Kuijt1967 Ixocactus

THE GENUS IXOCACTUS (LORANTHACEAE, S.S.
):
D E S C R I P T I O N OF ITS F I R S T SPECIES
JOB KUIJT
Kuijt, J. (University of British Columbia, Vancouver, Canada). The genus

lxocactus (Loranthaceae, s.s.): description of its first species. Brittonia 19: 62-
67. 1967.--The first species of lxocactus Rizzini, 1. hutchisonii Kuijt, is described
and illustrated. The genus is unique in the family in that it is squamate and lacks
inflorescences. Other remarkable features are the dense covering of spines,
tetracolpate pollen, the small number of pollen grains per anther, the small size
of the flower, and the occurrence of this parasite on stems and roots of other
mistletoes, lxocactus is believed to be related to Oryctanthus.
The mistletoe genus Ixocactus is at present a nebulous entity. Its only appearance
in botanical literature has been in Rizzini's (1952) generic treatment of Brazilian
mistletoes. Except for some incidental comments scattered through Rizzini's text,
Ixocactus appears only in his key (p. 118). The relevant statement in this key
therefore becomes the generic diagnosis, and I quote it in full: "Stirps aphylla. Flores
unissexuales, 4-6 in axilles squamarum solitarii." No specific epithet is given, no
specimen cited, and no locality provided. We shall see that this brief diagnosis
contains an important inaccuracy: the flowers are in reality perfect.
M y first acquaintance with this truly remarkable mistletoe genus came when
studying a recent Colombian collection by Mr. P. C. Hutchison of the University
of California, Berkeley. Some characters of these plants tallied with the Ixocactus
diagnosis, and when Dr. T. Lasser, Caracas, kindly lent me the material upon which
Rizzini had based his new genus, it became at once apparent that the Colombian and
Venezuelan collections were identical. I am thus in a position to describe the type
of the genus, i.e., its first (and only) species. Rizzini's specific epithet, not published
but written on the label of the Venezuelan collection, referred to the simplicity of
the flower. This epithet, especially in view of Rizzini's erroneous view of the unisexual
floral condition, is misleading. Instead, I take pleasure in naming this species after
Mr. Paul C. Hutchison, indefatigable botanical collector of the Andean region.
I x o c a c t u s h u t e h i s o n i i Kuijt, sp. nov.

Frutex squamatus ramosissimus, statu sicco olivaceus. Internodia usque ad 7 cm
longa, valde alternatim complanata, circumscriptione oblonga vel subrhomboidea,
ad 10 mm lata, latitudine maxima paulum sub nodo, minima paulum supra nodum.
Squamae foliaceae magnitudine infra 1 ram, obtuse triangulares; phyllotaxis decussata.
Ramuli laterales vel flores haud ultra 5 pro axilla. Flores hermaphroditi, maturitate
2 mm longi, infra 1 mm lati. Calyculus manifestus. Perianthii partes 4, late ovales,
ca 1 mm longae, antheris singulis sessilibus obsitae. Antherae sacculis polliniferis
binis, minutis, longitudinaliter dehiscentes. Pistillum obtuse conoideum, basi quad-
rilobatum, stigmate capitato. Stylus persistens. Fructus bacciformis, ovoideus, maxi-
mus metatus 3.5 x 2 ram. Grana pollinis quadricolpata, globosa. Affixiones haus-
toriales observatae simplices, sine radicibus epicorticalibus.
TYPE: Tamayo 2526 (VEN), Venezuela, Lara, Guarico, San Isidro, Sep. 1942.
OTHER MATERIAL SEEN: P. C. Hutchison & J. M . Idrobo 3008 (UC, COL, F, NY,
US, MO, K, M I C H ) . COLOMBIA, DEPT. VALLE DEL CAUCA, Pacific slope, western
Cordillera, Finca Kyburz, 1 km above and east from Bitaco, eastern slope above
Bitaco River; common on orange trees. 4500 ft, 16 Nov. 1963.
BRITTONIA19: 62--67. Jan.-Mar. 1967.
62
1967] K U I J T : IXOCACTUS 63
FIGS. 1-6. Ixocactus hutchisonii. FIG. 1. Habit. FIG. 2. Vascular structure of flower in Fig. 3.
FIG. 3. Flower, the two nearest petals removed. FIG. 4. M a t u r e flower. FIG. 5. Two adjacent
petals as seen from within the flower. FIc. 6. Fruit. (Hutchison & Idrobo 3008.)
64 BRITTONIA [VOL. 19
Olive-green, squamate shrubs of a foot or more in diameter. Phyllotaxis opposite

and decussate. Internodes up to 7 mm long, greatly compressed, oblong to somewhat
rhombic. Width of internode up to 10 mm below the node, 5 mm or less just above.
Leaf scales less than 1 mm in size, bluntly triangular. Lateral branches (or flowers)
at most 5 per axil, being one branch each above and below the primary axillary
branch, plus one in each prophyll of the same, the resultant pattern being a cross-
shaped one. Flowers perfect, 2 mm long at maturity, and less than 1 mm in diameter.
Ovary crowned by a distinct, smooth calyculus, within which the four perianth mem-
bers are found, these broadly oval, ca 1 mm in length, usually four in number but
sometimes three, each with a sessile anther. Anthers with two minute pollen sacs
each, dehiscing longitudinally; each pollen sac with no more than 30 pollen grains.
Pollen sacs of one anther sometimes at slightly different heights, and anthers of
adjacent petals normally inserted at different levels, at least in tetramerous flowers.
Pollen quadricolpate (one 3-colpate and one 5-colpate grain seen), ca 100 ~ in
equatorial diameter and 80/~ from pole to pole, with a dense covering of stout spines
and four conspicuous, longitudinal colps. Style bluntly conical, persistent, basally
four-lobed (tetramerous flowers), stigma capitate. Fruit an oval berry, largest seen
3.5 • 2 ram, stigma remaining prominent.
This extraordinary mistletoe genus superficially resembles Phoradendron ]ragilis
and some other squamate species of Viscaceae. It is only upon closer study that it
can be assigned to Loranthaceae instead. It lacks the nodal constrictions which char-
acterize nearly all Viscaceae. Its flowers are perfect, a condition unknown in Viscaceae.
Its calyculus also indicates a position in Loranthaceae, even though similar structures
are said to exist in Viscum (Schaeppi & Steindl 1942).
It is at once obvious that Ixocactus is the most reduced of Loranthacean genera.
The squamate habit is very rare in the family, indeed may otherwise be restricted to
Phrygilanthus aphyllus. The advanced position of Ixocactus is even more evident in
the flower, by far the smallest in the family. In fact, most Viscacean flowers are
larger than those of the present genus, thus effacing a convenient difference between
the two mistletoe families. Again, in the structure of the stamen the degree of reduc-
tion is extreme. The filament is absent. It is a remarkable fact that a tetramerous
flower produces no more than 240 pollen grains. In some ways the anther shows
similarities to those of Phthirusa and Oryctanthus, also of Loranthaceae.
Ixocactus pollen is unique in Santalales through its combination of exine charac-
ters and the tetracolpate condition. Except for Viscaceae, where small spines may
be scattered on the surface of the grain, virtually no sculptured exines are known in
the order. No other member of the order has predominately tetracolpate pollen grains
except Ongokea, Harmandia and Aptandra (Reed 1955), an isolated, tree-like group
of Olacaceae bearing no other resemblances to Ixocactus. This is not to deny the
very rare appearance of tetracolpate grains as teratological products in some other
mistletoe genera (Dixit 1962).
The position of axillary branches is interesting, especially in the case of the ones
above a n d below primary lateral ones. In exceptionally vigorous individuals of
Struthanthus a similar seriation of entire inflorescences may sometimes be seen.
It is also of great interest that at least several of the individuals collected were
parasitic on other mistletoes. One was attached to the stem of a Phthirusa, in all
likelihood P. pyrifolia (HBK.) Urban. A large plant parasitized a stout epicortical
root of a mistletoe (Fig. 12) which, judging by the appearance of the root and its
haustoria, may have been an Oryctanthus. In both these cases, only a slight swelling
was noticeable on the host organ attacked. Such hyperparasitism is known from a
number of tropical mistletoes (Kuijt 1964a).
1967] KUIJT: IXOCACTUS 65
Fits. 7-12. lxocactus hutchisonii, Hutchison & ldrobo 3008. FIG. 7. Pollen, polar and
equatorial view. Diameter of grain ca 100 ~. FIa. 8. Apical portion of branch showing scale
leaves. Fic. 9. Axillary branching pattern. Axillant scale leaf below; all other brackets represent
prophylls (see text). Fic. 10. Transection of closed flower. FIG. 11. Insertion on branch of
Phthirusa sp. FIc. 12. Insertion on epicortical root, perhaps of Oryctanthus sp.
A third attachment is illustrated in Fig. 11. I t is evident from the haustorial skele-
ton (not shown here) that a gradual transition, a harmonious fusion, is present be-
tween the host and the parasite. The darker wood becomes irregular in its surface
contours near the host wood, but no individual haustorial strands are formed. This
is typical of a hyperparasitic connection ( K u i j t 1964a) b u t does not necessarily
mean that the parasite is unable to form a more elaborate haustorial system on non-
mistletoes. Since a certain correspondence exists between the gross wood structure of
this host branch and that of O r y c t a n t h u s we can perhaps say that all three specimens
66 BRITTONIA [VOL. 19
were growing on other mistletoes. The host branch parasitized by the Venezuelan col-
lection also appears to be a Phthirusa-like plant, although I cannot be certain of it.
One may search in vain through the many Loranthacean genera for mistletoes
showing close affinities with Ixocactus. Even where superficial similarities attract
the attention, as with the near-sessile flowers of the paleotropical genera Sogerianthe
(Danser 1933) and Barathranthus (Prakash 1963), the remaining differences pro-
hibit a close alliance. The rare occurrence of single axillary flowers in SSruthanthus
oerstedii (Kuijt 1964b) is also probably unrelated to the absence of anything like an
inflorescence in Ixocacsus.
Considering the nearly perfect separation of genera in the Old World from those
of the New World in the family, it is probably advisable to look for possible affinities
of Ixocactus with neotropicaI genera. I would suggest that OryctanShus is the only
Loranthacean genus showing any affinities with Ixocactus worth mentioning. Spheri-
cal pollen, very exceptional among Loranthaceae, is known from OrycSanthus (Kuijt
1961). OrycSanthus, also, has flowers which are completely sessile on an axis, even
if this axis happens to be an inflorescence axis. In fact, when we ignore the dis-
crepancy in size and petal number, the flowers of Oryctanthus (and its close relative
Phthirusa) and Ixocactus are very similar indeed. The major differences separating
the two genera are found in the epicortical roots, foliage leaves, and inflorescences of
Oryctanthus, to which are added very striking pollen-grain differences. Even with
respect to the latter it is interesting to note that only these two genera have diverged
from the typical, smooth, nearly triangular Loranthacean pollen type in the Americas.
Geographically, Ixocactus seems to occupy the core of the Oryctanthus territory.
It is rather strange that Ixocactus has had to wait so long to be properly described.
The Venezuelan collection bears the vernacular name "tifia" (dye), which may either
indicate confusion with other mistletoes, or suggest common occurrence. The Colom-
bian material was gathered from cultivated orange trees. It seems reasonable to pre-
dict, therefore, that the apparent rarity of Ixocactus is to some degree a function of
its humble appearance.
A word may finally be said with regard to the vasculature of the flower as it ap-
pears from material cleared in N a O H and chloral hydrate (Fig. 2). In a tetramerous
flower four vascular strands enter the base of the ovary and run the length of it to
supply the four petals. In some cases, just below the stamen, a simple dichotomy is
seen, one branch leading into the tip of the connective of the anther, the other running
the length of the petal. In other cases no branch is given off, the trace running
directly into the connective, leaving the remainder of the petal without vascular
supply. This seems to be true especially in many petals bearing the stamen in the
lower position. One instance has been noted where, on the contrary, the stamen was
evascular and the trace led into the tip of the petal. The latter situation is often met
in Viscaceae. Four very delicate strands reach about halfway up the style. They
remain separate and unbranched when followed downward, but their contact with
petal traces could not be ascertained in cleared material. Indeed, in position these
traces alternate with those of the petals, and there may not be any xylem continuity
at this time. In trimerous flowers a corresponding organization exists, showing three
petal traces and three stylar traces.
I t is a pleasure to acknowledge the assistance of three colleagues, the first two
from Utrecht, the last from Berkeley: Dr. Karel U. Kramer for the Latin diagnosis,
and Drs. Frans A. Stafleu and Paul C. Silva in matters of nomenclature.
LITERATURE CITED
Danser, B.H. 1933. A new system for the genera of Loranthaceae-Loranthoideae, with a nomen-
clator for the Old World species of this subfamily. Verh. Kon. Akad. Wetens. Amsterdam,
1967] KUIJT: IXOCACTUS 67
Afd. Nat., 2d Sect., 29(6) : 1-128.

Dixit, S. N. 1962. Rank of the subfamilies Loranthoideae and Viscoideae. Bull. Bot. Survey
India 4: 49-55.
Kuijt, Job. 1961. Notes on the anatomy of the genus Oryctanthus (Loranthaceae). Canad. Jour.
Bot. 39: 1809-1816.
1964a. Critical observations on the parasitism of New World mistletoes. Canad. Jour.
Bot. 42: 1243-1278.
---. 1964b. A revision of the Loranthaceae of Costa Rica. Bot. Jahrb. 83: 250-326.
P r a k a s h , S. 1963. Morphological and embryological studies in the family Loranthaceae--X. Ba-
rathranthus axanthus (Korth.) Miq. Phytomorphology 13: 97-103.
Reed, C . F . 1955. The comparative morphology of the Olacaceae, Opiliaceae and Octoknemaceae.
Mem. Soc. Brot. 10: 29-79.
Rizzini, C . T . 1952. Pars generalis prodromi monographiae Loranthacearum Brasiliae terrarum-
que finitimarum. Arq. Jard. Bot. Rio de Janeiro 12.39-126.
Schaeppi, H., & F. Steindl. 1945. Bltitenmorphologische und embryologische Untersuchungen
an einiger Viscoideen. Beih. Viertelj. Schr. Naturf. Ges. Ziirich 9 0 : 1-46.

Kuijt1967 Ixocactus

Uploaded by

Document Information

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Kuijt1967 Ixocactus

Uploaded by

Copyright:

Available Formats

THE GENUS IXOCACTUS (LORANTHACEAE, S.S.

Kuijt, J. (University of British Columbia, Vancouver, Canada). The genus

I x o c a c t u s h u t e h i s o n i i Kuijt, sp. nov.

Olive-green, squamate shrubs of a foot or more in diameter. Phyllotaxis opposite

Afd. Nat., 2d Sect., 29(6) : 1-128.

You might also like