Brocchinia

Life-forms

Classification System: APG IV

Superregnum: Eukaryota
Regnum: Plantae
Cladus: Angiosperms
Cladus: Monocots
Cladus: Commelinids
Ordo: Poales

Familia: Bromeliaceae
Subfamilia: Brocchinioideae
Genus: Brocchinia
Species: B. acuminata – B. amazonica – B. cataractarum – B. cowanii – B. delicatula – B. gilmartiniae – B. hectioides – B. hitchcockii – B. maguirei – B. melanacra – B. micrantha – B. paniculata – B. prismatica – B. reducta – B. rupestris – B. serrata – B. steyermarkii – B. tatei – B. vestita – B. uaipanensis – B. wurdackiana

Name

Brocchinia Schult. & Schult. f., Syst. Veg. 7: 1250 (1830)
Type species: Brocchinia paniculata Schult. f. Syst. Veg. (ed. 15 bis) 7(2): 1250. (1830)

Synonyms

Heterotypic
Sequencia Givnish, Aliso 23: 18 (2007). This is not definitive see Navioideae Hams (1929) and Givnish (et al.) (2007 and 2011).

Note: Givnish et al., (2007) subsumed Ayensua into Brocchinia as Brocchinia uaipanensis, but this is not accepted by Govaerts et al. (2015).
References

Schultes, J.A. & Schultes, J.H. in J. J. Roemer & J. A. Schultes}} (1830) Systema Vegetabilium 7(2): lxx, 1250.
Gilmartin, A.J. (1988) Phylogenetic relationships of groups of genera within the subfamily Pitcairnioideae (Bromeliaceae). Systematic Botany, 13(2): 283–293.
Benzing, D.H., Givnish, T.J., and Bermudes, D. (1985) Absorptive trichomes in Brocchinia reducta (Bromeliaceae) and their evolutionary and systematic significance. Systematic Botany, 10(1): 81–91.
Givnish, T.J., Burkhardt, E.L., Happel, R.E. & Weintraub, J.D. (1984) Carnivory in the bromeliad Brocchinia reducta, with a cost/benefit model for the general restriction of carnivorous plants to sunny, moist, nutrient-poor habitats. American Naturalist 124: 479–497.
Givnish, T.J., Millam, K.C., Berry, P. E. & Sytsma, K. J. 2007. Phylogeny, adaptive radiation, and historical biogeography of Bromeliaceae inferred from ndhF sequence data. Aliso 23(1): 3–26. DOI: 10.5642/aliso.20072301.04 PDF Reference page.
Givnish, T.J., Barfuss, M.H.J., Van Ee, B., Riina, R., Schulte, K., Horres, R., Gonsiska, P.A., Jabaily, R.A., Crayn, D.M., Smith, J.A.C., Winter, K., Brown, G.K., Evans, T.M., Holst, B.K., Luther, H., Till, W., Zizka, G., Berry, P.E. & Sytsma, K.J. 2011. Phylogeny, adaptive radiation, and historical biogeography in Bromeliaceae: Insights from an eight-locus plastid phylogeny. American Journal of Botany 98(5): 872–895. DOI: 10.3732/ajb.1000059. PDF Reference page.
Govaerts, R. et al. 2015. Brocchinia in World Checklist of Selected Plant Families. The Board of Trustees of the Royal Botanic Gardens, Kew. Published on the internet. Accessed: 2015 July 14. Reference page. Includes Sequencia (2007) as a synonym.
Tropicos.org 2014. Brocchinia. Missouri Botanical Garden. Published on the internet. Accessed: 2014 Sept. 20. Notes Sequencia (2007) as incertae sedis
International Plant Names Index. 2014. Brocchinia. Published online. Accessed: Sept. 20. 2014.

Brocchinia is a genus of the botanical family Bromeliaceae, and is the sole genus of the subfamily Brocchinioideae, containing 20 species.[2][3] The genus is named for Giovanni Battista Brocchi, Italian naturalist (1772–1826). Brocchinia species are native primarily to the ancient Guayana Shield in southern Venezuela and Guyana, with some species extending into Colombia and northern Brazil.[1] Its species are generally restricted to areas of sand and sandstone of the Roraima Formation; a few occur on granite.[4]

Based on chloroplast DNA sequence variation, Brocchinia appears to be sister to all other bromeliads.[5] Calibration of the molecular family tree of bromeliads against the known ages of various fossil monocots suggests that Brocchinia lineage diverged from other bromeliads nearly 20 million years ago, and that some of the living species of Brocchinia began diverging from each other soon thereafter. The next lineage to diverge from other bromeliads included Lindmania and Connellia, which are also endemic to the Guayana Shield; at least one other subfamily Navioideae is also almost entirely restricted to that region.[5] Consequently, it is quite likely that the family as a whole arose in the Guayana Shield. Previously, Brocchinia was thought to be a member of the bromeliad subfamily Pitcairnioideae, based on its possession of winged seeds like those seen in other members of that group.[6][7][8] This view has now been overturned after the recognition that Pitcairnioideae as originally circumscribed more than a century ago represents several morphologically similar but independently derived groups that share only ancestral (plesiomorphic) characters, not true synapomorphies (shared derived characters).[5]

Brocchinia has undergone a spectacular adaptive radiation in mechanisms of nutrient capture, apparently in response to the unusually infertile, heavily leached substrates of the Guayana Shield.[9] At least two species, Brocchinia reducta and B. hechtioides, appear to be carnivorous.[9] Like most species of Brocchinia – and indeed, of many other bromeliad genera – these two species impound rain water in a tank formed by tightly overlapping leaves arranged in a basal rosette. In these species, however, the leaves are nearly vertical, their inner surfaces are covered with fine wax that readily exfoliates, the tank fluid is highly acid (pH c. 3.0) and emits a sweet, nectar-like odor, and the tank itself is full of the dead remains of ants (B. reducta) or bees and wasps (B. hechtioides). Recently, it has been shown that at least B. reducta secretes a phosphatase into its tanks.[10]

Another species, Brocchinia acuminata, is ant-fed myrmecophyte, apparently depending in part on nutrients and dead nestmates dropping into the tank from ants that live among the swollen, achlorophyllous leaf bases. The facultative epiphyte B. tatei – together with the tree-like B. micrantha with its massive, gutter-like leaf axils that hold liters of rainwater – captures a great deal of falling vegetable debris. One terrestrial population of B. tatei was discovered with heterocystous cyanobacteria in its tanks, suggesting nitrogen fixation.[9]

The earliest divergent members of the genus – including Brocchinia prismatica in the Prismatica clade, and such species as B. melanacra and B. vestita in the Maguirei clade[9] – lack tanks entirely and appear to depend solely on soil nutrients. Acquisition of the tank habit seems likely to have been the key innovation driving the evolution of specialized mechanisms of nutrient capture in Brocchinia, and is associated with thin leaf cross-sections and the later evolution of carnivory, ant-fed myrmecophily, epiphytism, and N fixation in the Acuminata and Reducta clades.[9] Each of the nutritionally specialized species have relatively large areas of live trichomes on the leaf bases that can absorb amino acids at high rates. Tanks and absorptive trichomes were later lost secondarily in Brocchinia steyermarkii, a terrestrial species common in wet sandy areas in the Gran Sabana.[11] Brocchinia melanacra is especially adapted to ground fires, with highly sclerotized leaf tips that protect that single bud in unexpanded leaves but appear to be useless (often dangling limply in the breeze) in fully expanded leaves. Brocchinia serrata, a highly aberrant taxon with tough, serrate leaves that is found only on a few mesetas in Colombia, has now been shown to be completely unrelated and has been described as the sole member of a new genus Sequencia, with its name reflecting its initial recognition based on DNA sequence data.[5] The ecology of the dwarf species – Brocchinia cataractarum, B. delicatula, B. rupestris – remain enigmatic.
Species

Brocchinia acuminata L.B. Smith
Brocchinia amazonica L.B. Smith
Brocchinia cataractarum (Sandwith) B. Holst
Brocchinia cowanii L.B. Smith
Brocchinia delicatula L.B. Smith
Brocchinia gilmartiniae G.S. Varadarajan
Brocchinia hechtioides Mez
Brocchinia hitchcockii L.B. Smith
Brocchinia maguirei L.B. Smith
Brocchinia melanacra L.B. Smith
Brocchinia micrantha (Baker) Mez
Brocchinia paniculata Schultes f.
Brocchinia prismatica L.B. Smith
Brocchinia reducta Baker
Brocchinia rupestris (Gleason) B. Holst
Brocchinia serrata L.B.Sm
Brocchinia steyermarkii L.B. Smith
Brocchinia tatei L.B. Smith
Brocchinia uaipanensis (Maguire) Givnish
Brocchinia vestita L.B. Smith
Brocchinia wurdackiana B. Holst

References

Kew World Checklist of Selected Plant Families
Givnish, Thomas J.; Barfuss, Michael H. J.; Van Ee, Benjamin; Riina, Ricarda; Schulte, Katharina; Horres, Ralf; Gonsiska, Philip A.; Jabaily, Rachel S.; Crayn, Darren M.; Smith, J. Andrew C.; Winter, Klaus (May 2011). "Phylogeny, adaptive radiation, and historical biogeography in Bromeliaceae: Insights from an eight-locus plastid phylogeny". American Journal of Botany. 98 (5): 872–895. doi:10.3732/ajb.1000059. hdl:2027.42/142109. PMID 21613186.
Gouda, E.J. & Butcher, D. (cont. updated) A List of Accepted Bromeliaceae Names [http://bromeliad.nl/bromNames/]. University Botanic Gardens, Utrecht (accessed: 20-12-2019).
Givnish TJ, Burkhardt EL, Happel RE, Weintraub JW. 1985. Carnivory in the bromeliad Brocchinia reducta, with a cost/benefit model for the general restriction of carnivorous plants to sunny, moist, nutrient-poor habitats. American Naturalist 124: 479-497.
Givnish TJ, Millam KC, Berry PE, Sytsma KJ. 2007. Phylogeny, adaptive radiation, and historical biogeography of Bromeliaceae inferred from ndhF sequence data. Aliso 23: 3-26.
Gilmartin AJ. 1988. Phylogenetic relationships of groups of genera within the subfamily Pitcairnioideae (Bromeliaceae). Systematic Botany, 13: 283-293.
Benzing DH. 1980. The Biology of the Bromeliads. California: Mad River Press.
Benzing DH. 2000. Bromeliaceae: Portrait of an Adaptive Radiation, Cambridge University Press, New York.
Givnish TJ, Sytsma KJ, Smith JE, Hahn WJ, Benzing DH, Burkhardt EL. 1997. Molecular evolution and adaptive radiation in Brocchinia (Bromeliaceae: Pitcairnioideae) atop tepuis of the Guayana Shield. Pp. 259-311 in TJ Givnish and KJ Sytsma, "Molecular Evolution and Adaptive Radiation", Cambridge University Press
Plachno BJ, Jankun A. 2005. Phosphatase activity in glandular structures of carnivorous plant traps, International Botanical Congress Vienna, P1716, The Jagiellonian Univ., Inst. of Botany, Dept. of Plant Cytology and Embryology, Krakow, Poland.
Benzing DH, Givnish TJ, Bermudes D. 1985. Absorptive trichomes in Brocchinia reducta (Bromeliaceae) and their evolutionary and systematic significance. Systematic Botany, 10(1): 81-91.