Mediterranean pines and their history

The assembly of Mediterranean pines in the sense ofMirov is inhomogeneous in respect to morphological, geographical and evolutionary affinities. Considering new or neglected characters (vegetative and particularly reproductive, cone scales, apophyses, mucros, seeds: Figs. 1–3) in extant populations and fossils, three groups are recognized. The group of coast and island pines extends from the Canary islands to the Himalaya region and is closely related to Caribbean and C. American taxa. This complex evidently has originated from haploxyl ancestors of sect.Parrya during the Mesozoic (Upper Jurassic/Lower Cretaceous) in the NW. Tethys area (Fig. 13).P. rzedowskii can be regarded as an extant survivor of this first phase of differentiation. The extremely variableP. canariensis (together withP. roxburghii in sect.Sula) marks a transitional phase towards the more advanced diploxyl species of sect.Pinea (withP. pinea, P. halepensis &P. brutia, and subsect.Oocarpae) and sect.Pinaster (withP. pinaster, etc.) (Figs. 4–12).—The second group consists of diploxyl mountain pines from the areas surrounding the Mediterranean. They are classified as members of the Eurasiatic sect.Pinus subsect.Sylvestres, have differentiated along the northern Parathetys area, and exhibit close links with E. Asiatic taxa. The third group includes the haploxyl mountain pinesP. cembra andP. peuce which can be regarded as western outposts of the circumpacific centred sect.Strobus with a pre-Tertiary origin.—As an appendix, an improved classification scheme is presented for the pine groups discussed.     

Phenotypic and phenological plasticity of an aquatic macrophyteMenyanthes trifoliata L.

Phenotypic and phenological properties ofMenyanthes trifoliata L. were investigated within and outside of aPhragmites australis (Cav.) Trin. ex. Steud. canopy in a floating peat mat in Mizorogaike Pond, Central Japan. Under theP. australis canopy,M. trifoliata adjusted its phenotypic properties to the conditions of decreased light by increasing leaf blade area, decreasing leaf blade thickness and elongating petioles. The earlier expansion of leaves ofM. trifoliata within theP. australis community than outside the community was advantageous in terms of allowing the plant in the mixed community to produce as much dry matter as possible before the foliage ofP. australis could overgrow it. Despite the harmful effect ofP. australis onM. trifoliata's vegetative growth and reproduction, the latter species can persist in aP. australis community by changing its phenotypic and phenological properties.     

The perennial taxa ofCrucianella (Rubiaceae) in SW. Asia and their eco-geographical differentiation

The perennial taxa ofCrucianella in Asia form a coherent group, apparently diploid (x = 11) and outbreeding throughout, and should be placed into sect.Roseae. This Irano-Turanian group has its centre of diversity in the mountain systems south of the Caspian Sea and reaches with outposts NE. and E. Anatolia, NE. Iraq, S. Iran and C. Asia. Four species and 13 subspecies (within the polymorphicC. gilanica) are recognized, described (partly as new), and illustrated (Figs. 1–6). Conspectus, keys and distribution maps (Figs. 7 and 8) as well as plesio- and apomorphic character states and data on size of areas are provided (Table 1). There is an obvious correlation between more plesiomorphic taxa with smaller areas in the distribution centre of the group, and more apomorphic taxa with larger areas towards its periphery (Fig. 9). These findings are linked to a working hypothesis on the evolution of the group.Dedicated to Prof. DrLothar Geitler on the occasion of his 90th birthday.     

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.     

Contrasting effects of habitat structure on the recruitment and mortality of an epibiotic macroalga

Field surveys over 2 years in contiguous beds of the seagrassesZostera capricorni andPosidonia australis showed that the green algaCodium duthieae was consistently more abundant inZ. capricorni than inP. australis. In 1 year, mature plants were also more abundant at the boundary between the seagrass beds than in either bed. Field experiments and programmes of sampling were used to investigate three potential explanations for the unusual distribution of this alga: (1) that the availability of substrata suitable for attachment of the alga differed between the two seagrass beds; (2) that mortality of matureC. duthieae differed between the seagrass beds; and/or (3) that the intensity of recruitment was different in the two seagrass beds.C. duthieae plants were exclusively epibionts of the bivalveAnadara trapezia. Detailed sampling showed that the abundance ofA. trapezia was similar in both seagrass beds and that the distribution of bivalves suitable as substrata forC. duthieae plants was not obviously related to proximity to the boundary between the beds. Two experiments investigated the survival ofC. duthieae plants in each bed. In the first, matureC. duthieae plants transplanted into theP. australis bed suffered similar rates of mortality to plants which were disturbed and moved within theZ. capricorni bed or which were left undisturbed in theZ. capricorni bed. Fewer of the host bivalves were recovered from theZ. capricorni bed, however, indicating that the mechanism of mortality differed between the beds, hosts being more frequently dislodged in theZ. capricorni bed. Removal of the leaves of the seagrasses had consistently greater effects on near-bottom current velocities in theZ. capricorni bed than in theP. australis bed and significantly increased mortality ofC. duthieae in theZ. capricorni bed. Survival of plants was greater in plots of artificial leaves ofP. australis placed in theZ. capricorni bed than in plots of the naturalZ. capricorni leaves or plots where the natural leaves were removed. Most mortality in theZ. capricorni bed was due to dislodgement of the alga and its bivalve substratum. Corresponding manipulations of leaves in theP. australis bed had consistently smaller effects on survivorship of both the alga and its host. Patterns in the recruitment of the alga most clearly reflected the distribution of adults.C. duthieae recruits were 5 times more abundant in theZ. capricorni bed and at the boundary between the two beds than in theP. australis bed. The results demonstrate how habitat structure, provided by the canopy of leaves of the two species of seagrass, can have contrasting effects on the recruitment and mortality of a macroalga. In the case ofC. duthieae, it appears that the differential pattern of recruitment is the primary determinant of the distribution of adult plants.     

Distribution and deployment ofPresbytis melalophos group in Sumatera,Indonesia

Pelage color and pattern, facial markings, cranial morphology, and vocalization have been studied comparatively in thePresbytis melalophos group in Sumatera. Pelage coloration of fetuses and neonates were described forP. melalophos andP. femoralis. Based on this study, three species —P. thomasi, P. melalophos, andP. femoralis — are recognized. Using field data that have been gathered since 1981, the distribution of these three species has been mapped. The resultant map reveals that no sympatry is known and that rivers form the interspecific boundaries. However, rivers apparently are not barriers to the dispersal of species in this group. Instead, distributions appear to be limited ecologically, by competitive exclusion. Historically, it appears that the ancestral stock initially dispersed throughout this area, then theP. melalophos stock split from theP. thomasi-P. femoralis stock, theP. thomasi stock split from theP. femoralis stock, and, finally, theP. femoralis stock dispersed to eastern Sumatera. It appears unlikely that the centrifugal speciation hypothesis based on the principle of metachromism applies to deployment of theP. melalophos group.     

Host habitat finding and host selection of theDrosophila parasitoidLeptopilina australis (Hymenoptera,Eucoilidae), with a comparison of the niches of EuropeanLeptopilina species

Summary Decaying petioles of giant hogweed,Heracleum mantegazzianum Sommier & Levier, are used as a breeding site by six species ofDrosophila and the drosophilidScaptomyza pallida. The most numerous parasitoid species associated with this community isLeptopilina australis. BecauseL. australis was previously unknown in western Europe, we present the characters to distinguish it form its close relativeL. clavipes. Experiments on host species selection and survival ofL. australis showed that this parasitoid mainly usesD. limbata as host. Olfactometer experiments showed thatL. australis is attracted by the odour of decaying hogweed stalks, especially when these contain larvae ofD. limbata. L. australis is also strongly attracted by the odour of stinkhorns, a habitat in which it has never been found in nature.D. phalerata is the dominant fly species in stinkhorns, and is not a host ofL. australis. We offer a possible functional explanation for this unexpected habitat choice, by showing thatD. transversa andD. kuntzei, both species found to breed in fungi, are also suitable hosts forL. australis. We also discuss habitat choice with regard to a proposed phylogeny of theLeptopilina species in temperate Europe. Finally, we discuss niche overlap ofL. australis with the otherLeptopilina species.     

Vesicular-arbuscular mycorrhizal fungi in coastal dune plant communities II. Spore formation ofGlomus spp. predominates under geographically separated patches ofElymus mollis

Five study sites in Japan with pure patches ofElymus mollis near to the shoreline were chosen. Abundance of spore formation ofGlomus spp. underE. mollis patches were studied at four sites to understand the factors determining the development of VA mycorrhizal fungal communities at the edge of coastal sand dune vegetation. At each study site, eight soil samples from two soil depths were collected four times (three times for Niigata) during a year. Spores ofGlomus spp. predominated at every site, but species composition ofGlomus spp. differed from site to site. In a further experiment to measure the soil depth of the highest spore densities underE. mollis, the highest density was found at depths where rhizomes ofE. mollis were present.Contribution No. 119, Laboratories of Plant Pathology and Mycology, Institute of Agriculture and Forestry, University of Tsukuba.     

Cytogenetic studies on an intergeneric hybrid betweenAgropyron tsukushiense andElymus mollis

Elymus mollis is distributed widely from Korea to Japan, Kamchatka and Alaska, the northern part of U.S.S.R., and Northern and Eastern Canada, Greenland and Iceland. This species is tetraploid (2n=28). A strain of this species collected in Hokkaido was crossed withAgropyron tsukushiense var.transiens collected in Mishima. From this cross, 22 F₁ plants were produced. Crossability calculated from the number of hybrid plants produced and the number of floret pollinated was 30.6%. The shape of the F₁ spikes was of theAgropyron type but the glumes were hairy as were those of theElymus parent. One of the characteristics distinguishingElymus fromAgropyron is the production of two spikelets at almost all nodes of the rachis. This character was not expressed in the F₁ plants. All pollen grains of the F₁ plants were completely abortive. The average chromosome pairing at the MI of the PMCs of the F₁ amounted to 2.03 bivalents and 30.95 univalents. Almost all bivalents ranging from one to seven were rod-shaped connected with interstitial or terminal chiasma. These results indicate a lack of genomic homology between the three genomes ofA. tsukushiense and the two genomes ofE. mollis. Contribution No. 37 from the Plant Germ-plasm Institute, Faculty of Agriculture, Kyoto University, Kyoto, Japan.     

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.     

Alchemilla austriaca,spec. nova,(Rosaceae) — eine neue Art aus den Ostalpen

Alchemilla austriaca is a new species which belongs to the group ofA. demissa, A. frigens, A. longana, A. longiuscula, A. semisecta, andA. sinuata. The holotype specimen as well as leaf and flower details are illustrated (Figs. 1–3). A complete character analysis is given, differences and similarities of allied species are presented in two tables, and the position of the group within the genus is discussed.A. austriaca so far is known only from the Austrian Alps and mainly from the central ranges (distribution map: Fig. 4). Its wet subalpine and alpine habitats are characterized by species lists.

     

Modulation of flight by the giant interneurons of the cockroach

In this paper, I have examined the behavioral functions of feedback loops between the cockroach (Periplaneta americana) giant interneurons (GIs) and the flight thoracic rhythm generator.

Paleogeography and historical phytogeography (paleochorology) in the Neophyticum

After presenting the methodological bases of historical phytogeography (paleochorology; Fig. 1), a new paleographic map series is introduced (Fig. 2). They serve for ten examples of paleochorological analyses, in which the major phases of the development of a group in space and time are documented. In addition to purely tropical taxa (Nypa, Ctenolophon), others which originated in the warm and humid N. Tethyan belt (Bombacaceae p.p.,Olacaceae p.p.,Symplocos, Alangium, Nepenthes) or which presently occur as eastern N. American/E. Asian disjuncts (Sarcococca andPachysandra) are discussed (Figs. 3–14). Finally, analyses of two very old angiosperm groups with complicated paleochorological patterns (Restionaceae andAscarina, Chloranthaceae p.p.; Figs. 15–16) are presented.Part I, to be followed by part II in this journal.     

Interactions of thePolycomb group of genes with homeotic loci ofDrosophila

Summary Members of thePolycomb (Pc) group of genes are required for the correct determination of segment identity, and are thought to be negative regulators of thebithorax andAntennapedia complexes. This hypothesis has been tested molecularly for only some members of thePc group. Here, we examine the distribution ofUltrabithorax (Ubx),Antennapedia (Antp), andSex combs reduced (Scr) proteins in the epidermis, central nervous system, and midgut of embryos homozygous for mutations in tenPc group genes. We show that zygotic loss of mostPc group genes causes ectopic expression ofUbx andAntp, but that there are differences in time and tissue-specificity. FivePc group mutations lack midgut constrictions and also exhibit ectopic or suppressedUbx expression and suppression ofAntp expression. Distribution ofAntp is upset earlier than distribution ofUbx in the central nervous system of everyPc group mutant affecting both genes. Loss of the zygotic products ofPolycomb, extra sex combs, andAdditional sex combs cause ectopic expression ofScr in epidermis, and allPc group genes exceptPsc have suppressedScr expression in the nervous system. These results are discussed with respect to the function of thePc group.