Fourleaf manyseed Polycarpon tetraphyllum, copyright Forest & Kim Starr.

Belongs within: Caryophyllales.
Contains: Polycarpaea, Scleranthus, Minuartia, Stellaria, Colobanthus, Spergularia, Cerastium, Sagina, Arenaria, Silene, Dianthus.

The Caryophyllaceae are a cosmopolitan family of herbaceous plants, most diverse in temperate regions. Familiar representatives include Dianthus (pinks), Stellaria (chickweeds) and Gypsophila (baby's breath). Some species have become widespread as pasture weeds. In members of the subfamily Illecebroideae, stipules are present on the leaves; other subfamilies usually lack stipules. Illecebroideae also differ from other subfamilies in having fruits as utricles rather than capsules. The Caryophylloideae have flowers with sepals more or less fused at the base, whereas Minuartoideae and Illecebroideae have free sepals.

Characters (from Allan 1961): Annual to perennial herbs; leaves usually simple, entire, opposite, often connate, or with scarious stipules. Flowers in simple to compound cymes, or solitary; usually perfect, hypogynous or perigynous; sepals 4-5, free or joined below; petals as many as sepals, often small or absent, free, often bifid or emarginate; stamens up to 10, anthers dehiscing longitudinally; ovary one-celled, at least in upper part, ovules 1 to many on basal or free-central placenta, embryo usually curved; fruit usually a dry capsule opening by teeth or valves.


<==Caryophyllaceae [Caryophyllineae, Illecebraceae, Paronychieae]
    |--Illecebroideae [Illecebraceae, Paronychoideae] T00
    |    |--Achyronychia cooperi H93
    |    |--Scopulophila rixfordii H93
    |    |--Cardionema ramosissimum H93
    |    |--Scleranthus A61
    |    |--Herniaria C06
    |    |    |--H. alpina C55a
    |    |    `--H. hirsuta PT98
    |    |         |--H. h. ssp. hirsuta H93
    |    |         `--H. h. ssp. cinerea [incl. Paronychia pusilla] H93
    |    `--Paronychia H93
    |         |--P. ahartii H93
    |         |--P. argentea C74
    |         |--P. brasiliana H90
    |         |--P. echinulata H93
    |         `--P. franciscana H93
    |--Minuartioideae [Alsineae, Alsinoideae] T00
    |    |--Minuartia KB96
    |    |--Stellaria C06
    |    |--Colobanthus C06
    |    |--Spergularia C06
    |    |--Cerastium H93
    |    |--Pseudostellaria jamesiana [=Stellaria jamesiana] H93
    |    |--Holosteum umbellatum H93
    |    |--Sagina H93
    |    |--Arenaria H93
    |    |--Moehringia macrophylla [=Arenaria macrophylla] H93
    |    |--Loeflingia squarrosa H93
    |    |    |--L. s. var. squarrosa [incl. L. pusilla] H93
    |    |    `--L. s. var. artemisiarum H93
    |    |--Moenchia H93
    |    |    |--M. erecta H93
    |    |    `--M. mantica Y98
    |    |--Alsine C55b
    |    |    |--A. biflora [=Stellaria biflora] C55b
    |    |    |--A. media C55b
    |    |    `--A. stricta [=Spergula stricta; incl. Arenaria uliginosa] C55b
    |    |--Spergula H93
    |    |    |--S. arvensis C55b
    |    |    |--S. fallax VB02
    |    |    |--S. pentandra C06
    |    |    `--S. saginoides C55a
    |    `--Polycarpon H93
    |         |--P. alsinaefolium C55b
    |         |--P. depressum H93
    |         |--P. peploides C74
    |         |--P. prostratum P03
    |         `--P. tetraphyllum (Linnaeus) Linnaeus 1759 PL04 [=Mollugo tetraphylla H90]
    `--Caryophylloideae [Silenoideae] T00
         |         Agrostemma H93
         |           |--A. aestivalis BO08
         |           |--A. flammea BO08
         |           `--A. githago JK80
         |         Petrorhagia H93
         |           |--P. armerioides (Seringe) Ball & Heywood 1964 PL04
         |           |--P. dubia [incl. Kohlrauschia velutina] H93
         |           |--P. nanteuilii H93
         |           |--P. prolifera GK00 [=Dianthus prolifer C06, Tunica prolifera H93]
         |           `--P. velutina GK00
         |         Velezia rigida H93
         |         Vaccaria H93
         |           |--V. hispanica [incl. V. segetalis] H93
         |           `--V. pyramidata H90
         |--Sileneae C06
         |    |--Silene C06
         |    `--Gypsophila C06
         |         |--G. australis H90
         |         |--G. cerastioides O88
         |         |--G. elegans H93
         |         |--G. paniculata H93
         |         |--G. petraea H09
         |         |--G. repens S96
         |         |--G. scorzonerifolia H93
         |         `--G. tubulosa C06
         `--Caryophylleae M99
              |--Caryophyllus aromaticus C55b
              |--Dianthus M99
              |--Saponaria C06
              |    |--S. caespitosa R84
              |    |--S. calabrica H90
              |    |--S. officinalis C55b
              |    `--S. vaccaria C55b
              `--Lychnis C06
                   |--L. alpina C55b
                   |--L. chalcedonica H90
                   |--L. coronaria C06
                   |--L. dioica R13
                   |--L. flos-cuculi BN99
                   |--L. githago [incl. Githago segetum] C06
                   `--L. vespertina C06

Caryophyllaceae incertae sedis:
  Geocarpon T00
  Dicheranthus YY22
  Melandrium YY22
    |--M. album GM96
    `--M. magellanicum D03
  Thylacospermum caespitosum O88
  Drymaria O88
    |--D. cordata O88
    |    |--D. c. ssp. cordata H90
    |    `--D. c. ssp. diandra H90
    `--D. villosa D07
  Lepyrodiclis holosteoides O88
  Bolanthus graecus (von Schreber) Barkoudah 1962 PL04
  Viscaria alpina KB96
  Corrigiola litoralis GK00
  Schiedea verticillata O94
  Polycarpaea LK14
  Hantsia pulchra (Chandler) Chandler 1960 CBH93
  Caryophyllidites polyoratus Muller 1970 CBH93

*Type species of generic name indicated


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[BO08] Bönsel, D., I. Ottich, A. Malten & G. Zizka. 2008. An updated list of the vascular plants of Frankfurt am Main (Pteridophyta & Spermatophyta). Senckenbergiana Biologica 88 (1): 111–121.

[BN99] Bungener, P., S. Nussbaum, A. Grub & J. Fuhrer. 1999. Growth response of grassland species to ozone in relation to soil moisture condition and plant strategy. New Phytologist 142: 283–293.

[C55a] Candolle, A. de. 1855a. 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.

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

[GK00] Gibson, N., & G. J. Keighery. 2000. Flora and vegetation of the Byenup-Muir reserve system, south-west Western Australia. CALMScience 3 (3): 323–402.

[GM96] Griffiths, A. J. F., J. H. Miller, D. T. Suzuki, R. C. Lewontin & W. M. Gelbart. 1996. An Introduction to Genetic Analysis 6th ed. W. H. Freeman and Company: New York.

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[H09] Heltmann, H. 2009. Der Königstein (Piatra Craiului), die Perle der Burzenländer Gebirge. Mauritiana 20 (3): 515–527.

[H93] Hickman, J. C. (ed.) 1993. The Jepson Manual: Higher Plants of California. University of California Press: Berkeley (California).

[JK80] John, J., & K.-P. Kolbe. 1980. The systematic position of the “Theales” from the viewpoint of serology. Biochemical Systematics and Ecology 8: 241–248.

[KB96] Konstantinou, M., & D. Babalonas. 1996. Metal uptake by Caryophyllaceae species from metalliferous soils in northern Greece. Plant Systematics and Evolution 203: 1–10.

[LK14] Lyons, M. N., G. J. Keighery, L. A. Gibson & T. Handasyde. 2014. Flora and vegetation communities of selected islands off the Kimberley coast of Western Australia. Records of the Western Australian Museum Supplement 81: 205–244.

[M99] Matthews, M. 1999. Heliothine Moths of Australia: A guide to bollworms and related noctuid groups. CSIRO Publishing.

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

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[R13] Reuter, O. M. 1913. Lebensgewohnheiten und Instinkte der Insekten bis zum Erwachen der sozialen Instinkte. R. Friedländer & Sohn: Berlin.

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[YY22] Yampolsky, C., & H. Yampolsky. 1922. Distribution of sex forms in the phanerogamic flora. Bibliotheca Genetica 3: 1–62.

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Last updated: 4 April 2020.

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