Boraginaceae

Cordia gerascanthus, from here.


Belongs within: Gentianidae.
Contains: Boragineae, Cynoglosseae, Lithospermeae, Ehretioideae, Heliotropioideae.

The Boraginaceae as recognised herein is a clade supported by molecular analyses (Långström & Chase 2002). It combines older concepts of the Boraginaceae with the Hydrophylloideae, previously treated as a separate family distinguished by the possession of capsular fruits. Some members of the family are grown as herbs (such as borage Borago officinalis) or ornamentals (such as forget-me-nots, Myosotis, and heliotropes, Heliotropium). Subfamilies within the Boraginaceae other than the Hydrophylloideae are primarily separated by characters of the gynoecium. Boraginoideae, for instance, have a deeply four-lobed ovary maturing to four nutlets and a basifixed style (Långström & Chase 2002).

Synapomorphies (from www.mobot.org): Plant more or less herbaceous; pyrrolizidine alkaloids; cork superficial to mid-cortical (pericyclic); sieve tubes with nuclear non-dispersive protein bodies; nodes 1:1; petiole bundle(s) arcuate; plant more or less roughly hairy, hairs with a basal cystolith or cystolith-like body, and/or walls calcified; leaves spiral; inflorescences terminal, cymose, cyme scorpioid; calyx free, corolla tube formation late; anther placentoid absent; nectary not vascularized; ovary with secondary septae, stigma dry; ovules 2/carpel, apical, pendulous; calyx persistent in fruit; micropylar endosperm haustoria present.

<==Boraginaceae [Ehretiaceae]
    |--Boraginoideae [Trigonotideae] LC02
    |    |  i. s.: Amphibologyne LC02
    |    |         Brachybotrys LC02
    |    |         Decalepidanthus LC02
    |    |         Mairetis LC02
    |    |         Mertensia LC02
    |    |           |--M. cryptocarpa D03
    |    |           `--M. maritima C55
    |    |         Moltkiopsis LC02
    |    |         Moritzia LC02
    |    |         Neatostema Johnston 1953 LC02, BR65
    |    |           `--N. apulum (Linnaeus) Johnston 1953 (see below for synonymy) BR65
    |    |         Omphalotrigonotis LC02
    |    |         Sinojohnstonia LC02
    |    |         Thaumatocaryum LC02
    |    |         Trigonocaryum LC02
    |    |         Zoelleria LC02
    |    |         Embadium Black 1931 BR65
    |    |           `--E. stagnense Black 1931 BR65
    |    |--Boragineae LC02
    |    |--Cynoglosseae LC02
    |    `--+--Lithospermeae LC02
    |       `--Echiochileae LC02
    |            |--Antiphytum hintoniorum LC02
    |            |--Ogastemma pusillum LC02
    |            `--Echiochilon [Echiochilinae] LC02
    |                 |--E. collenettei LC02
    |                 |--E. kotschyi [incl. Sericostoma pauciflorum var. qeshmensis Ghahreman & Attar 1996] LC02
    |                 `--E. pauciflorum (Stocks ex Wight) Långström & Chase 2002 (see below for synonymy) LC02
    `--+--Ehretioideae LC02
       |--Hydrophylloideae [Hydrophyllaceae] LC02
       |    |--Wigandia [Nameae] LC02
       |    |    |--W. caracasana LC02
       |    |    |--W. pruritiva J87
       |    |    `--W. urens MM96
       |    `--+--Hydrophylleae LC02
       |       |    |--Hydrophyllum canadense LC02
       |       |    `--Nemophila LC02
       |       |         |--N. insignis LC02
       |       |         `--N. maculata PT01
       |       `--Phacelieae LC02
       |            |--Emmenanthe penduliflora LC02
       |            `--Phacelia LC02
       |                 |--P. campanularia PT01
       |                 |--P. circinata D03
       |                 `--P. grandiflora LC02
       `--+--Heliotropioideae LC02
          `--Cordioideae [Cordiaceae] LC02
               |--Patagonula americana LC02
               `--Cordia LC02
                    |--C. alliodora HH03
                    |--C. anisodonta J87
                    |--C. borinquensis SWK87
                    |--C. dependens J87
                    |--C. dichotomum P03
                    |--C. gerascanthus HH03
                    |--C. gharaf PP07
                    |--C. lima J87
                    |--C. macrostachya LC02
                    |--C. myxa SR07
                    |--C. nitida SWK87
                    |--C. picardae J87
                    |--C. sonorae HH03
                    |--C. sulcata SWK87
                    `--C. trichotoma HH03

Boraginaceae incertae sedis:
  Chionocharis hookeri O88
  Lasiocaryum O88
    |--L. densiflorum O88
    |--L. diffusum O88
    `--L. munroi O88
  Microcaryum pygmaeum O88
  Wellstedia [Wellstedioideae] T00
  Tournefortia J87
    |--T. argentea H09
    |--T. bicolor J87
    |--T. glabra J87
    |--T. hirsutissima J87
    |--T. laurifolia C55
    `--T. selleana J87
  Messerschmidia argentea SJP60

Echiochilon pauciflorum (Stocks ex Wight) Långström & Chase 2002 [=Sericostoma pauciflorum Stocks ex Wight 1848] LC02

Neatostema apulum (Linnaeus) Johnston 1953 [=Myosotis apula Linnaeus 1753, Lithospermum apulum (Linnaeus) Vahl. 1938, Rhytispermum apulum (Linnaeus) Reichenbach 1858] BR65

*Type species of generic name indicated

REFERENCES

[BR65] Black, J. M., & E. L. Robertson. 1965. Flora of South Australia. Part IV. Oleaceae-Compositae. W. L. Hawes, Government Printer: Adelaide.

[C55] Candolle, A. de. 1855. Géographie Botanique Raisonée: Ou exposition des faits principaux et des lois concernant la distribution géographique des plantes de l’époque actuelle vol. 1. Librairie de Victor Masson: Paris.

[D03] Dusén, P. 1903. The vegetation of western Patagonia. In Reports of the Princeton University Expeditions to Patagonia, 1896-1899, vol. 8 – Botany (W. B. Scott, ed.) pp. 1-34. The University: Princeton (New Jersey).

[H09] Hedley, C. 1909. The Marine Fauna of Queensland: Address by the President of Section D. Australasian Association for the Advancement of Science: Brisbane.

[HH03] Hernández, J. R., & J. F. Hennen. 2003. Rust fungi causing galls, witches’ brooms, and other abnormal plant growths in northwestern Argentina. Mycologia 95 (4): 728-755.

[J87] Judd, W. S. 1987. Floristic study of Morne La Visite and Pic Macaya National Parks, Haiti. Bulletin of the Florida State Museum – Biological Sciences 32 (1): 1-136.

[LC02] Långström, E., & M. W. Chase. 2002. Tribes of Boraginoideae (Boraginaceae) and placement of Antiphytum, Echiochilon, Ogastemma and Sericostoma: A phylogenetic analysis based on atpB plastid DNA sequence data. Plant Systematics and Evolution 234: 137-153.

[MM96] Mound, L. A., & R. Marullo. 1996. The thrips of Central and South America: an introduction (Insecta: Thysanoptera). Memoirs on Entomology, International 6: 1-487.

[O88] Ohba, H. 1988. The alpine flora of the Nepal Himalayas: An introductory note. In The Himalayan Plants vol. 1 (H. Ohba & S. B. Malla, eds) The University Museum, University of Tokyo, Bulletin 31: 19-46.

[PP07] Pandey, R. P. & P. M. Padhye. 2007. Studies on phytodiversity of Arid Machia Safari Park-Kailana in Jodhpur (Rajasthan). Bulletin of the Botanical Survey of India 49 (1-4): 15-78.

[PT01] Pemberton, L. M. S., S.-L. Tsai, P. H. Lovell & P. J. Harris. 2001. Epidermal patterning in seedling roots of eudicotyledons. Annals of Botany 87: 649-654.

[SJP60] St. John, H. & W. R. Philipson. 1960. List of the flora of Oeno Atoll, Tuamotu Archipelago, south-central Pacific Ocean. Transactions of the Royal Society of New Zealand 88 (3): 401-403.

[SR07] Sankar, R. V., K. Ravikumar, N. M. Ganesh Babu & D. K. Ved. 2007. Botany of Anapady MPCA, Palghat district, Kerala with special emphasis on species of conservation concern. Bulletin of the Botanical Survey of India 49 (1-4): 165-172.

[SWK87] Snyder, N. F. R., J. W. Wiley & C. B. Kepler. 1987. The Parrots of Luquillo: Natural history and conservation of the Puerto Rican parrot. Western Foundation of Vertebrate Zoology: Los Angeles.

[T00] Thorne, R. F. 2000. The classification and geography of the flowering plants: Dicotyledons of the class Angiospermae (subclasses Magnoliidae, Ranunculidae, Caryophyllidae, Dilleniidae, Rosidae, Asteridae, and Lamiidae). The Botanical Review 66: 441-647.

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