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Classification System: APG IV

Superregnum: Eukaryota
Regnum: Plantae
Cladus: Angiosperms
Cladus: Eudicots
Cladus: Core eudicots
Cladus: Asterids
Cladus: Lamiids
Ordo: Gentianales

Familia: Rubiaceae
Subfamilia: Rubioideae
Tribus: Spermacoceae
Genus: Arcytophyllum
Species: A. lavarum – A. nitidum – A. serpyllaceum
Name

Arcytophyllum Willd. ex Schult. & Schult.f.
Synonyms

Heterotypic
Anotis DC., Prodr. 4: 431 (1830)
Mallostoma H.Karst., Fl. Columb. 2: 9 (1862)
Pseudrachicallis H.Karst., Fl. Columb. 2: 10 (1862)
Teinosolen Hook.f. in G.Bentham & J.D.Hooker, Gen. Pl. 2: 61 (1873)
Ereicoctis (DC.) Kuntze, Revis. Gen. Pl. 1: 281 (1891)
Terrellianthus Borhidi, Acta Bot. Hung. 54: 46 (2012)

References

Govaerts, R. et al. 2020. Arcytophyllum in World Checklist of Selected Plant Families. The Board of Trustees of the Royal Botanic Gardens, Kew. Published on the internet. Accessed: 2020 Mar. 16. Reference page.

Arcytophyllum is a genus of flowering plants in the family Rubiaceae. The genus contains 18 species, distributed from New Mexico to Bolivia.[1][2][3]

Description

Members of this genus are shrubs or dwarf shrubs that occupy mountainous habitats. Their leaves are oppositely arranged, small, sessile or sub-sessile, and possess inconspicuous secondary venation. Flowers are tetra- or pentamerous, with the stamens adnate to the white or pink funnelform or salverform corolla. The fruit is a capsule with septicidal dehiscence.
Species

Arcytophyllum aristatum Standl. - Colombia, Ecuador
Arcytophyllum cachirense (H.Karst.) K.Schum. - Colombia
Arcytophyllum capitatum (Benth.) K.Schum. - Colombia, Ecuador
Arcytophyllum ciliolatum Standl. - Ecuador, Perú
Arcytophyllum ericoides (Willd. ex Roem. & Schult.) Standl. - Ecuador, Perú
Arcytophyllum fasciculatum (A.Gray) Terrell & H.Rob. - New Mexico, Texas, Coahuila
Arcytophyllum filiforme (Ruiz & Pav.) Standl. - Bolivia, Perú, Ecuador, Colombia
Arcytophyllum lavarum K.Schum. - Costa Rica, Panama
Arcytophyllum macbridei Standl. - Amazonas (of Perú)
Arcytophyllum muticum (Wedd.) Standl. - Costa Rica, Panama, Colombia, Ecuador, Venezuela
Arcytophyllum nitidum (Kunth) Schltdl. - Colombia, Venezuela
Arcytophyllum peruvianum (Wernham) J.H.Kirkbr. - Perú
Arcytophyllum rivetii Danguy & Cherm. - Ecuador, Perú
Arcytophyllum serpyllaceum (Schltdl.) Terrell - Guatemala, Chiapas, Veracruz, Oaxaca
Arcytophyllum setosum (Ruiz & Pav.) Schltdl. - Bolivia, Perú, Ecuador, Colombia
Arcytophyllum thymifolium (Ruiz & Pav.) Standl. - Colombia, Ecuador, Perú
Arcytophyllum venezuelanum Steyerm. - Venezuela
Arcytophyllum vernicosum Standl. - Ecuador, Peru

Phylogeny and biogeography

Arcytophyllum is a monophyletic genus when A. serpyllaceum is excluded, as this species has been shown to be more closely related to Bouvardia but has not been formally transferred to another genus.[4][5] The remaining species are most closely related to the North American and Caribbean genus Houstonia (including Stenaria) and one species of the polyphyletic genus Oldenlandia (Oldenlandia microtheca) distributed in Mexico and Central America, indicating that Arcytophyllum may be one of the few cases in the family Rubiaceae of a southward migration to reach the Andes.[4] The center of diversity is in South America; namely in Ecuador, where 10 species are documented.[6]
Floral morphology and evolution

Flowers of Arcytophyllum are distylous, meaning that two morphs exist differing in the vertical orientation of anthers and stigmas. Pin morphs have stigmas positioned above the anthers, while thrum morphs have stigmas positioned below the anthers. A study of 11 species showed significantly higher seed production in thrum than pin morphs of A. aristatum, A. lavarum, and A. vernicosum, and a similar tendency in all other sampled species.[7] In contrast, pin morphs of A. lavarum cross pollinate more efficiently than thrum morphs, thus transmitting more of their genes to the next generation via pollen than seed. As better exporters of pollen to flowers of the opposite morph, long-styled A. lavarum give rise to disassortative pollen flow and a division in reproductive functions between the morphs. If pollinators promote the condition of distyly and asymmetric pollen flow over evolutionary time, it is plausible that gender specialization will evolve into dioecy.[8]
References

"Arcytophyllum in the World Checklist of Rubiaceae". Retrieved 10 June 2014.
Biota of North America Program, Arcytophyllum fasciculatum
Willdenow, Carl Ludwig von in Schultes, Josef (Joseph) August & Schultes, Julius Hermann. 1827. Mantissa in volumen tertium:Systematis vegetabilium caroli a Linné 3: 108
Andersson, L.; Rova, J.H.E.; Guarin, F.A. (2002). "Relationships, circumscription, and biogeography of Arcytophyllum (Rubiaceae) based on evidence from cpDNA". Brittonia. 54 (1): 40–49. doi:10.1663/0007-196x(2002)054[0040:rcaboa]2.0.co;2.
Groeninckx, I.; Dessein, S.; Ochoterena, H.; Persson, C.; Motley, T.J.; Kårehed, J.; Bremer, B.; Huysmans, S.; Smets, E. (2009). "Phylogeny of the Herbaceous Tribe Spermacoceae (Rubiaceae) Based on Plastid DNA Data". Annals of the Missouri Botanical Garden. 96 (1): 109–132. doi:10.3417/2006201. S2CID 56042261.
Ulloa, C.U.; Jørgensen, P.M. "Arcytophyllum". Trees and shrubs of the Andes of Ecuador. Aarhus University Press. Retrieved 6 February 2013.
Wolff, D.; Leide-Schumann, S. (2007). "Evolution of flower morphology, pollen dimorphism, and nectar composition in Arcytophyllum, a distylous genus of Rubiaceae". Organisms, Diversity & Evolution. 7 (2): 106–123. doi:10.1016/j.ode.2006.02.005.
García-Robledo, C.; Mora, F. (2007). "Pollination Biology and the Impact of Floral Display, Pollen Donors, and Distyly on Seed Production in Arcytophyllum lavarum (Rubiaceae)". Plant Biology. 9 (4): 453–461. doi:10.1055/s-2007-964962. PMID 17401810.

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