Efficient pollination and high reproductive success in two Brazilian Spiranthinae orchids: Insights on the evolutionary history of pollination within the Pelexia clade

The pollination process and breeding system of Pachygenium bonariense and Pachygenium pteryganthum were documented for native populations from Rio Grande do Sul, Brazil. The flowers secrete nectar (on average with 65% to 67% of the concentration of total sugars), which is concealed into an incurved spur. Both species are pollinated by native, large Apidae bees, and the pollinaria adhere under the ventral surface of their labrum while bees try to reach the nectar. P. bonariense is pollinated by native bumblebees Bombus (Fervidobombus) morio and P. pteryganthum by females of oil-gathering Centris (Melacentris) sp. Both orchid species are self-compatible but pollinator-dependent (i.e., unable to set fruit and seed in the absence of pollinators). They also displayed high natural reproductive success (from 22% to 90%) and Nilsson's male efficiency factors ranging from 2.2 to 3.6, suggesting that, during our observations, more than one flower was pollinated for each flower acting as the pollen donor. To enrich the discussion, additional unpublished data on the pollination and breeding system of other orchid species of the Pelexia clade (the Spiranthinae group containing the studied species) are also presented and briefly illustrated. A phylogenetic framework of the clade is used to ascertain well-supported evolutionary scenarios for pollination-related floral traits (viscidium, protandry and a group of pollinators). The characteristic “dorsally adhesive” viscidium can be considered a synapomorphy for the subclade formed by the genera Cyclopogon, Veyretia, Sarcoglottis, Pachygenium, Pelexia and Brachystele, with a reversal to the plesiomorphic condition (ventrally adhesive) at the latter genus. Pollination by bees is likely the rule within the subclade and may be regarded as plesiomorphic for the whole Spiranthinae subtribe.     

Flower Morphology and Pollination Mechanism in Three Sympatric Goodyerinae Orchids from Southeastern Brazil

The pollination biology of Aspidogyne argentea, Aspidogyne longicornuandErythrodes arietina was studied in south-eastern Brazil.The three species are self-compatible but are pollinator-dependent.These three orchid species offer nectar as a reward to pollinatorsand flower visitors. The twoAspidogyne spp. have a dorsally-adhesiveviscidium, a feature which precludes pollinators other thanbees. Erythrodes arietina flowers are protandrous and show aventrally-adhesive viscidium. Aspidogyne argentea is visitedby halictid bees (here ranked as probable pollinators) and Hesperiidaebutterflies. Aspidogyne longicornu is pollinated by femalesof Euglossa(Euglossini) and visited by the hummingbird Phaethornisruber and by the euglossine bees Eulaema seabrai(females) andEulaema cingulata(males). The pollinarium adheres to the ventralsurface of the bee labrum, a very difficult place for bees toclean. The dorsally adhesive viscidium in Aspidogyne parallelsthat of the Spiranthinae genera of the so-called ‘Pelexiaalliance’. This condition seems to be particularly adaptativeunder conditions of low-frequency pollinator visits. Erythrodesarietina is pollinated by bees of the genera Paratetrapediaand Osiris which carry the pollinarium on the dorsal surfaceof their proboscis. Occasionally, these bees remove pollinariafrom their mouthparts using their forelegs. In general, in thespecies studied, a combination of both pollinator behaviourand morphological peculiarities promotes cross-pollination.Copyright 2001 Annals of Botany Company Orchids, Goodyerinae, Aspidogyne, Erythrodes, pollination, bees, hummingbirds, butterflies, morphology, viscidium, protandry     

Gynostemium structure and relationships of theCorallorhizinae (Orchidaceae: Epidendroideae)

Several types of pollinium stalk are present in theCorallorhizinae. Govenia has a cellular epidermal pollinium stalk, whileTipularia has a unique stipe that results from prolongation and detachment of the rostellum.Dactylostalix andEphippianthus have no stipe, but do have a detachable viscidium.Aplectrum, Corallorhiza, Cremastra, andOreorchis have hamuli. Cladistic analysis of morphological characters suggests that these four genera form a coreCorallorhizinae, and are sister group to a clade includingTipularia, Calypso, andYoania (the latter two possessing a true tegula).Govenia is shown to be most closely related to cymbidioid genera (such asEulophia), sharing with them the cellular epidermal pollinium stalk andCymbidium type velamen, and indicating that removal ofGovenia fromCorallorhizinae is appropriate.Dactylostalix andEphippianthus appear to occupy a more basal position in the subfamily, further arguing that the subtribe as previously recognized is paraphyletic.     

The pollination mechanism in the 'Pelexia alliance' (Orchidaceae: Spiranthinae)

The biological function of the wedge–like, dorsally adhesive viscidium of the neotropical orchid genera Cyclopogon, Pelexia and Sarcoglottis (grouped in the so-called 'Pelexia alliance') is elucidated by the study of the pollination biology of three species occurring in Sao Paulo State, southeastern Brazil. Cyclopogon congestus is pollinated by the bee, Pseudoaugochloropsis graminea (Halictidae), Pelexia oestrifera by workers of Bombus (Fervidobombus) atratus (Apidae) and Sarcoglottis fasciculata by males and females of Euglossa cordata (Apidae: Euglossini). These three species offer nectar as a reward and are self-compatible, though they need pollinators to set fruits. In spite of the difference in flower sizes and in their pollinators' taxonomic groups, the pollination mechanism is essentially the same for these species. The pollinarium adheres to the ventral surface of the bee labrum. The viscidium needs to be dorsally pressed in order to liberate a glue which fixes the pollinarium to a bee. Pollination is achieved by the interaction of the orchid column and the mouthparts of the bees. Fivation to the ventral surface of the labrum is advantageous for the orchid, since it is a difficult place for the bees to clean. Another advantage is that, since the labrum is articulated, when the bees fold and close their mouthparts, the pollinarium remains protected under the bee's head, thus reducing the risks of pollen loss. Since the wedge-like, dorsally adhesive viscidiurn is a characteristic feature of Cyclopogon, Pelexia and Sarcoglottis , it is suggested that some kind of phylogenetic constraint may exist, impeding the occurrence of pollinators other than bees in these orchid genera. All other flower-visiting animals lack the labrum-like structure needed to fix the pollinarium.     

PTEROCARYOID FRUITS (JUGLANDACEAE) IN THE PALEOGENE OF NORTH AMERICA AND THEIR EVOLUTIONARY AND BIOGEOGRAPHIC SIGNIFICANCE

The Juglandaceae (walnut family) has an excellent fossil record of various organs extending back to the earliest Tertiary. Several genera which today are restricted to isolated geographic regions were widespread in the Northern Hemisphere during the Tertiary. This paper focuses upon the fossil record of the Pterocarya alliance of the subfamily Juglandoideae, tribe Juglandeae. The Pterocarya alliance includes only two modern genera, Pterocarya (five species) and Cyclocarya (one species), both restricted to Eurasia. Paleogene sediments of the Rocky Mountain Region have yielded three genera and four species referable to the Pterocarya alliance: Cyclocarya (two species), Pterocarya, and a new genus, Polyptera. Although three of the four Paleogene species described here are attributed to present day genera, each represents an extinct form, which cannot be accommodated by any single living species. These fossils, reviewed with other published reports, indicate that the Pterocarya alliance, like the Engelhardia alliance of the same family, was more diverse and much more widespread geographically in the Tertiary than it is today.     

Phylogenetic relationships in the Marcetia alliance (Melastomeae,Melastomataceae) and implications for generic circumscription

The Marcetia alliance of Melastomataceae is an exclusively Neotropical group that includes at least 12 genera of mostly herbs and subshrubs, occurring in the cerrado of central Brazil and savannas of the Amazon region and Guayana highlands. This study aimed to test the monophyly of genera in the Marcetia alliance, evaluate their phylogenetic relationships and generic boundaries, and investigate morphological characters as potential synapomorphies for delimiting clades or genera. We used nuclear (ITS, ETS) and plastid (accD‐psaI, atpH‐atpF, trnS‐trnG) DNA sequences of 107 terminals in 12 genera from the alliance. Aciotis, Fritzschia, Marcetia and Siphanthera were shown to be monophyletic and supported by molecular and morphological characters. Other genera with variable morphology and wider distributions, such as Acisanthera, Comolia, Ernestia and Macairea, were recovered as paraphyletic or polyphyletic. Most morphological characters analysed were found to be homoplastic, but when combined they are potentially useful for the diagnosis of genera and infrageneric groups. This study represents a major step in understanding internal relationships and provides the basis for a revision of the generic classification in the Marcetia alliance.     

Spiranthes hachijoensis (Orchidaceae), a new species within the S. sinensis species complex in Japan,based on morphological,phylogenetic, and ecological evidence

The systematics of the Old World Spiranthes sinensis (Pers.) Ames species complex (Orchidaceae) has been complicated by its wide distribution and morphological variations. Within the species complex, S. australis Lindl. has been generally accepted as the only Spiranthes Rich. species distributed on the Japanese mainland. The present study provides morphological, phylogenetic, and ecological evidence for the recognition of S. hachijoensis Suetsugu as a new species of the S. sinensis species complex on the Japanese mainland. Spiranthes hachijoensis is morphologically similar to S. hongkongensis S.Y. Hu & Barretto and S. nivea T.P. Lin & W.M. Lin, sharing a degenerated rostellum, pollinia without a viscidium, and distinctly trilobed stigma. However, the taxon can be morphologically distinguished from S. hongkongensis by its glabrous rachis, ovaries, and sepals, and from S. nivea by its papillate labellum disc, larger papillate basal labellum callosities, and glabrous rachis, ovaries, and sepals. The autogamy and flowering phenology (i.e., earlier flowering) of S. hachijoensis are most likely responsible for premating isolation from the sympatric S. australis. A MIG-seq-based high-throughput molecular analysis indicated that the genetic difference between S. hachijoensis and its putative sister species S. sinensis is comparable to, or even greater than, the genetic difference between pairs of other species within the S. sinensis species complex. Our multifaceted approach strongly supports the recognition of S. hachijoensis as a morphologically, phenologically, phylogenetically, and ecologically distinct species.

     

COMPOSITION OF VOLATILES IN RELATION TO TAXONOMY OF AMERICAN ALLIUMS

The North American species of Allium exclusive of A. schoenoprasum and A. tricoccum of Old World affinity are grouped on the basis of morphological similarity into eight discontinuous species alliances typified by A. acuminatum, A. campanulatum, A. canadense, A. cernuum, A. falcifolium, A. kunthii, A. sanbornii, and A. validum, respectively. Representatives of each of these alliances were compared with respect to volatile constituents responsible for characteristic odors by means of gas chromatography. Results indicate that these volatiles provide evidence of relationship useful in the classification of alliums. Uniformity was found in composition of volatiles in the representatives of the A. canadense, A. cernuum, A. kunthii, and A. sanbornii alliances. Variation was observed in the A. acuminatum, A. campanulatum, and A. falcifolium alliances. A. validum was the only species of its alliance studied. Vapors of A. validum contain mostly n-propyl sulfides (onion-like odor) as does the cultivated A. cepa. Methyl sulfides (cabbage-like odor) predominate in the A. sanbornii alliance. A few species of the A. acuminatum and A. falcifolium alliances contain mainly allyl sulfides (garlic-like odor).     

Relationships ofLomatium among other genera ofApiaceae

Lomatium is the largest North American genus ofApiaceae (=Umbelliferae). Relations amongLomatium and other genera have not been well understood. TheApiaceae-Araliaceae lineage and genera of North AmericanApiaceae were analyzed phenetically and cladistically in turn, and seven phenetic alliances of genera are identified. Data on frequencies of character-state combinations were assembled and usingSporne's notions of maximal and minimal co-variation of states, two character-states were identified that are derived among Umbellifer genera. Using cladistic analysis of the alliance consisting ofLomatium and six other genera we identified three sister groups toLomatium:Polytaenia alone;Polytaenia withPseudocymopterus, and a group of four genera. Not all genera in the group of seven withLomatium are supported by autapomorphies impeding phylogenetic reconstructions. Both phenetics and cladistics have helped to suggest and evaluate relationships; to call attention to groups requiring further study, and in particular to identify derived character-states.     

The so‐called primitive genera of Genisteae (Fabaceae): systematic and phyletic considerations based on karyological data

A karyological analysis of the so‐called primitive genera of Genisteae has shown that they have a relatively homogeneous chromosome complement: all species tend to have a somatic chromosome number 2n = 48, which can increase to 2n = 52, presumably as a result of hyperaneuploidy. Karyological data suggest that Argyrocytisus, Cytisophyllum and Petteria may be considered as distinct genera rather than being assigned to Cytisus, with 2n = 52 for the first of these and 2n = 50 for the other two genera. They may be interpreted as relict monotypic genera as a result of the presence of a stabilized aneuploidy. Karyological characters exclude a recent origin of Genisteae from Thermopsideae. On the contrary, they are consistent with the hypothesis that Genisteae and Thermopsideae are independently derived from a basic papilionoid stock, of which present day Sophoreae are the remainder. At least two lines would lead from Sophoreae to the taxa of the ‘genistoid alliance’, one to Thermopsideae and the other ‘podalyrioid alliances’ (Podalyrieae and Mirbelieae), with the prevailing basic number of x = 9, and the other to Genisteae, with a basic number of x = 12 persisting in some present day genera, including Cytisus s.l. From this lineage, a wide range of secondary basic numbers has been formed, mostly by descending aneuploidy. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 160 , 232–248.     

Characteristics of three spinifex alliances in Central Australia

Abstract. The three most extensive spinifex alliances in arid Australia are characterised across a spectrum of post-fire successional states by their plant species, soil chemical characteristics and rainfall. Mean annual rainfall and some nutrients are correlated with the distributions of the alliances; the Triodia pungens alliance occurs in the highest rainfall region with the highest nutrient levels, the Plectrachne schinzii alliance in the mid-rainfall region with medium nutrient levels, and the T. basedowii alliance in the lowest rainfall region with a wide range of nutrient levels. This control on distribution can be over-ridden by topographic position. Where the T. pungens alliance occurs in low rainfall areas it is on sites favoured by soil moisture concentrations higher than the surrounding areas. Non-spinifex plant species are weakly associated with these alliances and their distributions relate to mean rainfall. Soil nutrient levels are generally low and, except for carbon, do not differ significantly between alliances. Soil grain sizes do not differ between alliances.     

Phylogeny and taxonomy of rhizobia

Rhizobia are bacteria that form nitrogen-fixing nodules on the roots, or occasionally the shoots, of legumes. There are currently more than a dozen validly named species, but the true number of species is probably orders of magnitude higher. The named species are listed and briefly discussed. Sequences of the small subunit ribosomal RNA (SSU or 16S rRNA) support the well-established subdivision of rhizobia into three genera: Rhizobium, Bradyrhizobium, and Azorhizobium. These all lie within the alpha subdivision of the Proteobacteria, but on quite distinct branches, each of which also includes many bacterial species that are not rhizobia. It has been clear for several years that Rhizobium, on this definition, is still too broad and is polyphyletic: there are many non-rhizobia within this radiation. Recently, therefore, it has been suggested that this genus should be split into four genera, namely Rhizobium (R. leguminosarum, R. tropici, R. etli), Sinorhizobium (S. fredii, S. meliloti, S. teranga, S. saheli), Mesorhizobium (M. loti, M. huakuii, M ciceri, M. tianshanense, M. mediterraneum), and a fourth, unnamed, genus for the current R. galegae. The evidence and pros and cons are reviewed.     

Spiranthes ovalis var. Erostellata (Orchidaceae), a new autogamous variety from the Eastern United States

Examination of flowers from herbarium specimens ofSpiranthes ovalis revealed two kinds of column structure. Some plants, including the type, have flowers with a normally developed rostellum and viscidium; other plants that have flowers in which these structures are entirely lacking are here described asSpiranthes ovalis var.erostellata. Self-pollination in the new variety was proven by pollinia removal and observation of plants in insect-proof cages. The var.erostellata has a somewhat smaller perianth with sepals 3.5–5 instead of 4–6.1 mm long and is more common and more widespread than the typical variety.     

Selfing potential in Epipactis palustris, E. helleborine and E. atrorubens (Orchidaceae)

Factors enabling and limiting self-fertilization were examined in populations of Epipactis palustris, E. helleborine and E. atrorubens located in NE Poland. In these species we found self-compatibility within the same flower (even 100% of fruit set from induced autogamy) and the same inflorescence (reaching 90–100% of fruit set from induced geitonogamy). Facultative autogamy was found only in E. palustris (even 82.5% of fruit set in a bagged inflorescence), due to underdevelopment of the clinandrium and projection of pollinia over the rostellum. Allogamy was predominant in E. helleborine (maximum to 2.8% of fruits set spontaneously) and E. atrorubens (maximum to 3.4%), attributable to a well-developed rostellum and clinandrium and to the position of pollinia. Autonomous selfing takes place during flower wilting, when the viscidium becomes evanescent and the pollinia are fragile. Selfing is enabled mostly by pollinator behaviour, promoting both geitonogamy and autogamy, and influencing mixed-mating.     

Systematic relationships of hymenolepidid cestodes of rodents and shrews inferred from sequences of 28S ribosomal RNA

Haukisalmi, V., Hardman, L. M., Foronda, P., Feliu, C., Laakkonen, J., Niemimaa, J., Lehtonen, J. T. & Henttonen, H. (2010). Systematic relationships of hymenolepidid cestodes of rodents and shrews inferred from sequences of 28S ribosomal RNA. —Zoologica Scripta, 39, 631–641. This study attempts to elucidate systematic relationships of hymenolepidid cestodes of rodents (18 species), shrews (13 species) and bats (one species) using sequences of partial 28S ribosomal RNA, with special reference to the genus Rodentolepis. The main finding is the presence of four multispecies clades of hymenolepidid cestodes showing pronounced morphological variation and frequent colonizations between unrelated hosts. Neither the hymenolepidid cestodes of shrews nor rodents were monophyletic. Also, the genus Rodentolepis sensu Vaucher in Czaplinski & Vaucher (1994, Keys to the Cestode Parasites of Vertebrates. Commonwealth Agricultural Bureaux International, Cambridge) is clearly non‐monophyletic. Although rostellar morphology is obviously a key feature on specific and generic levels, on higher systematic levels it seems to be a rather poor indicator of phylogenetic affinity in hymenolepidid cestodes. The presence of clades with more than one rostellar type (armed rostellum present, rudimentary unarmed rostellum present and rostellum absent) also conflicts with the proposed subfamilial and tribal classifications of hymenolepidid cestodes. The overall evidence suggests that the recent trend of splitting hymenolepidid cestodes into multiple genera will produce a more stable and practical classification than the earlier practice of favouring a few, morphologically variable genera. New classifications of hymenolepidid cestodes should, however, consider both morphological and molecular evidence.     

Plant communities of the montane mesophilous grasslands (Polygono bistortae–Trisetion flavescentis alliance) in central Europe: Formalized classification and syntaxonomical revision

Abstract

The research conducted here presents a syntaxonomical revision of the montane mesophilous meadows of the Polygono bistortae-Trisetion flavescentis alliance in central Europe – the Slovak part of the western and eastern Carpathians. These typical semi-natural grasslands occur mainly as small islands over the calcareous bedrocks. Associations of this alliance have tight relationships to the Arrhenatherion, Mesobromion and Nardo-Agrostion alliances. Formal definitions based on combination of the species groups were used regarding the diversity and geographical range of the Polygono bistortae-Trisetion flavescentis alliance. Following the formal definitions from the eight associations reported previously for Slovakia, only four of them can be placed within the Polygono bistortae-Trisetion flavescentis: Campanulo glomeratae-Geranietum sylvatici, Geranio sylvatici-Trisetetum, Crepido mollis-Agrostietum capillaries and Geranio-Alchemilletum crinitae. A comparison of traditional and formalized classification, and the advantages/disadvantages of the formalized classification are discussed.     

Phylogeny of the Pantomorus–Naupactus complex based on morphological and molecular data (Coleoptera: Curculionidae)

The Pantomorus–Naupactus complex is a Neotropical group of broad‐nosed weevils (Coleoptera: Curculionidae) including several parthenogenetic species usually assigned to the genera Naupactus Dejean, Pantomorus Schoenherr, Asynonychus Crotch, Aramigus Horn, Eurymetopus Schoenherr and Graphognathus Buchanan. Sixteen species were studied to test hypotheses on the monophyly of these genera, and on the origin of the parthenogenetic lineages. A matrix of 30 morphological characters and 999 positions of the Cytochrome Oxidase I gene, was analyzed with separate partitions and simultaneously, under equal and implied weights, and with different transversion/transitions costs. The ILD test indicates that the incongruence between the molecular and morphological data is not significant. Under equal weights, the molecular data resulted in a single tree and morphology in 34 trees; under implied weights morphology gave a different tree, and under TV:TS ≥ 4:1 molecular and combined analyses resulted in the same optimal tree. According to the latter, Naupactus includes Graphognathus, and is thus paraphyletic and basal regarding remaining genera, Pantomorus is polyphyletic and includes Aramigus and Asynonychus, and Eurymetopus is monophyletic. The species in which apomictic parthenogenesis has been verified (Aramigus tessellatus, Asynonychus cervinus and Graphognathus lecuoloma), belong to different clades of the Pantomorus‐Naupactus complex, with basal sexual relatives. © The Willi Hennig Society 2005.     

More than meets the eye: a morphological and phylogenetic comparison of long‐spurred,white‐flowered Satyrium species (Orchidaceae) in South Africa

Superficial similarities among unrelated species are often a result of convergent evolution and can cause considerable taxonomic confusion. A case in point is Satyrium eurycalcaratum , described here as a new species, which has been confused with several other Satyrium spp. with similar long‐spurred, white flowers. A phylogenetic analysis, based on molecular data, indicated that S. eurycalcaratum is not closely related to any of the species with which it has been previously confused. A comparative analysis of morphological characters in the seven South African Satyrium spp. with long‐spurred, white flowers showed that each of these, including S. eurycalcaratum , is characterized by a unique combination of traits. Despite the similarity in pollination syndrome characters, such as spur length and flower colour, variation in rostellum structure was particularly pronounced and four distinctive forms were present. There was no phylogenetic signal in patterns of interspecific rostellum variation, as some closely related species had different rostella, whereas some distantly related species shared similar rostellum structures. We therefore conclude that the use of rostellum traits in conjunction with phylogenetic evidence can resolve species delimitations among orchid species that share the same pollination syndrome. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 166 , 417–430.     

TWO NEW GENERA OF BRAZILIAN BAMBOOS RELATED TO GUADUA (POACEAE: BAMBUSOIDEAE: BAMBUSEAE)

Of all the genera of woody bamboos described from America, only three have pseudospikelets—Atractantha, Elytrostachys, and Guadua. The former two have pseudospikelets with an elongated rachilla internode that precedes the floret, making it pedicellate; they also share a type of leaf anatomy that separates them from the latter. The two new genera herein described, Criciuma and Eremocaulon, show most similarity to species of Guadua, both in spikelet morphology and in leaf anatomy. The study of these new taxa has helped to clarify the limits of New World genera with pseudospikelets and strengthened the case for maintaining Guadua as a genus distinct from its Old World counterpart, Bambusa. We are now able to discern a New World line of bamboos that includes Criciuma, Eremocaulon, and Guadua, separate from an Old World line that includes Bambusa, Dendrocalamus, and several other genera.     

NUMBERS AND DISTRIBUTIONS OF VASCULAR BUNDLES IN SCAPES OF THREE ALLIANCES OF THE GENUS ALLIUM L. (LILIACEAE)

The arrangement and numbers of vascular bundles were documented for the species of the Allium cernuum Roth, Allium validum Wats., and Allium kunthii G. Don alliances. These characters proved consistent and diagnostically useful in the A. cernuum and A. validum alliances, but inconsistent in the A. kunthii alliance.