheric<ll, 1.53 X 1-2.5 cm. Leaves 5-9, hysteranthous,
short petiolate, petiole bases encased by
numerous linear-lanceolate cataphylls,
these 3-4 cm X 7.5-10 mm; petiole 1.5-5
cm X 1-2 mm, pale to mid-green; leaf
lamina ovate to ovate-spathulate, elliptic
ovate or elliptic, 1.5-4 X 1-2 cm, apex obtuse to sub-acute, base cuneate, ca. 3-5
primary lateral veins per side, margins
smooth to somewhat crispulate, lamina
mid to dark green, very occasionally spotted silver-grey. Injlorescence appearing
in autumn, sweetly perfumed; peduncle
2-7 cm X 3-5 mm, dirty white, clothed
with numerous cataphylls. Spathe 5-6 cm
long; spathe limb narrowly lanceolate,
2.5-3 cm X 5-7 mm wide, strongly cucullate, the margins recurved, apex acute, interior and exterior pale greenish white
spotted and mottled pale purple; spathe
tube ellipsoid, 2-3 cm X 1-2 cm wide, the
margins fused for their entire length, interior and exterior pale greenish white :::':
mottled with pinkish brown, pale purple
or purple-brown, the rim of opening
stained brown or yellow. Spadix shorter
than to sub-equal to the spathe, 3-5 cm
long, appendix sessile, slender cylindric to
filiform, 3-4 cm X 0.5-2.5, dull reddish
brown, dark red to purple-red towards the
apex. Staminodes absent. Staminate
flowers in a cylindrical zone 8-10 X 2-3
mm diam., anthers cream. Interstice 5-11
X 0.5-2 mm, cream. Staminodes a few
scattered vestigial filaments may be found
on the interstice, usually above the pistillate flowers. Pistillate flowers in a hemispherical cluster 1-2.5 X 2-4 mm; ovaries
oblong-ovoid, 1-1.25 X 0.5 mm, pale
green, stigma sessile, sub-capitate, 0.25
mm in diam., pale green. Itifructescence
globose, 2-3 cm in diam., consisting of ca.
30 berries; berries 3.5-4 X 3-4 mm, dirty
white when ripe, seed ovoid, 3-4 mm in
diam., testa slightly reticulate, pale brown.
2n = 26 (Petersen 1989).
Distribution-Crete.
Ecology-Open, grazed scrub, disturbed habitats, road and track margins in
limestone-derived red clays and screes, of-
33
ten in clay-filled holes in limestone, limestone pavement. Alt. 0-1,050 m.
Etymology-Named for Peter Davis,
who collected the material used to describe the species.
Biarum davisii is an attractive species
which, until the discovery of B. ditschianum, ranked as the most unusual taxon in
the genus. Apart from its remarkably small
size, characteristics such as the basically
ovate foliage, deeply urceolate spathe
tube, pinkish brown spathe and sweet lilac-like smell when in blossom are all
unique in the genus.
Riedl's (1980b) suggestion that B. davisii
might eventually prove to be a subspecies
of B. olivieri is highly improbable. The sessile stigmas, densely aggregated staminate
flowers, rudimentalY spathe limb, ovate to
ovate-spathulate, long-petioledleaf blades
and unique inflorescence colour are clearly very different to the states found in B.
olivieri. Aside from these intrinsic floral
and vegetative differences, the species occur in fundamentally different environments. Biarum davisii is a plant of limestone screes and red clay pockets on calcareous hillsides. Biarum olivieri occurs
uniquely in consolidated sand and waterdeposited silt in flat fields (pers. observ.,
Koach 1988).
The inflorescence colour and smell suggests that B. davisii has a different pollination syndrome from the remainder of
the genus and Biarum davisii is visited by
bees on Crete (Boyce, pers. observ.; Akeroyd, 1988).
Biarum davisii is widespread but rather
local on Crete. Where it occurs it is often
abundant, forming extensive colonies,
however, the small size together with the
fleeting appearance of the inflorescences
means that B. davisii is much overlooked
and this has led to the belief that it is rare.
More recent observations, together with
data on herbarium sheets, suggest that this
is not the case and that it occurs in most
parts of the island. The Cretan populations
of B. davisii are remarkably uniform,
plants from opposite ends of the island be-
34
ing virtually indistinguishable morphologically.
21. Biarum marmarisense (P.c. Boyce)
P.c. Boyce stat. nov. Basionym: Biarum davisii marmarisense P.c. Boyce
in Aroideana 10(4):14 (1987) ("marｭ｡ｲｩｳ･ｮｾｬＧＩＮ＠
Type: TURKEY, Mugla,
Marmaris, Bozburun, Taslica Ki.)yli,
T.Baytop et al (holotype EGE (accession no. EGE 8796); isotypes E, K).
Tuber globose to rather hemispherical,
1.5-3 X 1-2.5 cm. Leaves 5-9, hysteranthous, long petiolate, petiole bases
encased by numerous linear-Ianceolate
cataphylls, these 3-9 cm X 7.5-15 mm;
petiole 7-11 cm X 1-2.5 mm, pale to midgreen; leaf lamina ovate to ovate-spathulate, elliptic ovate or elliptic, 1.5-6.5 X 13 cm, apex obtuse to sub-acute, base cuneate to slightly cordate, ca. 6-9 primary
lateral veins per side, margins smooth to
somewhat crispulate, lamina mid to dark
green. Inflorescence appearing in autumn, strongly and sweetly scented, peduncle 2-11 cm X 3-7 mm, dirty white,
clothed with numerous cataphylls. Spathe
7-11 cm long; spathe limb narrowly lanceolate, 3-5 cm X 5-10 mm wide, strongly
cucullate, the margins recurved, apex
acute, interior and exterior pale greenish
white spotted and mottled pale purple;
spathe tube ellipsoid, 2-4.5 cm X 1-2.25
cm wide, the margins fused for their entire
length, interior and exterior pale greenish
white ± mottled with pinkish purple, the
rim of opening stained purple-pink. Spadix shorter than to sub-equal to the
spathe, 3.5-9 cm long, appendix sessile,
slender cylindric to filiform, 3-7 cm X 0.52.5 mm, dull reddish brown, dark red to
purple-red towards the apex. Staminate
flowers in a cylindrical zone 15-16 X 23 mm diam., anthers cream. Interstice 511 X 0.5-2 mm, cream. Staminodes usually absent. Pistillate flowers in a hemispherical cluster 1-2.5 X 2-4 mm; ovaries
oblong-ovoid, 1-1.25 X 0.5 mm, pale
green, stigma sessile, sub-capitate, 0.25
mm in diam., pale green. Infructescence
globose, 2-3 cm in diam., consisting of ca.
AROIDEANA, Vol. 29
30 berries; berries 3.5-4 X 3-4 mm, dirty
white when ripe, seed ovoid, 3-4 mm in
diam., testa slightly reticulate, pale brown.
2n = 22, 24 (Gill, 1988).
Distribution-SW Turkey, Greece (Simi
Island.)
Ecology-Limestone garigue, frequently
in heavy red limestone-derived clays. Sealevel-ISO mas!.
Etymology-Coming from the Marmaris
Peninsula of southwestern Turkey.
Gill (1988) in an exhaustive study of B.
davisii concluded that there were sufficient grounds for the recognition of the
Simi population as a variety of B. marmarisense, using characters similar to
those used to distinguish B. marmarisense
from B. davisii. My own view is that more
research into the Turkish populations is required to evaluate the characters before
such a move is made.
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