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Lobelioideae , the largest subfamily within Campanulaceae , includes 33 genera and approximately1200 species. It is characterized by resupinate flowers with zygomorphic corollas and connate anthers and is widely distributed across the world. The systematics of Lobelioideae has been quite challenging over the years, with different scholars postulating varying theories. To outline major progress and highlight the existing systematic problems in Lobelioideae , we conducted a literature review on this subfamily. Additionally, we conducted phylogenetic and biogeographic analyses for Lobelioideae using plastids and internal transcribed spacer regions. We found that former studies have reached agreement on the southern African origin of Lobelioideae , herbaceous habit and Asian origin of giant lobelioids, the convergent evolution of giant rosette lobelioids, and lastly, the multiple cosmopolitan and independent radiation of lobelioids in Africa, Pacific Basin, and the Hawaiian Islands. Also, Apetahia Baill., Sclerotheca A.DC., and Cyanea Gaudich. are paraphyletic, while Lobelia L., Pratia Gaudich., Centropogon C.Presl, Siphocampylus Pohl, and Isotoma Lindl. are polyphyletic. The taxonomy of these genera, especially Lobelia , is particularly quite frustrating. This calls for further reappraisals using both morphological and molecular data.

Eleven plants were collected at the Zongo Valley to evaluate their antioxidant and photo-protector properties. In this paper we report a strong correlation between high antioxidant activity and strong UV-A and/or UV-B absorptions. The most active species, tested at 10µg/ml with the DPPH assay, were Fuchsia boliviana (leaves), Baccharis pentlandii (flowers), Rubus floribundus (fruits), Fuchsia boliviana (flowers and fruits) and Brachyotum microdon (flowers). All the mentioned species have important UV- B and/or UV-A absorptions. This DPPH/UV technique could be used to preliminary screen vegetable samples and to select those with DPPH values above 83% and strong UV-A and/or UV-B absorptions. The chosen samples can then be evaluated with other more expensive in vitro assay (TEAC, ABTS or FRAP) to finally confirm their activities with the in vivo test. To our knowledge, this is the first time that the antioxidant properties of Distichia muscoides , Souroubea fragilis , Brachyotum microdon , Monnina bridgesii , Baccharis pentlandii , Thibaudia crenulata , Siphocampylus tupaeformis , Cobaea scandens , Fuchsia boliviana and Rubus floribundus are reported. In addition, this is the first time that Siphocampylus tupaeformis and Thibaudia crenulata are presented in a publication as well as the study of their photo-protector and antioxidant properties.

Two species of Siphocampylus (Campanulaceae: Lobelioideae) from the Central Andes of Peru and Bolivia are described, illustrated, and discussed with reference to related species. One species, Siphocampylus antonellii, is endemic to high elevation grasslands of Calca, Peru, while the second, Siphocampylus siberiensis, is endemic to cloud forests of Cochabamba, Bolivia. Both species are robust shrubs that produce tubular pink flowers that are likely pollinated by hummingbirds.

During the revision of Siphocampylus for the Flora of Argentina, we detected names which need lectotypification. As a result of our analysis of original materials and protologues, lectotypes are designated for the following names: S. fiebrigii var. intermedius, S. foliosus, S. foliosus var. minor, S. lorentzii, S. nemoralis, S. nemoralis var. grisebachii, S. nemoralis fo. hieronymi, S. orbignianus, and S. verticillatus. Fil: Salomón, Luciana. Consejo Nacional de Investigaciones Científicas y Técnicas. Instituto de Botánica Darwinion. Academia Nacional de Ciencias Exactas, Físicas y Naturales. Instituto de Botánica Darwinion; Argentina Fil: Nicola, Marcela Viviana. Consejo Nacional de Investigaciones Científicas y Técnicas. Instituto de Botánica Darwinion. Academia Nacional de Ciencias Exactas, Físicas y Naturales. Instituto de Botánica Darwinion; Argentina Fil: Nomdedeu, Soledad María. Consejo Nacional de Investigaciones Científicas y Técnicas. Instituto de Botánica Darwinion. Academia Nacional de Ciencias Exactas, Físicas y Naturales. Instituto de Botánica Darwinion; Argentina

thomesianus, Siphocampylus, S. Moricand, 1847: 142, pl. 85. Type locality: in Brasilia prope S. Thomé prov. Bahiensis [Blanchet] (Campanulaceae)., Published as part of Breure, Abraham S. H. & Tardy, Emmanuel, 2016, From the shadows of the past: Moricand senior and junior, two 19 th century naturalists from Geneva, with their newly described taxa and molluscan types, pp. 113-138 in Revue suisse de Zoologie 123 (1) on page 119, DOI: 10.5281/zenodo.46292

Resumo O presente manuscrito apresenta o tratamento florístico de Campanulaceae no Parque Nacional do Itatiaia. A família está representada na área por três gêneros, Lobelia com duas espécies: L. camporum e L. fistulosa, Siphocampylus, com quatro espécies: S. duploserratus, S. longepedunculatus, S. umbellatus e S. westinianus e Wahlenbergia, com uma espécie: W. linarioides. As espécies foram encontradas majoritariamente na parte alta do parque, geralmente associadas a afloramentos rochosos dos campos de altitude, com vegetação graminoide ao redor. Somente S. longepedunculatus foi coletado em sub-bosque de floresta ombrófila, em local úmido. São fornecidas descrições, chave de identificação, além de comentários sobre distribuição geográfica, hábitat e variação morfológica das espécies. Abstract This manuscript presents the floristic treatment of Campanulaceae in the Itatiaia National Park. The family is represented in the area by three genera, Lobelia with two species: L. camporum and L. fistulosa, Siphocampylus, with four species: S. duploserratus, S. longepedunculatus, S. umbellatus e S. westinianus, and Wahlenbergia, with one species: W. linarioides. The species were mostly found in the upper part of the park, usually in the rocky outcrops of the high fields, surrounded by graminoid vegetation. Only S. longepedunculatus was collected in rain forest understory, in moist places. Descriptions, an identification key, as well as comments on geographic distribution, habitat and morphological variation of species are provided.

A 3.7 kilobase region of chloroplast DNA that includes atpB, rbcL, and their intergenic spacer was sequenced in 61 samples from 45 species of South American Lobeliaceae plus two outgroup samples from Australia. A clade of four hexaploid Lobelia species from Chile is sister to a clade comprising Lysipomia, Siphocampylus, Centropogon, and Burmeistera. Lysipomia is a monophyletic group of small cushion-forming plants endemic to the high Andes, and is sister to the clade comprising the remaining three shrubby genera, which are most diverse in the Andes, but also extend to Central America, Mexico, and the West Indies. Siphocampylus has capsular fruit and is inferred to be paraphyletic relative to fleshy-fruited Centropogon and Burmeistera, but fleshy fruits have evidently evolved repeatedly, making Centropogon polyphyletiC. Burmeistera is primarily bat-pollinated and monophyletic, having evolved from one group of species in Centropogon. The phylogenetic relationships within Burmeistera indicate that t...

Relationships within the subfamily Lobelioideae in Campanulaceae are inferred from DNA sequence variation in the rbcL and ndhF genes, and the trnL-F region including the trnL intron and the trnL-F intergenic spacer. Results derived from Bayesian and parsimony analyses provide evidence for the long-suspected paraphyly of the genus Lobelia, comprising over 400 species as presently circumscribed. The perennial dwarf herbs belonging to the Andean genus Lysipomia are sister to a group comprising the Neotropical shrubs Burmeistera, Centropogon, and Siphocampylus. Giant lobelioids from the Hawaiian Islands, Brazil, Africa, and Sri Lanka form a strongly supported group. Character optimizations on the phylogenetic tree reveal that shifts in fruit types and lignification have occurred much more commonly than generally assumed. The main clades in the subfamily are outlined, which largely contradict previous classifications based on morphology.

The Campanulaceae are represented in Peru by 11 genera and around 180 species (Brako & Zarucchi, 1993; Ulloa Ulloa et al., 2004), mainly shrubs and herbs. Here we recognize 32 endemic species in three genera. Siphocampylus and Centropogon account for the largest number of endemic species. Endemic Campanulaceae are found mostly in the Very Humid Montane Forests region, between 800 and 3600 m elevation. We applied IUCN categories and criteria to 23 species. Nine endemic species have been recorded within the Peruvian parks system., La familia Campanulaceae es reconocida en el Perú por presentar 11 géneros y alrededor de 180 especies (Brako & Zarucchi, 1993; Ulloa Ulloa et al., 2004), principalmente arbustos y hierbas. En este trabajo reconocemos 32 especies endémicas en tres géneros, incluidas en su mayoría en los géneros Siphocampylus y Centropogon. Los taxones endémicos se encuentran principalmente en la región Bosques Muy Húmedos Montanos, entre los 800 y 3600 m de altitud. Se aplicaron las categorías y criterios de la UICN a 23 especies. Nueve especies endémicas se encuentran representadas dentro del Sistema Nacional de Áreas Naturales Protegidas por el Estado.

Siphocampylus is a neotropical genus that comprises 221 species distributed from Costa Rica to Argentina and in the Greater Antilles. Twenty-eight species have been reported from Brazil, mainly occupying mountainous terrain. The floral venation patterns and the origin of the hypanthium in eight Brazilian species, including three varieties, are described. Eleven ovarian vascular bundles depart from the siphonostele or receptacular stele: five of these bundles result from sepalar and staminal adnation and are alternate to five petalar bundles; the remaining bundle is central carpellary. The staminal bundles diverge from the sepalar bundles at the sinus, while the carpellary bundles form a cross, resulting in four ventral bundles; two of these feed the ovules; the other two feed the style. Apparently, the dorsal carpellary bundles diverge at the same site and then ramify profusely. The venation pattern observed is unprecedented in Siphocampylus and is quite different from other reports on genera of Campanulaceae. Further, these findings suggest that the origin of the hypanthium is appendicular, increasing knowledge of venation in this group, thus providing data for phylogenetic considerations.

The morphology of branched foliar trichomes in Lobelioideae (Campanulaceae) was studied via scanning electron microscopy. Trichomes of this type are found only in the Neotropical endemics Burmeistera, Centropogon, and Siphocampylus, and in the Hawaiian endemic Cyanea. The 37 species examined included all species with branched trichomes in Burmeistera, Centropogon sect. Burmeisteroides, Centropogon sect. Niveopsis, Centropogon sect. Wimmeriopsis, Cyanea, and Siphocampylus, plus about one- fourth of the 100 species comprising Centropogon sect. Siphocampyloides. Trichomes observed conformed to eight structural types. Dendritic, multangulate, multangulate-dendritic, echinoid, and furcate hairs were found only in Centropogon sect. Siphocampyloides, Cyanea, and Siphocampylus. Echinoid-dendritic hairs were confined to Centropogon sect. Burmeisteroides, stellate hairs to Centropogon sect. Wimmeriopsis, and multiseriate- multangulate hairs to Burmeistera and Centropogon sect. Niveopsis. Within Centropogon, the distribution of trichome types is consistent with the currently accepted classification of the genus. Correlations with other data, both morphological and molecular, suggest that branched trichomes have evolved in the subfamily repeatedly, perhaps as many as 11 times.

 Premise of the study: The species-rich Neotropical genera Centropogon , Burmeistera , and Siphocampylus represent more than half of the ~1200 species in the subfamily Lobelioideae (Campanulaceae). They exhibit remarkable morphological variation in fl oral morphology and habit. Limited taxon sampling and phylogenetic resolution, however, obscures our understanding of relationships between and within these genera and underscores our uncertainty of the systematic value of fruit type as a major diagnostic character.  Methods: We inferred a phylogeny from fi ve plastid DNA regions ( rpl32 - trnL , ndhF - rpl32 , rps16 - trnK , trnG - trnG - trns , rbcL ) using maximum-likelihood and Bayesian inference. Ancestral character reconstructions were applied to infer patterns of fruit evolution.  Key results: Our results demonstrate that the majority of species in the genera Centropogon , Burmeistera , and Siphocampylus together form a primarily mainland Neotropical clade, collectively termed the “centropogonids.” Caribbean Siphocampylus , however, group with other Caribbean lobelioid species. We fihigh support for the monophyly of Burmeistera and the polyphyly of Centropogon and mainland Siphocampylus . The ancestral fruit type of the centropogonids is a capsule; berries have evolved independently multiple times.  Conclusions: Our plastid phylogeny greatly improves the phylogenetic resolution within Neotropical Lobelioideae and highlights the need for taxonomic revisions in the subfamily. Inference of ancestral character states identifi es a dynamic pattern of fruit evolution within the centropogonids, emphasizing the diffi culty of diagnosing broad taxonomic groups on the basis of fruit type. Finally, we identify that the centropogonids, Lysipomia , and Lobelia section Tupa form a Pan-Andean radiation with broad habitat diversity. This clade is a prime candidate for investigations of Neotropical biogeography and morphological evolution.

Recent surveys of seed coat morphology in Lobelioideae (Campanulaceae) have demonstrated the systematic utility of such data in the subfamily and led to a revision of the supraspecific classification ofLobelia. Expanding upon these studies, we examined via scanning electron microscopy 41 seed accessions, emphasizing lobelioid genera in which only one or no species had been examined. Most conformed to previously described testal patterns. However, five species of the endemic Hawaiian genus Cyanea, comprising the molecularly defined Hardyi Clade, had a unique testal pattern (here termed Type F), characterized by laterally compressed, almost linear, areoles with rounded, knob-like protuberances on the radial walls at opposite ends. This offered a convenient synapomorphy for recognition of a clade originally defined on a molecular basis. A second unique testal pattern was found in the related Hawaiian endemics Brighamia and Delissea, thus supporting their close relationship. In this type (here termed Type G), the seed coat is irregularly wrinkled (rugose), creating broad, rounded ridges that run more-or-less perpendicular to the long axis of the seed and thus to the long axis of the testal cells. Seed coat morphology also supported the monophyly of all 124 species of Hawaiian Lobelioideae and their probable derivation from Asian species of Lobelia subg. Tupa. Additional studies supported close relationships between (1) the neotropical genera Centropogon and Siphocampylus; (2) the western American genera Legenere and Downingia; and (3) Jamaican Hippobroma and Lobelia sect. Tylomium, a group endemic to the West Indies.

Summary In the present, final part of a series of contributions on nectaries, further, miscellaneous case histories are presented because of their histological or ecological interest. In the special section, extrafloral and floral nectaries of members of six families are treated. A general survey concludes the paper. Extranuptial glands attracting ant guards are associated with the flowers of Becium and Ocimum (Lamiaceae); they represent novel cases in a family hitherto believed to lack this kind of organs. The glands develop in an unusual way by transformation of abscission scars of floral bracts. The five floral but extranuptial glands on the calyx of Siphocampylus betulifolius (Lobeliaceae), with a similar function, are unusual in sharing the same histological structure and drainage via stomatal sap-holes with the closely adjoining nuptial nectary, a disk. Three case histories refer to zoophilous unisexual flowers with only one of the functional sexual morphs rewarding with nectar. In Pachysandra and Sarcococca (monoecious Buxaceae) the male flowers are nectariferous. The nectarless pistillate flowers are inadvertently pollinated by bees while these exploit the staminate flowers. Two other instances, although heterogeneous in detail, exhibit the reverse situation in that only the pistillate flowers yield nectar: the male capitula of the dioecious composite Heterothalamus alienus are girdled by female-sterile ray florets with copious nectar; bees taking it become dusted with pollen when they move over the dry staminate disk florets. They may subsequently pollinate the female heads, which entirely consist of nectariferous pistillate ray florets. In the hummingbird-pollinated Begonia ferruginea it is also the pistillate flowers that contain a presumably nectareous fluid, while the dry staminate ones fallaciously attract the visitors by mimicking the pistillate flowers. Although basically epigynous, the protandrous revolver flowers of the little-known genus Pentaphragma (Pentaphragmataceae)have maintained the subbasal position of their five separate nectaries on the ovary wall, the nectar being accessible to bees via five receptacular channels. In the concluding chapter, possible evolutionary pathways leading to floral nectaries, particularly floral disks, are discussed. Hydathodes of vegetative provenance rather than extrafloral nectaries are likely precursors of most floral nectaries. Developmental modules comprising a single stomatal sap-hole or groups of them, associated with peri stomatal gland cells, may have aggregated in flowers to fonn disks. Separate nectariferous scales as found in flowers of the Cucurbitaceae, Crassulaceae and the Geraniales also reveal similarities with the respective leaf hydathodes. The organophylesis of the single scale seated in the axial spur of Pelargonium is detailed as an example. A clustered distribution of stomatal sap-holes on disks can be partly interpreted as revealing their origin from meristic hydathodal modules. Examples of the diversity of these patterns are given, which are little explored and may provide useful taxonomic characters.

This contribution presents and describes Siphocampylus baccae and S. sevegnaniae, two narrowly endemic new species from waterfalls and rocky cliffs in Santa Catarina state, southern Brazil. We assess the conservation status of both species as critically endangered (CR) according to the IUCN criteria. We present descriptions, photographs, a distribution map and taxonomic comments on the new species along with an identification key to the species of Siphocampylus that occur in Santa Catarina.

Most species of the neotropical genusSiphocampylus are believed to be bird-pollinated. The pollination biology ofSiphocampylus sulfureus was studied in a montane region in SE Brazil. This species has features intermediate between ornithophilous and chiropterophilous syndromes: it presents a striking combination of yellowish flowers with strong odour (chiropterophilous features), and diurnal anthesis and sucrose-dominated nectar (ornithophilous features). Major pollinators were hummingbirds by day, and a phyllostomid bat by night.Siphocampylus sulfureus may be viewed as a recent derivate from the presumed ornithophilous stock within sect.Macrosiphon, and thus benefits from the activity of both diurnal and nocturnal vertebrate pollinators.

From Siphocampylus verticillatus , two novel piperidine alkaloids have been isolated, namely meso -8,10-di- n -propyllobelidiol[ meso -2,6-bis(2′-hydroxypentyl)- N -methylpiperidine] and 8-ethyl-10- n -propyllobelidiol[6-(2′-hydroxybutyl-2-(2″-hydroxypentyl)- N -methylpiperidine].

Almost 11% of the 400 unidentified specimens of South American Lobelioideae (Campanulaceae) examined recently cannot be equated with any known species. They are described here as 17 new species: 6 of Burmeistera, 6 of Centropogon, and 5 of Siphocampylus. Several are characterized by morphological characters or combinations of characters that are unique or unusual in their respective genera or in the subfamily as a whole. Six of the novelties are endemic to Ecuador, four to Peru7 three each to Venezuela and Colombia, and one to Bolivia.

Siphocampylus platysiphon (Campanulaceae: Lobelioideae) is described as a new species from northern Peru. It is a member of subsect.Megastomi in sect.Brachysiphon and is most closely related toS. giganteus, a species common to the north in Ecuador and Colombia.