Contributions to the knowledge of sieve-element plastids inGunneraceae and allied families

P-type sieve-element plastids were found in theGunneraceae, while S-type plastids are present in theHaloragaceae andHippuridaceae. The specific characters of the sieve-element plastids (e.g., their size and the morphology of their contents) are discussed in relation to other taxa of theRosidae containing P-type plastids and to the systematic position of theGunneraceae. Contributions to the Knowledge of P-Type Sieve-Element Plastids in Dicotyledons, III. — For other parts of this series see (I.:)Behnke (1982 b) and (II.:)Behnke (1985).     

Further studies of the sieve-element plastids of theCaryophyllales includingBarbeuia,Corrigiola, Lyallia,Microtea, Sarcobatus,andTelephium

The sieve-element plastids of 69 species of theCaryophyllales were investigated by transmission electron microscopy. All contained the specific subtype-P3 plastids characterized by a peripheral ring of protein filaments. The presence or absence of an additional central protein crystal and their shape being either polygonal or globular as well as the average sizes of the sieve-element plastids are useful features in the characterization of some families.—Barbeuia contains sieve-element plastids that confirm its placement within thePhytolaccaceae. Lyallia differs fromHectorella by including small starch grains in their sieve-element plastids, which otherwise by their globular crystals negate a closer connection to theCaryophyllaceae. The lack of a central protein crystal in its form-P3fs plastids placesMicrotea best within theChenopodiaceae. Sarcobatus, a so far uncontested member of theChenopodiaceae, contains form-P3cf plastids, i.e., including a central crystal not found elsewhere in this family.Telephium andCorrigiola, shifted back and forth betweenMolluginaceae andCaryophyllaceae, have form-P3cf(s) plastids with a polygonal crystal which favor their placement within theCaryophyllaceae.     

Ultrastructure of sieve-element plastids inCaryophyllales (Centrospermae), evidence for the delimitation and classification of the order

The orderCaryophyllales (Centrospermae) was found to contain specific P-type sieve-element plastids which are characterized by protein inclusions composed of ring-shaped bundles of filaments and of central crystalloids. The sieve-element plastids of 14 families (140 species investigated) fit into this overall characterization, and more specific details are used to delimit the families and arrange them within the order.Phytolaccaceae, the basic family of the order display much diversity: the crystalloids inside their plastids are either globular (most genera) or polygonal (Stegnosperma), starch may also be present (Phytolacca).Nyctaginaceae, with starch inBougainvillea sieve-element plastids, can be derived directly fromPhytolacca. Globular crystalloids are present in most of the families, as inDidiereaceae, Cactaceae, Aizoaceae-Tetragoniaceae, Portulacaceae-Basellaceae-Halophytaceae-Hectorellaceae. Caryophyllaceae andLimeum ofMolluginaceae contain polygonal crystalloids (otherMolluginaceae with globular crystalloids). Crystalloids are entirely absent fromChenopodiaceae (incl.Dysphaniaceae) andAmaranthaceae. The probable relationships between these families are presented diagrammatically in Fig. 13. Bataceae, Gyrostemonaceae, Vivianiaceae, Theligonaceae, Polygonaceae, Plumbaginaceae, Fouquieriaceae, Frankeniaceae, andRhabdodendraceae—all at some time included into theCaryophyllales (Centrospermae) or doubtfully referred to them—develop S-type (or different P-type) sieve-element plastids. Their direct connection to theCaryophyllales therefore is excluded. Finally, evolutionary trends of theCaryophyllales are discussed.Presented in the Symposium Evolution of Centrospermous Families, during the XIIth International Botanical Congress, Leningrad, July 8, 1975.     

P-type sieve-element plastids and the systematics of theArales (sensuCronquist 1988)—with S-type plastids inPistia

The sieve-element plastids of 126 species of theArales were investigated by transmission electron microscopy. With the exception ofPistia (with S-type plastids) all contained the monocotyledon specific subtype-P2 plastids characterized by cuneate protein crystals. While the species studied from bothAcoraceae andLemnaceae have form-P2c plastids (i.e., with cuneate crystals only), those of theAraceae belong to either form P2c (14 species), P2cs (the great majority) or P2cfs (Monstera deliciosa, only, with form-P2cs plastids in the otherMonstera species studied). The form-P2cs plastids of theAraceae are grouped into different categories according to the quantity and quality of their protein and starch contents. The subfamilyLasioideae is redefined to comprise all aroid P2c-taxa and those P2cs-genera that contain only one or very few starch grains. Only little starch is also recorded in the sieve-element plastids ofGymnostachys (Gymnostachydoideae), with the other plastid data denying a close relationship toAcorus. While equal amounts of starch and protein are generally present in sieve-element plastids of the subfamiliesPothoideae, Monsteroideae, Colocasioideae, Philodendroideae, andAroideae, maximum starch content and only very few protein crystals are found in form-P2cs plastids ofCalla (Calloideae),Ariopsis (Aroideae), andRemusatia (Colocasioideae?). In the latter, both morphology and size of sieve-element plastids are close to those ofPistia.—In theAraceae the diameters of the sieve-element plastids exhibit a great size range, but are consistent within a species and within a defined part of the plant body. Comparative data are mainly available for stem and petiole sieve-element plastids.—The accumulated data are used to suggest an affiliation of the species to subfamilies and to discuss the phylogeny of theArales. Forms and sizes of their plastids support a separation of bothAcoraceae andLemnaceae from theAraceae. The presence of S-type plastids inPistia does not favour direct and close relationships to the form-P2c genusLemna.—The prevailing form-P2cs plastids might support proposals that place theArales (together with also form-P2cs plastid containingDioscoreales) in the neighbourhood of basal dicotyledons. BesidesAsarum andSaruma (Aristolochiaceae), with monocotyledonous form-P2c plastids,Pistia (with dicotyledonous S-type plastids) gives another example for a link between the two angiosperm classes.     

Sieve-element plastids,exine sculpturing and the systematic affinities of theBuxaceae

The sieve-element plastids of members of several genera in theBuxaceae (Buxus, Pachysandra andSarcococca) were found to be of the specific subtype PVI, which contains a central globular protein crystal.Simmondsia (Simmondsiaceae) andDaphniphyllum (Daphniphyllaceae), on the other hand, were found to contain S-type sieve-element plastids. The occurrence of the highly restricted PVI plastids in theBuxaceae mitigates against a close relationship between theBuxaceae andSimmondsia, Daphniphyllum andEuphorbiaceae. Exine sculpturing of theBuxaceae andSimmondsiaceae also shows no close similarities. Both of these EM characters are discussed in connection with other available data and with respect to earlier systematic treatment of these families.     

Serologische Untersuchungen zur Gliederung derBrassicaceae

The serological investigations support the opinion ofJanchen (1942) to combine the generaBunias, Isatis, andSisymbrium in the tribeSisymbrieae; Cheiranthus, Erysimum, andMatthiola in the tribeHesperideae; andBrassica, Crambe, Sinapis, andSuccowia in the tribeBrassiceae. They further underline the central position of theSisymbrieae and the isolated position of theHeliophileae. In accordance withEigner (1973) theBrassiceae are placed closer to theSisymbrieae than inJanchen; the same holds for thePringleeae. No serological justification could be found to uniteArabis andBarbarea in the tribeArabideae, andAlyssum andLunaria in theAlysseae. From the antigen-systems used among the representatives ofJanchen's Lepidieae the generaLepidium andNeslia show remarkable correspondence both toCamelina andThlaspi, but not toCochlearia which appears distant fromCamelina andThlaspi also.

Teil 1/Part 1.     

Eine blütenmorphologische Analyse derLagoecieae (Apiaceae)

An analysis of the morphology, anatomy and ontogeny of the flowers, particularly of the gynoecium ofLagoecieae is presented. 1. The gynoecial model of angiosperms can be applied to all three generaArctopus, Lagoecia andPetagnia. 2. In the case ofArctopus an additional Apikalseptum is developed. 3. In the synascidiate region of the gynoecium the adaxial carpel is reduced inArctopus andPetagnia, the abaxial inLagoecia. 4. The reduced carpel produces either one mature ovule inArctopus, a rudimentary ovule inPetagnia, or none inLagoecia. 5.Petagnia andLagoecia have a completely pseudomonomerous gynoecium. 6.Arctopus displays many flower characteristics which lack in theSaniculoideae but occur in theHydrocotyloideae. 7. ForPetagnia andLagoecia an independent phylogenetic development within theSaniculoideae is assumed.

Herrn Univ.-Prof. Dr.Walter Leinfellner zum 70. Geburtstag gewidmet.     

Chemosystematics ofSuillaceae andGomphidiaceae (suborderSuillineae)

The chemotaxonomic findings relating to the generaBoletinus, Suillus, Gastroboletus, Gomphidius, andChroogomphus are summarized and discussed, using published data as well as our own hitherto unpublished evidence of pigments and chromogens. The study confirms repeatedly made claims that these genera are closely related. In addition to the presence of pigments which are typical for most members of theBoletales (e.g., pulvinic acid derivatives, terphenyl quinones, cyclopentenones), prenylated phenols and quinones can also be constantly detected here (with the exception ofBoletinus), just as inRhizopogon. Accordingly,Suillus is more closely related to theGomphidiaceae andRhizopogonaceae than to the remaining boletes. It is therefore necessary to establish a new family (Suillaceae which includeBoletinus, Suillus, andGastrosuillus) and a new suborder (Suillineae which includeSuillaceae, Gomphidiaceae, andRhizopogonaceae) within theBoletales. Chemosystematics ofBoletales 2. For part 1 seeBesl & al. (1986). Dedicated to emer. Univ.-Prof. DrFriedrich Ehrendorfer on the occasion of his 70th birthday     

S-type sieve-element plastids and anthocyanins inVivianiaceae: Evidence against its inclusion intoCentrospermae

The presence of S-type sieve-element plastids and anthocyanins in theVivianiaceae indicates that it is not a member ofCentrospermae (Caryophyllales).     

Form-Pfs Plastids,Stem Anatomy and Systematic Affinities of Stylobasium Desf. (Stylobasiaceae)

The vascular system of the stem of Stylobasium was investigated during its primary and secondary phases with both light and electron microscopic methods. It contains collateral bundles arranged in a ring, separated by rays which undergo regular cambial growth. The phloem consists of short sieve elements connected to sieve tubes by simple sieve plates, companion cells of the same length, and phloem parenchyma cells. During their autophagy-like differentiation and maturation, typical of all angiosperms, the sieve elements of Stylobasium have a peculiar feature, whereby they develop and retain form-Pfs plastids (containing protein filaments and starch). The sieve-element plastids of the two Stylobasium species, and of some 100 species belonging to taxa of which Stylobasium had been considered to be a possible member, have been studied by transmission electron microscopy. With the exception of a few species with form-Pcs plastids (containing a single small protein crystal in addition to starch), the great majority of taxa studied are characterized by S-type sieve-element plastids (containing starch only). The presence of form-Pfs plastids in Stylobasium supports its separation into the unigeneric Stylobasiaceae and the placement of this family close to other form-Pfs or form-Pcfs-containing taxa. While other characters would exclude an affiliation to the Magnolianae (form-Pfs plastids in Canella) or Caryophyllales (form-Pfs plastids in Microtea), an association with the form-Pcfs families Connaraceae and Mimosaceae is positively considered and corresponds to their frequent allocation close to the Rutales and Sapindales. Within the Rutales/Sapindales the sizes of sieve-element plastids (average diameter) range from very large (e.g. in the Julianaceae) to comparatively small (e.g. in Aceraceae) and are used to group the families. The sieve element characters of the Coriariaceae (tiny plastids with almost no starch, wide sieve plate pores, copious P-protein) suggest their removal from Rutales/Sapindales into the neighbourhood of the Cucurbitaceae.     

Host-specificity of a root borer,Bembecia chrysidiformis [Lep.: Sesiidae], a potential control agent forRumex spp. [Polygonaceae] in Australia

Bembecia chrysidiformis (Esper) [Lep.: Sesiidae] was examined for its natural history and specificity toRumex spp. (Polygonaceae) which are weeds in Australia. Adults of this southern European insect appear in late spring to summer. Eggs are laid on the dried, seed bearing stems of perennialRumex plants. The larvae tunnel inside the root during summer through to the next spring. In nature, the larvae are round inRumex species of the subgeneraRumex andAcetosa. In host-specificity tests with 1st instar larvae, the roots of a number of genera within thePolygonaceae were attacked. Larvae died on a range of plants from other families except inPersea americana Miller(Lauraceae), Helianthemum nummularium (L.) Miller (Cistaceae) andQuercus ilex L. (Fagaceae) where larvae fed on the stems. The insect was judged safe for release in Australia by assessing aspects of its biology, its known host plants, and the lack of reported attack on other plants.      

Taxonomy and phylogeny of the tribePlucheeae (Asteraceae)

The tribePlucheeae (Benth.)A. Anderb., has been analysed cladistically by means of a computerized parsimony program (Hennig 86), using theArctotideae as outgroup. The results of the analysis are presented in a consensus tree and one cladogram. Four major monophyletic subgroups can be recognized: TheColeocoma group (3 genera), thePterocaulon group (3 genera), theLaggera group (6 genera), and thePluchea group (12 genera). All recognized genera are described and most genera are supplied with taxonomical notes including comments on their taxonomic status. Genera such asBlumea, Pluchea, andEpaltes are demonstrated to be unnatural assemblages.Monarrhenus andTessaria are both closely related to thePluchea complex. The old generic nameLitogyne Harv. has been taken up for one species ofEpaltes, the genusRhodogeron is reduced to a synonym ofSachsia, and the following new combinations are made;Litogyne gariepina (DC.)A. Anderb., andSachsia coronopifolia (Griseb.)A. Anderb.     

Weitere Untersuchungen über Proteinkörper in Zellkernen und ihre taxonomische Bedeutung

In 83 species of the familiesMonotropaceae, Apocynaceae, Oleaceae, Scrophulariaceae, Lentibulariaceae, Bignoniaceae, Martyniaceae, Myoporaceae, Verbenaceae, Lamiaceae, Campanulaceae, andLiliaceae, protein bodies in the cell nuclei have been found, in 68 of these species for the first time. On the basis of their structure in accordance with morphological characters the generaBurgsdorfia, Hesiodia, Olisia, andPhlomoides of theLamiaceae are accepted;Lamium is divided intoLamium, Lamiastrum andOrvala; two new combinations are established:Kickxia campyloceras (Rech. fil. &Esfandiari)Speta andEtornotus papilionaceus (Burm. in L.)Speta. Deviating shape or lack of protein bodies corroborate former taxonomic decisions, e.g. the transfer ofMonotropa toMonotropaceae or the separation ofGaleopsis andLadanum; Veronica schmidtiana should not be included inPseudolysimachion. Systematic affinities are discussed primarily withinScrophulariaceae because nuclear protein bodies have been studied already in many species of this family. ForCampanula patula two 2 x populations are reported.

Herrn Professor Dr. L.Geitler zum 80. Geburtstag gewidmet.     

Classical morphological features of centrospermous families

The orderCentrospermae (Caryophyllales, Chenopodiales) as treated inA. Englers Syllabus, 12th edition (1964), is compared with several other modern and older systems with the result that no less than 11–13 (and more) families are considered to be centrospermous in the strict sense; to these may be added thePolygonales and, doubtfully, thePlumbaginales andBatidales. As indicated by their name Centrospermae their main character is the central or basal placentation in combination with campylotropy (or amphitropy) of the ovules, seeds with perisperm, and coiled or curved embryos in peripheral position. Other outstanding features are found in the embryology; the ovules are bitegmic-crassinucellate, a nucellar cap is present, as well as an endostome and air spaces; the pollen is trinucleate. Anomalous secondary thickening in stems and roots often occurs. The pollen morphology, specific P-type sieve-element plastids, and the presence or absence of betalains are also important characters. Other floral features, especially the structure of the gynoecium, the androecium, the perianth and the receptacle, as well as the morphology of the inflorescences are of taxonomic importance. The putative relationships of theCaryophyllidae can perhaps best be resolved on the basis of more detailed morphological investigations (e.g. the so-called apocarpy, the development of the androecium, the pollen morphology, chromosome numbers, etc.).Presented in the Symposium Evolution of Centrospermous Families, during the XIIth International Botanical Congress, Leningrad, July 8, 1975.     

Elektronenmikroskopische Untersuchungen zur Frage der verwandtschaftlichen Beziehungen zwischenTheligonum undRubiaceae: Feinbau der Siebelement-Plastiden und Anmerkungen zur Struktur der Pollenexine

Theligonum cynocrambe and 13 species ofRubiaceae contain S-type sieve-element plastids, wide-spread in Dicotyledons. Alignment ofTheligonum toCaryophyllales (Centrospermae), especiallyPhytolaccaceae, is unlikely, because this order is characterized by specific P-type plastids. SEM investigations show the pollen exine ofTheligonum to be microreticulate, with additional supratectate spinules.Asperula and other genera of the tribeRubieae have a tectum perforatum (punctitegillate sexine), also with supratectate verrucae or spinulae.—Thus ultrastructure suggests (but not definitely proves) relationships betweenTheligonum andRubiaceae, while affinities betweenTheligonum andCaryophyllales are excluded.

     

Relationships in theAcanthaceae and related families as suggested by cladistic analysis ofrbcL nucleotide sequences

A parsimony analysis of DNA sequences of the chloroplast-encoded generbcL from twelve members of theAcanthaceae s.l., including members of the sometimes segregateThunbergioideae andNelsonioideae, and other families in theBignoniales sensuThorne (1992) is presented. The results largely agree with the classification of theAcanthaceae presented byBremekamp (1965) andThorne (1992) and supportNelsonioideae as a sister group to the rest of theAcanthaceae. Thunbergioideae are placed as a sister toAcanthaceae s.str.Acanthus andAphelandra, both representatives ofAcanthoideae, form a sister group toRuellioideae. An analysis of branch support found that many branches throughout theBignoniales are weakly upheld. This points to the need for further studies in the group using more sequences ofrbcL as well as other data. None of the families ofBignoniales as presently circumscribed (includingAcanthaceae s.l.) were strongly supported, although the larger clade containing the families of theBignoniales was robust.     

Cytotaxonomical studies on AsianAnnonaceae

Chromosome numbers of 42 species and 3 varieties from 24 genera of theAnnonaceae have been determined (Table 1); reports for 15 of the genera are new. Among Asian genera 2n = 14 occurs only in the specializedMezzettia, while 2n = 16 is wide-spread and also has been found inAnaxagorea with some primitive characters. 2n = 18 is reported for 11 genera, and tetraploidy (2n = 36) has been observed inPolyalthia. Therefore, an original basic number of x = 8 or x = 9 is suggested at least for the Asian genera of theAnnonaceae.—Cytotaxonomical notes on the critical speciesPolyalthia rumphii andP. affinis are given, and the new combinationNeouvaria parallelivenia (Boerl.)Okada & Ueda is proposed.     

The chloroplast genome arrangement ofLobelia thuliniana (Lobeliaceae): Expansion of the inverted repeat in an ancestor of theCampanulales

A clone-bank ofSac I restriction fragments was constructed from the chloroplast DNA (cpDNA) ofLobelia thuliniana E. B. Knox (Lobeliaceae). These cloned fragments and a set of 106 clones spanning the tobacco chloroplast genome were used as probes to determine the cpDNA restriction fragment arrangement forSac I and six other restriction enzymes (BamH I,EcoR V,Hind III,Nci I,Pst I, andXho I) and the chloroplast genome arrangement ofL. thuliniana relative to tobacco, which has been fully sequenced and is collinear with the hypothesized ancestral genome arrangement of angiosperms. The results confirm and refine our previous understanding of the chloroplast genome arrangement in the large single-copy region (LSC) and reveal (1) a roughly 11 kilobase (kb) expansion of the inverted repeat (IR) into the small single-copy region (SSC) and (2) apparent sequence divergence of the DNA segment inL. thuliniana that corresponds to ORF1901 in tobacco. The expansion of the IR into the SSC is present in all other examined members ofLobeliaceae, Cyphiaceae, andCampanulaceae, which indicates that the IR expansion was an early event in the cpDNA evolution of theCampanulales. The IR expansion into the SSC was not present inSphenoclea, which additionally supports exclusion of this genus from theCampanulaceae.     

Accumulation of seed storage proteins and the taxonomy ofPoaceae

The accumulation of specific seed proteins is a taxonomically valuable feature and can be used to additionally characterize plant taxa. To date, mainly crop proteins have been analysed in thePoaceae. In this investigation seed proteins from 147 species were screened with emphasis on legumin-like proteins and prolamins. The groups resulting from evaluation of the protein profiles correspond with well-known subfamilies and tribes.Panicoideae are clearly separated fromPooideae. WithinPooideae, theBromeae plusTriticeae tribes revealed obvious similarities.Lolium, Festuca andVulpia, generally included in the tribeFestuceae, revealed a protein profile similar to the profile of theBromeae/Triticeae. Legumin-like proteins are accumulated abundantly inBambusoideae andPooideae exceptBromeae/Triticeae, however, only the species included in theAveninae subtribe produce soluble (globulin-type) legumins as already known fromAvena sativa. Dedicated to emer. Univ.-Prof. DrFriedrich Ehrendorfer on the occasion of his 70th birthday     

Interspecific hybridizations among species of theElymus semicostatus andElymus tibeticus groups (Poaceae)

Interspecific hybridizations were made between species of theE. semicostatus group, viz.,E. semicostatus (Nees exSteud.)Meld.,E. validus (Meld.)B. Salomon,E. abolinii (Drob.)Tzvel., andE. fedtschenkoi Tzvel., and species of theE. tibeticus group, viz.,E. pendulinus (Nevski)Tzvel.,E. tibeticus (Meld.)Singh,E. shandongensis B. Salomon, andE. gmelinii (Ledeb.)Tzvel., as well as among species within theE. tibeticus group. All species are tetraploid (2n = 4x = 28) and possess SY genomes. Meiotic pairing data from 24 hybrids involving 17 interspecific combinations are presented. The average number of chiasmata per cell ranged from 17.91 to 26.20 in hybrids within theE. tibeticus group, compared with 7.26 to 22.04 in hybrids between the two species groups. Despite the extensive collection of cytological data, there was no definite evidence for confirming or disproving the separate existence of the two groups.