Pollination of the southwestern Opuntias

This paper presents an overview of pollination systems inOpuntia sens. lat. in the southwestern United States. The floral syndrome of large, colorful, bowl-shaped, diurnal flowers is present in all of the prickly-pear species and most of the cholla (Cylindropuntia) species in this area. Pollination by medium-sized and large bees is established for some of the species with this floral syndrome and is predicted for the others. The same floral syndrome is widespread inOpuntia in other geographical areas.—Deviations from this floral syndrome occur in certain species of southwestern chollas, one of which has nocturnal disc-shaped flowers, and in several tropical American species groups or segregate genera with red hummingbird flowers. But such cases are relatively infrequent. Divergence between species with respect to floral syndrome and pollination system is not a common feature in the evolutionary pattern ofOpuntia. Pollination of North American Cacti, IV.—Previons parts of this series:Grant & Grant 1979a,Grant & al. 1979,Grant & Grant 1979b.     

The pollination spectrum in the southwestern American cactus flora

The cacti of the American Southwest, defined as the region from southern California to Texas, are surveyed for types of pollination systems and their frequencies. Four types of pollination systems are known to occur in the southwestern cactus flora: bee, hummingbird, hawkmoth, and bat pollination. Two other modes are suspected but not documented: miscellaneous smallinsect pollination and autogamy.—Bee flowers comprise a wide and nearly continuous series of size classes from very small to very large. The large bee flowers, with perianths 5.5 to 12.5 cm in diameter, form a prominent but arbitrarily delimited subcategory in the bee pollination system. Promiscuous flowers and autogamous flowers, if they occur, are included with the medium-sized and small bee flowers in our present classification, due to lack of information, and will have to be separated out when our knowledge is more complete. The overwhelming majority of species in the southwestern cactus flora are bee flowers. Between 39 and 44% of the species are large bee flowers. Another 50 to 56% of the species are classified as medium-sized and small bee flowers; while some of these may turn out to be promiscuous flowers and autogamous flowers, most of them are undoubtedly bee flowers.—Hummingbird, hawkmoth, and bat pollination are conspicuous but statistically minor components of the pollination spectrum, occurring in only one or a few species (see Table 2). Hummingbird pollination has arisen from bee pollination in the Southwest. The species or species groups with hawkmoth and bat pollination, on the other hand, are basically tropical groups which developed their advanced pollination systems in the tropics.Pollination of North American Cacti, V. See alsoGrant & Grant (1979a),Grant & al. (1979),Grant & Grant (1979b),Grant &Hurd (1979).     

Pollination ofOpuntia lindheimeri and related species

The large, yellow, bowl-shaped flowers ofOpuntia lindheimeri, O. discata, O. phaeacantha major, andO. compressa in Texas are visited by various species of beetles and bees. The beetles and small bees (Perdita, Dialictus) are pollen thieves. The pollinators are the medium-sized and larger bees (Melissodes, Diadasia, Lithurge, Megachile, Agapostemon, etc.). Different species of theOpuntia lindheimeri group have the same pollination system and there is no evidence of any floral isolation between them. The pollination system of these species ofOpuntia in Texas is essentially the same as that ofEchinocereus fasciculatus andFerocactus wislizeni in Arizona.Pollination of North American Cacti, II.—SeeGrant & Grant (1979).     

Pollination ofOpunta basilaris andO. littoralis

The flowers ofOpuntia basilaris andO. littoralis in southern California are visited commonly by beetles(Carpophilus, Trichochrous) and bees (especially anthophorids, megachilids, and halictids), but are pollinated mainly by the bees. This agrees with observations presented in the previous papers in this series for other cactus species in Arizona and Texas. The available evidence indicates that the large, diurnal, cup-shaped flowers in cacti of the American Southwest are primarily bee-pollinated. Our earlier view that theseOpuntia flowers are also pollinated to a significant extent bynitidulid andmelyrid beetles must be modified now in the light of further evidence. Some pollination probably is carried out by small beetles, but it probably represents only a small proportion of the total pollination.Pollination of North American Cacti, III.—See alsoGrant & Grant (1979) andGrant & al. (1979).     

Zwei neue Arten der GattungSaponaria (Caryophyllaceae) aus Afghanistan und Pakistan

Saponaria stenopetala sp.n. in Eastern Afghanistan is close toS. pachyphylla Rech. f. andS. subrosularis Rech. f.—The nearest allies ofS. makranica sp.n. from Western Pakistan and Southeastern Iran areS. kermanensis Bornm. andS. floribunda (Kar. & Kir.)Boiss.

Flora Iranicae praecursores 36–37. — Praecursores praecurrentes in Pl. Syst. Evol.139, 313–317 (1982).     

Die Klebstoffhaare an den Antheren vonCyclanthera pedata (Cucurbitaceae)

The highly specialized, two-celled hairs fringing the anthers ofCyclanthera pedata (L.)Schrad. and its relatives aid in pollination by producing a glue which sticks the coarse pollen grains onto insect visitors. This was proven by direct observations, confirming older assumptions. The adhesive is extruded from the big basal cell which bursts upon the slightest touch to the small top cell.—Turgor and, possibly, swelling pressure of the sticky material, are responsible for plasmoptysis and ejaculation. Cell contraction is due to tension of cuticular clips and, to a minor degree, to elasticity of the pectinic wall. A preformed apical dehiscence line disrupts when the top cell is bent aside.—The anthers hairs of a flower develop synchronously. From light and electron microscopic observations, the adhesive appears not to be a product of internal secretion. It originates from the endopolyploid giant nucleus of the basal cell, which finally degenerates by liquefying and swelling. The numerous peculiar elaioplasts apparently do not contribute to the hair's function.—The anther hairs are homologous to, and possibly have evolved from the so-called explosion hairs occurring on the green parts of manyCucurbitaceae.

Herrn Professor Dr.Walter Leinfellner zum 70. Geburtstag gewidmet.     

Taxonomic status of the species of the green algal genusNeochloris

Komárek has recently reviewed the various species assigned to the green algal genusNeochloris Starr (Chlorococcales, Chlorococcaceae) and removed those with uninucleate vegetative cells to a new genus,Ettlia. Watanabe & Floyd, unaware ofKomárek's work, also reviewed the species ofNeochloris and distributed them among three genera—Neochloris, Chlorococcopsis gen. nov., andParietochloris gen. nov.—on the basis of details of the covering of the zoospore and the arrangement of the basal bodies of the flagellar apparatus. This paper reconciles these two treatments and makes additional recommendations at the ranks of genus, family, order, and class.     

Pollenfertility and seed formation in theOrnithogalum umbellatum/angustifolium complex (Liliaceae/Scilloideae)

The pollen fertility and seed formation of six species of theOrnithogalum umbellatum/angustifolium complex and of seven related species were studied. Four types of pollen grains could be recognized. The pollen fertility varied greatly in this complex and is not related to the ploidy level. The seed formation ofO. umbellatum showed an adaptation to a subcontinental-Mediterranean climate, that ofO. angustifolium to an Atlantic climate. In both cases raindrops seem to be important for pollination, in view of the absence of insect pollinators. After open pollination 113 seedlings were obtained in four species. Their chromosome numbers were determined. Nearly all the cultivated seedlings were aneuploid, which points to a positive selection of euploids in nature, because aneuploid individuals are rare in the wild.Biosystematic Studies on theOrnithogalum umbellatum/angustifolium Complex III.—Previous parts of this series are Part I: Taxonomy. Proceeding Kon. Ned. Acad. Wet. series C,85 (4), 563–574 (1982) andvan Raamsdonk (1984).     

Pollen morphology ofSonchus and related genera,and a general discussion

The pollen morphology of the taxa belonging to the generaAetheorhiza Cass.,Launaea Cass.,Reichardia Roth andSonchus L. in the Iberian Peninsula has been studied with light and electron microscopy. The pollen is 3(-4)-zonocolporate and echinolophate (without polar lacunae, but in general with prelacunae), with equatorial ridges and 15–20 lacunae: 3–4 poral, 6–8 abporal and 6–8 paraporal. Small to medium size, P × E = 19–36 × 23–42 µm; sometimes two different sizes have been found. Exine 3–9 µm thick and ornamentation microreticulate and echinate. The results clearly show the relationships between genera. Moreno-Socías, E., Mejías, J. A., Díez, M. J., 1994: Morfología polínica deLactuceae (Asteraceae) en la Península Ibérica, I.Lactuca y géneros relacionados. — Acta Bot. Malacitana.19: 103–113.     

Omphalotaceae fam. nov. undPaxillaceae,ein chemotaxonomischer Vergleich zweier Pilzfamilien derBoletales

TheOmphalotaceae fam. nov., which include the generaOmphalotus andLampteromyces, are defined on the basis of characteristic sesquiterpenes and of their ability to cause white-rot. Anatomical and morphological features of the representatives of these genera support the creation of this new family. The occurrence of pigments, typical of theBoletales, and of cyanophilous spores, indicate membership of theOmphalotaceae in the orderBoletales. Relationships to the other families of this order — especially to thePaxillaceae — are discussed. The possible functional significance of fungal metabolites is considered.

Herrn Prof. Dr.J. Poelt zur Vollendung seines 60. Lebensjahres gewidmet.—Veränderte Fassung eines Vortrages auf der Tagung der Deutschen Botanischen Gesellschaft, Wien, September 1984.     

Fünf neue Arten der GattungAnthemis (Compositae) aus Iraq und Afghanistan

Five new species ofAnthemis sect.Anthemis from the Flora Iranica region are described:A. gillettii (subsect.Anthemis) from NW. Iraq and adjacent Iran is allied toA. damascena.—A. kurdica (subsect.Anthemis) also grows in Iraqi Kurdistan.—A. hamrinensis (subsect.Rascheyana; akin toA. plebeia) is distributed in the Jabal Hamrin region at the extreme outer margin of the Zagros chains.—A. kandaharica (subsect.Anthemis) andA. freitagii (subsect.Rascheyana) are distributed in Afghanistan.

Anschrift des Herausgebers: Hofrat Univ.-Prof. Dr.Karl Heinz Rechinger, Beckgasse 22, A-1130 Wien, Österreich.     

Six new species ofGagea (Liliaceae) from the Flora Iranica area

Six new species are described:Gagea anonyma, G. Staintonii, G. siphonantha, G. Grey-Wilsonii, G. chloroneura. All belong to subgen.Platyspermum (Boiss.)Miscz. Florae Iranicae praecursores63–68. — Praecursores praecurrentes: Pl. Syst. Evol.151, 281–293 (1986).     

Two newVeronica species (Scrophulariaceae) of Turkey and Iraq

Veronica davisii M. A.Fischer, sp. n., limited to the mountains of Kurdistan (S. E. Turkey and N. Iraq), is related to the Turkish-CaucasianV. gentianoides Vahl.—V. montbretii M. A.Fischer, sp. n., a local endemic of Erzincan prov. (E. Anatolia), shows affinities to the CaucasianV. liwanensis C. Koch and to the Turkish endemicV. oltensis Woron. & Schelk. from Erzurum province.     

Genetic mapping,distribution, and properties of an aconitase isozyme in Anopheles albimanus (Diptera:Culicidae)

Electrophoretic analysis of the developmental stages and tissues of Anopheles albimanus showed that qualitatively similar allozymes of aconitase (Acon-2) occur at all stages, and the enzyme is widespread in every larval and adult tissues. Relative heat stabilities of the allozymes were investigated by electrophoresis of heated aqueous extracts and by heating the enzyme in situ in acrylamide gels after electrophoretic separation in Tris-citrate and Tris-maleate buffer systems. The pupal aconitase in the crude extract is more stable to heat than the larval and adult enzyme. The presence of citrate ions in the gel increased the stability of aconitase to heat. Studies of substrate specificities indicated that cis-aconitic acid is the best substrate but citric acid can also serve as a substrate. Zymograms developed with isocitric acid as a substrate showed no aconitase electromorphs and produced only isocitrate dehydrogenase bands. Aconitase has a pH optimum of 8.0 and this enzyme is completely inhibited if treated in situ with ethylenediaminetetra-acetic acid (EDTA), p-chloromercuribenzoate (PCMB), and urea at concentrations higher than 5mm, 5×10^–5 m, and 2 m, respectively. Acon-2¹⁰⁰ and Acon-2¹⁰⁵ do not respond differently to the above treatments. Genetic crosses involving a holandric translocation, pericentric inversions, visible mutants, and allozyme markers were analyzed to map the aconitase (Acon-2) locus on the left arm of chromosome 3. The gene sequence (and map distances) on 3L is centromere—esterase-8 (Est-8)—2—esterase-4 (Est-4)—25—esterase-2 (Est-2)—9—Acon-2—5—phosphoglucomutase (Pgm)—7—esterase-6 (Est-6).     

Neoschumannia (includingSwynnertonia), a primitive genus of theAsclepiadaceae-Stapelieae

A comparison of the West AfricanNeoschumannia Schltr. and the East AfricanSwynnertonia S. Moore reveals that the two monotypic genera must be united.Swynnertonia is sunk into synonymy ofNeoschumannia and the new combinationNeoschumannia cardinea (S. Moore)Meve is made for the East African species.Neoschumannia is shown to belong to the tribeStapelieae. The taxon exhibits a very unusual character combination: the growth form of a woody liana is combined with a tripartite corona unique within theAsclepiadaceae. The morphology of the corona suggests a position ofNeoschumannia close to the base of theStapelieae —Ceropegiinae alliance.     

Megaspore wall growth inSelaginella (Lycopodiatae)

Different stages of megaspore and megasporangial development inSelaginella argentea (Wallich)Spring,S. bigelowii Unerw., andS. kraussiana (Kze.)A. Br. have been seen and studied. Megaspore wall units give positive reactions for polysaccharides and protein in young megaspores, and become the thick and resistant wall typical of the genus only later.—Units forming the exospore and the spaces between units enlarge from widths of 5–10nm early during development up to over 200 nm at pregermination stages. The spaces enlarge first. Initially they are circular and mostly about 70 nm in diameter. Later, spaces toward the inner part of the exospore enlarge more than those near the outer surface. During pregermination, wall spaces range in size from 4 to 50 times the width of units with the larger spaces located near the inner surface. As a result the exospore would be under tension to spring outward during germination when the laesurae are lysed.—A gap in the exospore, shaped like a half-moon in polar sections, forms in equatorial and distal portions of the spore. This gap becomes enormous, three times the volume of the central space plus the mesospore, and is filled with lipids and other nutrients. Late in development, during the period of tapetal cell degeneration, the gap contents are moved into the central space and the gap is closed.—Late in development the mesospore is degraded. Its products, along with gap contents, seem to be added to the contents of the central cavity and appear as reserve storage globules. A primary wall-like endospore is formed during this period, at the inner surface of the exospore. During germination this endospore develops further at its inner surface.—Changes in the size and shape of megasporangia occur independently of the size of megaspores.Megaspore development inSelaginella. II. For first part seeMorbelli & Rowley (1993).     

New and noteworthyHabenaria spp. (Orchidaceae) from New Guinea and adjacent islands

5 new taxa ofHabenaria, namelyH. bougainvillae, H. elongata R. Br. var.leptophylla, H. ensigera, H. rechingeri andH. trichoglossa, are described and illustrated, with reference to affinities to related Australian and Indo-malayan species. The occurrence in New Guinea of severalHabenaria spp. typical for a savanna-like vegetation, led to look more thoroughly at these taxa:H. elongata R. Br. andH. ochroleuca R. Br., considered so far to be endemic in Northern Australia, andH. khasiana Hook. f., hitherto only known from southeastern Asia.Studies in the subtribeHabenariinae Bentham (Orchidaceae), 2.—Part 1: Candollea34, 357 (1979).Dedicated to Hofrat Prof. DrKarl Heinz Rechinger on the occasion of his 80th birthday. — On April 30, in the year 1927, the author and his brotherOtto Renz metKarl Heinz Rechinger on a small steam-boat in a stormy Aegaen Sea, travelling from Piraeus to the Northern Sporades Islands:Karl Heinz with destination to Chelidromia, the author andO. Renz to Skopelos. By this lucky chance a lasting friendship began.     

Penicillin derivatives induce chemical structure-dependent root development,and application for plant transformation

We investigated five penicillin derivatives that are popularly used for transformation experiments with Agrobacterium rhizogenes—penicillin G, carbenicillin, ampicillin, amoxicillin and cephalexin—for their effects on the growth and morphology of Beta vulgaris, Capsicum annuum and Glehnia littoralis roots. Attention was given to the relationship between their chemical structures and functions. Ampicillin was found to stimulate root elongation but inhibit root branching, whereas carbenicillin inhibited root elongation but promoted root branching. Root cultures were also exposed to hydrolyzed products of these antibiotics—i.e. phenylmalonic acid (PM), phenylglycine and 6-aminopenicillanic acid (6-APA): PM inhibited root elongation the most, while root elongation was supported best by 6-APA. These results indicate that both the side chains and the major component of penicillin derivatives affect root development and that the nature of the side chains is responsible for the responses. Ampicillin but not carbenicillin was used in subsequent experiments described herein to eliminate bacteria and to support root growth of transformants of the recalcitrant plants.Abbreviations Amox: Amoxicillin - Amp: Ampicillin - 6-APA: 6-Aminopenicillanic acid - Carb: Carbenicillin - Ceph: Cephalexin - HG: d(–)-2-p-Hydroxyphenylglycine - PA: Phenylacetic acid - PenG: Penicillin G - PG: d(–)--Phenylglycine - PM: Phenylmalonic acid Communicated by L.C. Fowke     

An Iranian floristic link with Socotra inHypericum (Hypericaceae)

Hypericum dogonbadanicum Assadi, a local species from theQuercus brandtii forests of the Zagros Mountains in south-western Iran, belongs to sect.Campylosporus (Spach)R. Keller and is most closely related to a Socotran endemic,H. socotranum Good subsp.smithii N. Robson. Despite the inclusion of these forests in the Irano-Turanian Element of the Iran flora byZohary (1963),H. dogonbadanicum appears to belong rather to the Mediterranean dwarf-shrub maquis type of community and therefore to be a member ofZohary's (1963) Mediterranean/Irano-Turanian group of Iranian species.—The key position of the Socotran endemics to the understanding of the inter-sectional relationships is discussed.Dedicated to Prof.K. H. Rechinger on the occasion of his 80th birthday.     

Über zwei diploide,in Südwest-Anatolien endemische Arten aus dem Komplex derVeronica cymbalaria (Scrophulariaceae)

Veronica lycica Lehm. is a distinct, diploid member of theVeronica cymbalaria group, endemic in Lycia (S.W. Anatolia). Closely related is the newV. stamatiadae M. Fischer etW. Greuter which is also diploid and seems to be restricted to the small Greek island Ro close to the South coast of Lycia. The chromosome numbers for both species are reported for the first time (2n = 18).

Untersuchungen über den PolyploidkomplexVeronica cymbalaria agg., II. — Der erste Beitrag dieser Serie:Fischer (1975).