Stand of silver birch Betula pendula, copyright Percita.

Belongs within: Faganae.

The Betulaceae, birches and related taxa, is a group of deciduous trees and shrubs with pinnately-veined, sharp-toothed, simple leaves found mostly in temperate regions of the Northern Hemisphere. Flowers are borne in unisexual inflorescences with staminate flowers forming elongate, pendulous catkins.

Characters (from Flora of North America): Trees and shrubs, deciduous. Winter buds stipitate or sessile, with either 2–3 valvate scales (stipules) or few to many imbricate scales (or occasionally naked); terminal bud absent. Leaves alternate, spirally arranged, 2–3-ranked, simple; stipules deciduous, distinct; petioles present. Leaf blade sometimes lobed, pinnately veined, margins toothed, serrate to nearly entire; surfaces glabrous to tomentose, abaxially often with resinous glands. Inflorescences unisexual; staminate catkins pendulous, elongate, cylindric, conspicuously bracteate, consisting of crowded, reduced, 1–3-flowered clusters; pistillate inflorescences either of erect to pendulous bracteate catkins, or of compact 2--3-flowered clusters subtended by leafy involucres; bracts often nearly foliaceous or woody in infructescences. Staminate flowers bracteate; stamens (1–)4–6; anthers 2-locular, dehiscing by longitudinal slits, pollen sacs often more or less distinct; pistillode sometimes present. Pistillate flowers small, highly reduced; pistil 1, 2(–3)-carpellate; ovary inferior, usually 2-locular proximally, 1-locular distally; placentation axile; ovules 1–2 per locule, pendulous; styles 2, distinct or nearly so; stigmas dry; staminodes usually absent. Fruits nuts, nutlets, or 2-winged samaras, 1-seeded, without multibracteate cupule, often subtended or enclosed by foliaceous hull developed from 2–3 bracts; seed coat membranous; endosperm present, thin at maturity; embryo straight, as long as seed; cotyledons flat or greatly thickened, oily.

Betulaceae [Betulanae, Betuloideae]
    `--Betuleae CBH93
         |--Alnus CBH93
         |    |--A. acuminata F11
         |    |--A. arguta BT87
         |    |--A. glutinosa WM09
         |    |--A. hirsuta M03
         |    |    |--A. h. var. hirsuta A80
         |    |    `--A. h. var. sibirica A80
         |    |--A. incana HP-W05
         |    |--A. japonica [=A. maritima var. japonica] LO98
         |    |--A. jorullensis WO92
         |    |--A. maccoyi Ettingshausen 1886 F71
         |    |--A. maximowiezii M03
         |    |--A. nepalensis D07
         |    |--A. rhombifolia H93
         |    |--A. rubra PHK96
         |    |--A. rugosa JVHN90
         |    |--A. sinuata WM09 [=A. viridis ssp. sinuata H93]
         |    |--A. tenuifolia ZB01 [=A. incana ssp. tenuifolia H93]
         |    `--A. viridis HP-W05
         `--Betula WB03
              |--B. alba WEE70
              |--B. alleghaniensis BFD01
              |--B. ermanii WB03
              |--E. glandulosa RJ11
              |--B. lenta CS05
              |--B. lutea S96
              |--B. mandshurica S00
              |--B. maximowicziana LO98
              |--B. nana K02
              |--B. nigra CB06
              |--B. occidentalis Z83
              |--B. papyrifera JVHN90
              |--B. pendula BS01
              |--B. platyphylla LO98
              |    |--B. p. var. platyphylla LO98
              |    `--B. p. var. japonica LO98
              |--B. populifolia JVHN90
              |--B. pubescens W01
              `--B. utilis O88

Betulaceae incertae sedis:
  Ostryopsis YY22
  Ostrya YY22
    |--O. carpinifolia SM08
    `--O. virginiana H03
  Carpinus SP04
    |--C. betulus SM08 [incl. C. caucasica SP04]
    |--C. caroliniana V72
    |--C. orientalis G98
    `--C. tschonoskii [incl. C. laxiflora] LO98
  Betuloxylon oligocenicum Kaiser 1880 CBH93
  Coryloxylon nemejcii Prakash et al. 1971 CBH93
  Alnipollenites CBH93
    |--A. eminens Miki 1977 CBH93
    |--A. trina (Stanley) Norton in Norton & Hall 1969 YB02
    `--A. verus (Potonié) Potonié 1960 YB02
  Betulaceoipollenites Jarzen & Norris 1975 CBH93

*Type species of generic name indicated


[A80] Aoki, J. 1980. A revision of the oribatid mites of Japan. I. The families Phthiracaridae and Oribotritiidae. Bulletin of the Institute of Environmental Science and Technology, Yokohama National University 6 (2): 1–89.

[BS01] Bannister, P., & G. L. Strong. 2001. The distribution and population structure of the temperate mistletoe Ileostylus micranthus in the Northern Cemetery, Dunedin, New Zealand. New Zealand Journal of Botany 39: 225–233.

[BFD01] Bowman, J., G. J. Forbes & T. G. Dilworth. 2001. The spatial component of variation in small-mammal abundance measured at three scales. Canadian Journal of Zoology 79: 137–144.

[CB06] Christopher, C. C., & G. W. Barrett. 2006. Coexistence of white-footed mice (Peromyscus leucopus) and golden mice (Ochrotomys nuttalli) in a southeastern forest. Journal of Mammalogy 87 (1): 102–107.

[CBH93] Collinson, M. E., M. C. Boulter & P. L. Holmes. 1993. Magnoliophyta (‘Angiospermae’). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 809–841. Chapman & Hall: London.

[CS05] Connors, M. J., E. M. Schauber, A. Forbes, C. G. Jones, B. J. Goodwin & R. S. Ostfeld. 2005. Use of track plates to quantify predation risk at small spatial scales. Journal of Mammalogy 86 (5): 991–996.

[D07] Dash, S. S. 2007. Useful plants of Kabi Sacred Grove, Sikkim. Bulletin of the Botanical Survey of India 49: 79–88.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1–167.

[F11] Fraga, R. M. 2011. Family Icteridae (New World blackbirds). In: Hoyo, J. del, A. Elliott & D. A. Christie (eds) Handbook of the Birds of the World vol. 16. Tanagers to New World Blackbirds pp. 684–807. Lynx Edicions: Barcelona.

[G98] Gruber, J. 1998. Beiträge zur Systematik der Gattung Dicranolasma (Arachnida: Opiliones, Dicranolasmatidae). I. Dicranolasma thracium Staręga und verwandte Formen aus Südosteuropa und Südwestasien. Annalen des Naturhistorischen Museum in Wien. Serie B. Botanik und Zoologie 100: 489–537.

[H03] Healy, R. A. 2003. Mattirolomyces tiffanyae, a new truffle from Iowa, with ultrastructural evidence for its classification in the Pezizaceae. Mycologia 95 (4): 765–772.

[HP-W05] Heitland, W., & H. Pschorn-Walcher. 2005. Biology and parasitoids of the peculiar alder sawfly, Platycampus luridiventris (Fallen) (Insecta, Hymenoptera, Tenthredinidae). Senckenbergiana Biologica 85 (2): 215–231.

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

[JVHN90] Jennings, D. T., W. M. Vander Haegen & A. M. Narahara. 1990. A sampling of forest-floor spiders (Araneae) by expellant, Moosehorn National Wildlife Refuge, Maine. Journal of Arachnology 18: 173–180.

[K02] Koponen, S. 2002. Ground-living spiders in bogs in northern Europe. Journal of Arachnology 30 (2): 262–267.

[LO98] Lack, H. W., & H. Ohba. 1998. Die Xylothek des Chikusai Kato. Willdenowia 28: 263–276.

[M03] Mockford, E. L. 2003. New species and records of Psocoptera from the Kuril Islands. Mitt. Mus. Nat.kd. Berl., Dtsch. Entomol. Z. 50 (2): 191–230.

[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.

[PHK96] Prescott, L. M., J. P. Harley & D. A. Klein. 1996. Microbiology 3rd ed. Wm. C. Brown Publishers: Dubuque (Iowa).

[RJ11] Rising, J. D., A. Jaramillo, J. L. Copete, P. G. Ryan & S. C. Madge. 2011. Family Emberizidae (buntings and New World sparrows). In: Hoyo, J. del, A. Elliott & D. A. Christie (eds) Handbook of the Birds of the World vol. 16. Tanagers to New World Blackbirds pp. 428–683. Lynx Edicions: Barcelona.

[SP04] Schneeweiss, G. M., T. Palomeque, A. l. E. Colwell & H. Weiss-Schneeweiss. 2004. Chromosome numbers and karyotype evolution in holoparasitic Orobanche (Orobanchaceae) and related genera. American Journal of Botany 91 (3): 439–448.

[SM08] Schönhofer, A. L., & J. Martens. 2008. Revision of the genus Trogulus Latreille: the Trogulus coriziformis species-group of the western Mediterranean (Opiliones: Trogulidae). Invertebrate Systematics 22: 523–554.

[S00] Siddiqi, M. R. 2000. Tylenchida: Parasites of plants and insects 2nd ed. CABI Publishing: Wallingford (UK).

[S96] Smith, D. R. 1996. Aulacidae (Hymenoptera) in the mid-Atlantic states, with a key to species of eastern North America. Proceedings of the Entomological Society of Washington 98 (2): 274–291.

[V72] Voss, E. G. 1972. Michigan Flora. Part I. Gymnosperms and Monocots. Cranbrook Institute of Science and University of Michigan Herbarium.

[WO92] Walter, D. E., & D. J. O'Dowd. 1992. Leaves with domatia have more mites. Ecology 73 (4): 1514–1518.

[WM09] Wang, H., M. J. Moore, P. S. Soltis, C. D. Bell, S. F. Brockington, R. Alexandre, C. C. Davis, M. Latvis, S. R. Manchester & D. E. Soltis. 2009. Rosid radiation and the rapid rise of angiosperm-dominated forests. Proceedings of the National Academy of Sciences of the USA 106 (10): 3853–3858.

[WB03] Wang, Q.-M., F.-Y. Bai, J.-H. Zhao & J.-H. Jia. 2003. Bensingtonia changbaiensis sp. nov. and Bensingtonia sorbi sp. nov., novel ballistoconidium-forming yeast species from plant leaves. International Journal of Systematic and Evolutionary Microbiology 53: 2085–2089.

[W01] Wohltmann, A. 2001. Closely related species of Parasitengonae (Acari: Prostigmata) inhabiting the same areas: features facilitating coexistence. In: Halliday, R. B., D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff (eds) Acarology: Proceedings of the 10th International Congress pp. 121–135. CSIRO Publishing: Melbourne.

[WEE70] Woodward, T. E., J. W. Evans & V. F. Eastop. 1970. Hemiptera (bugs, leafhoppers, etc.) In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 387–457. Melbourne University Press.

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

[YB02] Yi, S., & D. J. Batten. 2002. Palynology of Upper Cretaceous (uppermost Campanian-Maastrichtian) deposits in the South Yellow Sea Basin, offshore Korea. Cretaceous Research 23: 687–706.

[ZB01] Zhang, N., & M. Blackwell. 2001. Molecular phylogeny of dogwood anthracnose fungus (Discula destructiva) and the Diaporthales. Mycologia 93 (2): 355–365.

[Z83] Zink, R. M. 1983. Evolutionary and systematic significance of temporal variation in the fox sparrow. Systematic Zoology 32 (3): 223–238.

Last updated: 5 September 2020.


Japanese chinquapin Castanopsis cuspidata, copyright Kenpei.

Belongs within: Fagaceae.

Castanopsis, the chinquapins, is a genus of trees found in tropical and subtropical eastern Asia. Many species are valued for their wood but a number have been threatened by over-logging.

Characters (from Flora of China): Trees evergreen. Winter buds ovoid to ellipsoid, with decussate scales. Stipules extrapetiolar. Leaves alternate, distichous, or for a few species spirally arranged. Inflorescences usually unisexual, erect, spicate or paniculate. Male flowers in fascicles of 3-7, rarely solitary and scattered; perianth 5- or 6(-8)-lobed; stamens (8 or)9-12; rudimentary pistil very small, densely covered with curved woolly hairs. Female flowers solitary or in clusters of 3-5(-7) per cupule; staminodes when present opposite perianth lobes; ovary 3-loculed; styles (2 or)3(or 4); stigmas punctiform or shallow terminal pores. Cupules solitary on rachis, actinomorphic or zygomorphic, rarely indehiscent, completely or partially enclosing nut; bracts sparsely to densely covering outside of cupule, spinelike or rarely scalelike or tubercles (cupule measurement always includes bracts). Nuts 1-3 per cupule, maturing after 2nd year or rarely in 1st year; abortive ovule apical. Germination hypogeal; cotyledons slightly convex, rarely cerebriform rugose.

    |--C. acuminatissima C78
    |--C. benthami Ettingshausen 1883 F71
    |--C. cuspidata [=Quercus cuspidata] LO98
    |    |--C. c. var. cuspidata LO98
    |    `--C. c. var. sieboldii LO98
    |--C. densinervia H03
    |--C. endertii H03
    |--C. indica D07
    |--C. lancifolia D07
    |--C. sclerophylla LI89
    `--C. tribuloides D07

*Type species of generic name indicated


[C78] Clunie, N. M. U. 1978. The vegetation. In: Womersley, J. S. (ed.) Handbooks of the Flora of Papua New Guinea vol. 1 pp. 1–11. Melbourne University Press: Carlton South (Australia).

[D07] Dash, S. S. 2007. Useful plants of Kabi Sacred Grove, Sikkim. Bulletin of the Botanical Survey of India 49: 79–88.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1–167.

[H03] Heads, M. 2003. Ericaceae in Malesia: vicariance biogeography, terrane tectonics and ecology. Telopea 10 (1): 311–449.

[LO98] Lack, H. W., & H. Ohba. 1998. Die Xylothek des Chikusai Kato. Willdenowia 28: 263–276.

[LI89] Liang L.-R. & K. Ishikawa. 1989. Occurrence of Gamasellus (Acarina, Gamasida, Ologamasidae) on Tian-mu Mountains in east China. Reports of Research, Matsuyama Shinonome Junior College 20: 143–152.

Last updated: 5 September 2020.


European ash Fraxinus excelsior, copyright Matthieu Sontag.

Belongs within: Lamiales.

The Oleoideae contains the olive Olea europaea and related taxa in which the corolla has four lobes or is absent, and the ovule is pendulous from the summit of the cell (Black & Robertson 1965). Olea europaea is native to the Mediterranean region and widely cultivated for its fruit, eaten after curing, or oil from its seeds.

    |--Myxopyreae T00
    `--Schrebereae T00

Oleoideae incertae sedis:
  Olea K09
    |--O. europaea Linnaeus 1753 PL04
    |    |  i. s.: O. e. var. sylvestris R-RR-GM-S98
    |    |--O. e. ssp. europaea PT98
    |    `--O. e. ssp. oleaster PT98
    |--O. macintyrei Ettingshausen 1886 F71
    |--O. paniculata BR65
    `--O. welwitschii K09
  Hesperelaea T00
  Fraxinus BR65
    |--F. americana SK03
    |--F. anomala H93
    |--F. caroliniana M83
    |--F. chinensis HSS13
    |--F. dipetala H93
    |--F. excelsior BR65
    |--F. floribunda SN88
    |--F. lanuginose T03
    |--F. latifolia H93
    |--F. longicuspis S00
    |--F. micrantha SN88
    |--F. nigra B14
    |--F. ornus R-CT01
    |--F. oxycarpa Willdenow 1806 [incl. F. oxyphylla von Bieberstein 1808] BR65
    |--F. pennsylvanica MC03
    |--F. rostrata C55
    |--F. saxonica HL08
    |--F. sieboldiana LO98
    `--F. velutina [incl. F. velutina var. coriacea] H93

*Type species of generic name indicated


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

[B14] Bouchard, P. (ed.) 2014. The Book of Beetles: A lifesize guide to six hundred of nature's gems. Ivy Press: Lewes (United Kingdom).

[C55] 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.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1–167.

[HL08] Henniger, M., & R. Leder. 2008. Eozäne Makroflora der Querfurter Mulde. Mauritiana 20 (2): 229–251.

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

[HSS13] Hirschfeld, E., A. Swash & R. Still. 2013. The World's Rarest Birds. Princeton University Press: Princeton (New Jersey).

[K09] Kelbessa, E. 2009. Three new species of Acanthaceae from Ethiopia. Kew Bulletin 64 (1): 57–65.

[LO98] Lack, H. W., & H. Ohba. 1998. Die Xylothek des Chikusai Kato. Willdenowia 28: 263–276.

[MC03] Mostert, L., P. W. Crous, J. Z. Groenewald, W. Gams & R. C. Summerbell. 2003. Togninia (Calosphaeriales) is confirmed as teleomorph of Phaeoacremonium by means of morphology, sexual compatibility and DNA phylogeny. Mycologia 95 (4): 646–659.

[M83] Myers, R. L. 1983. Site susceptibility to invasion by the exotic tree Melaleuca quinquenervia in southern Florida. Journal of Applied Ecology 20: 645–658.

[PT98] Panitsa, M., & D. Tzanoudakis. 1998. Contribution to the study of the Greek flora: flora and vegetation of the E Aegean islands Agathonisi and Pharmakonisi. Willdenowia 28: 95–116.

[PL04] Pohl, G., & I. Lenski. 2004. Zur Verbreitung und Vergesellschaftung von Pennisetum orientale Rich. in Nordeuböa (Griechenland) (Poaceae, Paniceae). Senckenbergiana Biologica 83 (2): 209–223.

[R-CT01] Ragusa-di Chiara, S., & H. Tsolakis. 2001. Phytoseiid faunas of natural and agricultural ecosystems in Sicily. In: Halliday, R. B., D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff (eds) Acarology: Proceedings of the 10th International Congress pp. 522–529. CSIRO Publishing: Melbourne.

[R-RR-GM-S98] Ramil-Rego, P., M. Rodríguez-Guitián & C. Muñoz-Sobrino. 1998. Sclerophyllous vegetation dynamics in the north of the Iberian peninsula during the last 16,000 years. Global Ecology and Biogeography Letters 7: 335–351.

[S00] Siddiqi, M. R. 2000. Tylenchida: Parasites of plants and insects 2nd ed. CABI Publishing: Wallingford (UK).

[SK03] Snell, K. L., & H. W. Keller. 2003. Vertical distribution and assemblages of corticolous myxomycetes on five tree species in the Great Smoky Mountains National Park. Mycologia 95 (4): 565–576.

[SN88] Suzuki, M., & S. Noshiro. 1988. Wood structure of Himalayan plants. In: Ohba, H., & S. B. Malla (eds) The Himalayan Plants vol. 1. The University Museum, University of Tokyo, Bulletin 31: 341–379.

[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.

[T03] Tsurusaki, N. 2003. Phenology and biology of harvestmen in and near Sapporo, Hokkaido, Japan, with some taxonomical notes on Nelima suzukii n. sp. and allies (Arachnida: Opiliones). Acta Arachnologica 52: 5–24.

Last updated: 25 November 2018.


Dryandra nivea, from Wikimedia.

Belongs within: Proteaceae.

Dryandra has been recognised as a genus of shrubs and small trees found in south-western Western Australia. Recent revisions have suggested its synonymisation with the related genus Banksia, from which it was previously distinguished by having flat rather than conical inflorescences.

Characters (from A. S. George): Shrubs or small trees, many prostrate. Leaves serrate to pinnatifid or pinnatipartite, sometimes almost pinnatisect, sometimes bipinnatifid, sometimes entire, hairy becoming glabrous adaxially, closely tomentose or woolly between nerves abaxially. Inflorescence terminal, axillary or on short lateral branchlet, commonly sessile, capitate; receptacle concave, flat or convex; involucre usually prominent, of many imbricate flat or subulate bracts, persistent; flowering usually centripetal; flowers 15–250 per head; floral bracts usually enlarged after flowering. Perianth straight or curved; tepals separating almost to base at anthesis. Pistil straight or curved, with style often exserted to one side before anthesis; pollen presenter usually erect, narrower, as wide as or wider than style. Follicles few per head, woody but not massively so, commonly striate, in 2 species cartilaginous. Seed usually with a terminal wing, rarely elliptic with an annular wing and then without a separator but with 2 supplementary outer wings.

    |--D. arborea G04a
    |--D. armata OS04
    |--D. bipinnatifida GK00
    |--D. carduacea RL05
    |--D. erythrocephala G04b
    |--D. ferruginea G04b
    |    |--D. f. ssp. ferruginea G04b
    |    `--D. f. ssp. flavescens G04b
    |--D. floribunda S35
    |--D. fraseri OS04
    |--D. lindleyana JK08
    |    |--D. l. ssp. lindleyana OS04
    |    `--D. l. ssp. sylvestris OS04
    |--D. nivea S35
    |--D. nobilis OS04
    |--D. pallida G04b
    |--D. porrecta GK00
    |--D. praeformosa Ettingshausen 1886 F71
    |--D. praemorsa JK08
    |--D. purdieana G04b
    |--D. rufistylis OS04
    |--D. sessilis JK08
    |--D. squarrosa JK08
    `--D. viscida G04b

*Type species of generic name indicated


[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1-167.

[G04a] Gibson, N. 2004a. Flora and vegetation of the Eastern Goldfields Ranges: part 6. Mt Manning Range. Journal of the Royal Society of Western Australia 87 (2): 35-47.

[G04b] Gibson, N. 2004b. Flora and vegetation of the Eastern Goldfields Ranges: part 7. Middle and South Ironcap, Digger Rock and Hatter Hill. Journal of the Royal Society of Western Australia 87 (2): 49-62.

[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.

[JK08] Johnstone, R. E., & T. Kirkby. 2008. Distribution, status, social organisation, movements and conservation of Baudin's cockatoo (Calyptorhynchus baudinii) in south-west Western Australia. Records of the Western Australian Museum 25 (1): 107-118.

[OS04] Obbens, F. J., & L. W. Sage. 2004. Vegetation and flora of a diverse upland remnant of the Western Australian wheatbelt (Nature Reserve A21064). Journal of the Royal Society of Western Australia 87 (1): 19-28.

[RL05] Rafferty, C., & B. B. Lamont. 2005. Selective feeding by macropods on vegetation regenerating following fire. Journal of the Royal Society of Western Australia 88 (4): 155-165.

[S35] Solomon, M. E. 1935. On a new genus and two new species of Western Australian Aleyrodidae. Journal of the Royal Society of Western Australia 21: 75-91.


Rough horsetail Equisetum hyemale, copyright Liné1.

Belongs within: Monilophyta.

The Equisetopsida includes the modern horsetails, Equisetum, and their fossil relatives. Members of this group have whorled leaves, and most have a eustele stem structure with a hollow pith surrounded by vascular bundles (Doyle 1998).

<==Equisetopsida [Equisetophyta, Sphenophyta, Sphenophytae, Sphenopsida]
    |  i. s.: Equisetires keuperiana W70
    |         Schizoneura Z02
    |           |--S. australis Etheridge 1893 F71
    |           `--S. paradoxa W70
    |         Phyllotheca Z02
    |           |--P. australis F71
    |           |--P. robusta F71
    |           `--P. wonthaggiensis [=Equisetites wonthaggiensis] KDD98
    |         Archaeocalamites radiatus D98, F71 [=Calamites radiatus F71]
    |--Calamostachys A38 [Calamostachyales Z02]
    |--Sphenophyllum [Sphenophyllales] D98
    |    |--S. antiquum A38
    |    `--S. cuneifolium A38
    `--Equisetales D98
         |--Calamites suckowi D98, A38
         `--Equisetum Linnaeus 1753 CD07 (see below for synonymy)
              |  i. s.: E. endoi BO02
              |         E. filum BO02
              |         E. inglisii D06
              |         E. renaultii BO02
              |         E. robustum SM03
              |         E. × schaffneri D06
              |--E. subg. Equisetum D06
              |    |--E. × dubium [=E. arvense × E. telmateia] D06
              |    |--E. × litorale [E. arvense × E. fluviatile] D06
              |    |--E. × rothmaleri [E. arvense × E. palustre; incl. E. × rogesianum] D06
              |    |--E. × wallichianum Page 1974 (see below for synonymy) D06
              |    |--E. sect. Equisetum D06
              |    |    |--*E. fluviatile Linnaeus 1753 [incl. E. maximum (nom. inv.)] D06
              |    |    |--E. bogotense D06
              |    |    |--E. diffusum Don 1825 (see below for synonymy) D06
              |    |    |--E. × dycei Page 1981 [E. fluviatile × E. palustre] D06
              |    |    |--E. × font-queri [E. palustre × E. telmateia] D06
              |    |    |--E. palustre Linnaeus 1753 (see below for synonymy) D06
              |    |    `--E. telmateia Ehrh. 1783 D06, CD07
              |    `--E. sect. Heterophyadica Br. 1839 D06
              |         |--E. (sect. *Heterophyadica) arvense Linnaeus 1753 (see below for synonymy) D06
              |         |--E. × calderi [E. arvense × E. pratense] D06
              |         |--E. pratense D06
              |         |--E. × suecicum [E. arvense × E. pratense] D06
              |         `--E. sylvaticum D06
              `--E. subg. Hippochaete (Milde) Baker 1887 [=Hippochaete Milde 1865] D06
                   |--E. × ferrissii [E. hyemale var. affine × E. laevigatum] D06
                   |--E. × mexicanum [E. hyemale var. affine × E. myriochaetum] D06
                   |--E. × moorei [E. hyemale var. hyemale × E. ramosissimum ssp. ramosissimum] D06
                   |--E. × naegelianum [E. ramosissimum ssp. ramosissimum × E. variegatum] D06
                   |--E. sect. Hippochaete [=Hippochaete sect. Euhippochaete Farwell 1916] D06
                   |    |  i. s.: E. scirpoides D06
                   |    |         E. × trachyodon [=Hippochaete trachyodon; Equisetum hyemale × E. variegatum] D06
                   |    |         E. variegatum D06
                   |    `--E. subsect. Perennantia (Doell) Hauke 1962 (see below for synonymy) D06
                   |         `--E. (sect. *Hippochaete, subsect. *Perennantia) hyemale Linnaeus 1753 (see below for synonymy) D06
                   |              |--E. h. var. hyemale D06
                   |              `--E. h. var. affine D06
                   |--E. sect. Ambigua (Farwell) Hauke 1962 [=Hippochaete sect. Ambigua Farwell 1916] D06
                   |    |--E. (sect. *Ambigua) laevigatum Br. 1844 D06
                   |    |--E. myriochaetum D06
                   |    `--E. ramosissimum Desf. 1800 [=Hippochaete ramosissima] D06
                   |         |--E. r. ssp. ramosissimum [incl. E. ramosissimum var. altissimum, E. elongatum] D06
                   |         `--E. r. ssp. debile (Roxburgh) Hauke 1962 D06 (see below for synonymy)
                   `--E. sect. Incunabula Hauke 1962 [=Hippochaete subg. Incunabula (Hauke) Holub 1972] D06
                        `--E. (sect. *Incunabula) giganteum Linnaeus 1763 (see below for synonymy) D06

Equisetum Linnaeus 1753 CD07 [incl. Allostelites Börner 1912 D06; Equisetaceae, Gonopterides, Iulospermae, Peltasporae, Peltata, Peltiflorae, Peltigerae]

Equisetum subsect. Perennantia (Doell) Hauke 1962 [=E. II Sclerocaulon b. Perennantia Doell 1857] D06

Equisetum (sect. *Heterophyadica) arvense Linnaeus 1753 [incl. E. boreale, E. plantula Griffith 1849, E. saxicola] D06

Equisetum diffusum Don 1825 [incl. E. bicarinatum, E. diffusum var. caespitosum, E. leave, E. mekongense Page 1974, E. diffusum var. nudum, E. diffusum var. paucidentatum Page 1974, E. diffusum var. polystachyum Milde 1867, E. diffusum var. ramosum non E. ramosum Payot. 1860] D06

Equisetum (sect. *Incunabula) giganteum Linnaeus 1763 [incl. E. elongatum var. affine non E. hyemale var. affine, E. brasiliense, E. caracasanum, E. elongatum var. dolosum, E. martii, E. poeppigianum, E. elongatum var. scaberrimum] D06

Equisetum (sect. *Hippochaete, subsect. *Perennantia) hyemale Linnaeus 1753 [=Hippochaete hiemale, H. hyemalis] D06

Equisetum palustre Linnaeus 1753 [incl. E. palustre var. americana Victorin 1927, E. arenarium Opiz 1819, E. nodosum Hoppe 1794, E. pratense de Schlechtendal 1823-1824 (preoc.), E. ramosum Payot. 1860, E. palustre var. szechuanense Page 1974, E. tenellum Fries 1846, E. tuberosum Hectot ap. De Candolle 1815, E. veronense Pollini 1816] D06

Equisetum ramosissimum ssp. debile (Roxburgh) Hauke 1962 D06 [=E. debilis D06, E. debile Roxburgh ex Vaucher 1822 I88, Hippochaete debilis (Vaucher) Holub 1972 D06; incl. E. huegelii Milde 1861 D06, E. pallens Wallich 1828 D06, E. debile var. pashan Mahabale 1938 D06]

Equisetum × wallichianum Page 1974 [E. arvense × E. diffusum; incl. E. scoparium Wallich 1828 (n. n.)] D06

*Type species of generic name indicated


[A38] Alcock, F. J. 1938. Geology of Saint John region, New Brunswick. Geological Survey of Canada Memoir 216: 1-65.

[BO02] Barale, G., & M. Ouaja. 2002. La biodiversité végétale des gisements d’âge Jurassique supérieur-Crétacé inférieur de Merbah El Asfer (Sud-Tunisien). Cretaceous Research 23: 707-737.

[CD07] Cantino, P. D., J. A. Doyle, S. W. Graham, W. S. Judd, R. G. Olmstead, D. E. Soltis, P. S. Soltis & M. J. Donoghue. 2007. Towards a phylogenetic nomenclature of Tracheophyta. Taxon 56 (3): E1-E44.

[D06] Datta, A. 2006. The family Equisetaceae in India. Bulletin of the Botanical Survey of India 48 (1-4): 1-58.

[D98] Doyle, J. A. 1998. Phylogeny of vascular plants. Annual Review of Ecology and Systematics 29: 567-599.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1-167.

[I88] Iwatsuki, K. 1988. An enumeration of the pteridophytes of Nepal. In The Himalayan Plants vol. 1 (H. Ohba & S. B. Malla, eds) The University Museum, University of Tokyo, Bulletin 31: 231-339.

[KDD98] Krassilov, V. A., D. L. Dilcher & J. G. Douglas. 1998. New ephedroid plant from the Lower Cretaceous Koonwarra fossil bed, Victoria, Australia. Alcheringa 22 (2): 123-133.

[SM03] Saldarriaga, J. F., M. L. McEwan, N. M. Fast, F. J. R. Taylor & P. J. Keeling. 2003. Multiple protein phylogenies show that Oxyrrhis marina and Perkinsus marinus are early branches of the dinoflagellate lineage. International Journal of Systematic and Evolutionary Microbiology 53: 355-365.

[W70] Wills, L. J. 1970. The Triassic succession in the central Midlands in its regional setting. Quarterly Journal of the Geological Society of London 126: 225-283.

[Z02] Zherikhin, V. V. 2002. Ecological history of the terrestrial insects. In History of Insects (A. P. Rasnitsyn & D. L. J. Quicke, eds) pp. 331-388. Kluwer Academic Publishers: Dordrecht.


Astrammina rara, from micro*scope.

Belongs within: Harosa.
Contains: Morozovella, Komokiacea, Hemisphaerammininae, Hyperamminidae, Allogromiidae, Lagynidae, Lagenida, Fusulinida, Tubothalamea, Bathysiphon, Saccamminidae, Xenophyophorea, Globothalamea.

The Foraminifera are a diverse group of aquatic, mostly marine, amoeboid organisms with granular cytoplasm forming slender anastomosing pseudopodia. Most Foraminifera construct some form of resistant test, and as a result they have an extensive fossil record. There are, however, some basal forms such as Reticulomyxa filosa that lack a test. If present, the test may be composed of foreign particles (such as sand grains) glued together with a cement, or it may be secreted directly by the foram itself. Classifications of forams have been heavily influenced by whether the test is agglutinated or secreted, and whether it is divided into multiple, discrete chambers, but it has become clear as our understanding of foram evolution has improved that both these features evolved on multiple occasions (Pawlowski et al. 2013). Multi-chambered tests are found within the Lagenida, Fusulinida, Tubothalamea and Globothalamea. Calcareous tests have also arisen independently within the Lagenida, Tubothalamea and Globothalamea.

Basal, unilocular forams may have a membranous or pseudochitinous test, or one composed of agglutinated sand grains and/or other foreign particles, and have been divided respectively between the Lagynacea and Ammodiscacea on the basis of this feature (Loeblich & Tappan 1964a). Once again, however, the different conditions are not phylogenetically distinct and the classification of many basal forams remains uncertain. Various basal forams may be free-living or attached to the substrate, and the test may be dominated by a single central chamber or elongate and/or branching.

Characters (from Loeblich & Tappan 1964a, as Foraminiferida + Granuloreticulosia): Cytoplasm minutely granular; pseudopodia delicate and reticulate. Protoplasmic body usually protected by test composed of one or more interconnected chambers; wall may be imperforate, finely or coarsely perforate, and may be chitinous, or composed of agglutinated particles or secreted material (rarely silica or aragonite, more commonly calcite), may have canal system of varying complexity; commonly with one or more large openings or apertures in addition to smaller wall perforations which may be present; pseudopodia protruding from apertures and perforations. Reproduction characterised by alternation of sexual and asexual generations, though one generation may be secondarily repressed; gametes commonly flagellate with two or three flagella, more rarely amoeboid.

Foraminifera (see below for synonymy)
    |--Astrammina Rhumbler in Wiesner 1931 PH03, LT64a [incl. Armorella Heron-Allen & Earland 1932 LT64a]
    |    |--*A. rara Rhumbler in Wiesner 1931 LT64a
    |    |--A. limicola G96
    |    |--A. sphaerica (Heron-Allen & Earland 1932) [=*Armorella sphaerica] LT64a
    |    `--A. triangularis PH03
    `--+--Allogromiidae PHT13
       `--+--+--Cribrothalammina alba PHT13
          |  `--Reticulomyxa Nauss 1949 PHT13, AS12 [Athalamea]
          |       `--R. filosa PH03
          `--+--+--Lagynidae PHT13
             |  `--+--Vellaria PHT13
             |     `--Psammophaga simplora PHT13, PH03
             `--+--+--+--Lagenida PHT13
                |  |  `--Fusulinida GAR03
                |  `--+--Tubothalamea PHT13
                |     `--+--Bathysiphon PHT13
                |        `--Micrometula Nyholm 1952 PHT13, LT64a
                |             `--*M. hyalostriata Nyholm 1952 LT64a
                `--+--+--Cylindrogullmia alba PH03
                   |  `--+--+--Toxisarcon synsuicidica PH03
                   |     |  `--+--+--Gloiogullmia PH03
                   |     |     |  `--Saccamminidae PH03
                   |     |     `--Rhabdammina Sars in Carpenter 1869 PH03, LT64a (see below for synonymy)
                   |     |          |--*R. abyssorum Sars in Carpenter 1869 [=*Arrhabdamum abyssorum] LT64a
                   |     |          |--R. irregularis C40
                   |     |          `--R. linearis Brady 1879 (see below for synonymy) LT64a
                   |     `--+--+--Xenophyophorea PHT13
                   |        |  `--Shinkaiya lindsayi PHT13
                   |        `--Rhizamminidae [Arrhizamnia, Rhizammininae, Testulosiphoninae] PH03
                   |             |--Marsipella Norman 1878 [=Armarsipellum Rhumbler 1913] LT64a
                   |             |    `--*M. elongata Norman 1878 [=*Armarsipellum elongatum] LT64a
                   |             `--Rhizammina Brady 1879 PHT13, LT64a (see below for synonymy)
                   |                  |--*R. algaeformis Brady 1879 [=*Arrhizammum algaeforme] LT64a
                   |                  |--R. globigerinifera Hofker 1930 [=*Testulorhiza globigerinifera] LT64a
                   |                  `--R. indivisa Brady 1884 [=*Testulosiphon indivisum] LT64a
                   `--Globothalamea PHT13

Foraminifera incertae sedis:
  Neoendothyra PE02
  Spheroidinellopsis seminulina KS02
  Lingulogavelinella Malapris 1965 BL79
    `--L. albiensis RC02
  Archaealveolina reicheli HS02
  Haynesina germanica LA03
  Radicula Christiansen 1958 LT64a
    `--*R. limosa Christiansen 1958 LT64a
  Globospirillina Antonova 1964 BL79
    `--G. clara BL79
  Mormosina kijastschensis BL79
  Arborammina G96
  Ellesmerella Mamet & Roux in Mamet et al. 1987 VH01
  Orobias Eichwald 1860 (n. d.) LT64b
    `--*O. aequalis Eichwald 1860 LT64b
  Consobrinella consobrina CM01
  Nodosarchaediscus CM01
    |--N. demaneti CM01
    `--N. saleei CM01
  Omphalotis minima CM01
  Schizocladus [Schizocladea] C-S03
  Planocassidulina H03
    |--P. norcrossi (Cushman 1933) (see below for synonymy) H03
    |--P. sublimbata (Asano & Nakamura 1937) [=Cassidulina sublimbata] H03
    `--P. yabei (Asano & Nakamura 1937) [=Cassidulina yabei] H03
  Loxoendothyra parakosvensis E03
  Palaeospiroplectammina parva E03
  Radiosphaera basilica E03
  Archaeoglobigerina FM03
    |--A. australis Huber 1990 FM03
    `--A. mateola Huber 1990 FM03
  *Paraglobivalvulina mira K04
  Dagmarita chanakchiensis K04
  Morozovella MO81
  Vanhoeffenella Rhumbler 1905 [=Arvanhoeffenum Rhumbler 1913] LT64a
    `--*V. gaussi Rhumbler 1905 [=*Arvanhoeffenum gaussi] LT64a
  ‘Paralagena’ Sabirov 1986 non Dall 1904 MSV00
  Septatounayella rauserae ZP86
  Eodiscus guangdongensis ZP86
  Shouguania annectena ZP86
  Rosita fornicata G-B87
  Praeorbulina curva H90
  Neolenticulina peregrina JW99
  Rutherfordoides tenuis JW99
  ‘Nautilus’ umbilicatus Linnaeus 1758 S64
  Cinibides thiara B49
  Dicarinella M08
    |--D. asymmetrica (see below for synonymy) M08
    |--D. concavata M08
    `--D. primitiva M08
  Radotruncana calcarata M08
  Contusotruncana fornicata M08
  Spiroplectinella semicomplanata M08
  Marginula breoni F71
  Schizamminidae LT64a
    |--Jullienella Schlumberger 1890 LT64a
    |    `--*J. foetida Schlumberger 1890 LT64a
    `--Schizammina Heron-Allen & Earland 1929 (see below for synonymy) LT64a
         |--*S. labyrinthica Heron-Allen & Earland 1929
         `--S. wedmoriensis (Chapman 1895) (see below for synonymy) LT64a
  Astrorhiza Sandahl 1858 (see below for synonymy) LT64a
    |--*A. limnicola Sandahl 1858 [=*Arastrorhizum limnicola] LT64a
    |--*Astrodiscus’ arenaceus Schulze 1875 LT64a
    |--A. arenaria C40
    |--*Haeckelina’ gigantea Bessels 1875 LT64a
    |--A. granulosa C40
    `--‘Ammodiscus’ lindahli Carpenter & Jeffreys 1870 (see below for synonymy) LT64a
  Komokiacea TH77
  Eoforschia moelleri R79
  Orbulinoides beckmanni P79
  Adherentina Spandel 1909 (n. d.) LT64b
    `--*A. rhenana Spandel 1909 LT64b
  Aeolides de Montfort 1808 (n. d.) LT64b
    `--*A. squammatus de Montfort 1808 LT64b
  Arthrocena Modeer 1791 (n. d.) LT64b
  Aspidodexia Ehrenberg 1872 (n. d.) LT64b
    `--*A. lineolata Ehrenberg 1872 LT64b
  ‘Auriculina’ Costa 1856 nec Grateloup 1838 nec Gray 1847 (n. d.) LT64b
    `--*A. crenata Costa 1856 LT64b
  ‘Buliminopsis’ Rzehak 1895 non Heude 1890 (n. d.) LT64b
    `--*B. conulus Rzehak 1895 LT64b
  Calatharia Zalessky 1926 LT64b
    `--*C. perforata Zalessky 1926 LT64b
  Cameroconus Meunier 1888 (n. d.) LT64b
    `--*C. marmoris Meunier 1888 LT64b
  ‘Canopus’ de Montfort 1808 non Fabricius 1803 (n. d.) LT64b
    `--*C. fabeolatus de Montfort 1808 LT64b
  ‘Cantharus’ de Montfort 1808 non Röding 1798 (n. d.) LT64b
    `--*C. calceolatus de Montfort 1808 LT64b
  Canthropes de Montfort 1808 (n. d.) [=Cantharipes Agassiz 1846, Canthropus Pallas in Oken 1815] LT64b
    `--*C. galet de Montfort 1808 LT64b
  Cepinula Schafhäutl 1851 (n. d.) LT64b
  Cerataria Zalessky 1926 (n. d.) LT64b
    `--*C. pulchella Zalessky 1926 LT64b
  Chelibs de Montfort 1808 (n. d.) [=Celibs Sherborn 1893] LT64b
    `--*C. gradatus de Montfort 1808 LT64b
  Cidarollus de Montfort 1808 (n. d.) LT64b
    `--*C. plicatus de Montfort 1808 LT64b
  Cimelidium Ehrenberg 1858 (n. d.) LT64b
    `--*C. homeri (Ehrenberg 1858) [=Guttulina homeri] LT64b
  Clypeocyclina Silvestri 1908 (n. d.) LT64b
  Colpopleura Ehrenberg 1844 (n. d.) LT64b
    `--*C. ocellata (Ehrenberg 1838) [=Rotalia ocellata] LT64b
  Cortalus de Montfort 1808 (n. d.) LT64b
    `--*C. pagodus de Montfort 1808 LT64b
  Craterularia Rhumbler 1911 (n. d.) LT64b
  Crustula Allix in Lecointre & Allix 1913 (n. d.) LT64b
    `--*C. complanata Allix in Lecointre & Allix 1913 LT64b
  Cyclopavonina Silvestri 1937 (n. d.) LT64b
    `--*C. cyclica Silvestri 1937 LT64b
  Cylindria de Gregorio 1930 (n. d.) LT64b
    `--*C. minuta de Gregorio 1930 LT64b
  Dexiopora Ehrenberg 1861 (n. d.) LT64b
    `--*D. megapora Ehrenberg 1861 LT64b
  Dorbignyaea Deshayes 1830 (n. d.) LT64b
  ‘Dujardinia’ Gray 1858 non Quatrefages 1844 (n. d.) LT64b
    `--*D. mediterranea Gray 1858 LT64b
  Elliptina Harting 1852 (n. d.) LT64b
    |--E. inflata LT64b
    `--E. truncata LT64b
  Epistominites Zalessky 1926 (n. d.) LT64b
    `--*E. formosulus Zalessky 1926 LT64b
  Glandiolus de Montfort 1808 (n. d.) LT64b
    `--*G. gradatus de Montfort 1808 LT64b
  Hedbergina Brönnimann & brown 1956 (n. d.) LT64b
    `--*H. seminolensis (Harlton 1927) [=Globigerina seminolensis] LT64b
  Hemistegina Kaufmann 1867 (n. d.) LT64b
    `--*H. rotula Kaufmann 1867 LT64b
  Hemisterea Ehrenberg 1872 (n. d.) LT64b
    `--*H. nautilus Ehrenberg 1872 LT64b
  Hemisticta Ehrenberg 1872 (n. d.) LT64b
    `--*H. amplificata Ehrenberg 1872 LT64b
  ‘Heterostomum’ Ehrenberg 1854 non Diesing 1850 (n. d.) LT64b
    `--*H. cyclostomum Ehrenberg 1854 LT64b
  Lagenopsis de Gregorio 1930 (n. d.) LT64b
    `--*L. maliarda de Gregorio 1930 LT64b
  Lekithiammina de Folin 1887 (n. d.) LT64b
    `--*L. aculeata de Folin 1887 LT64b
  ‘Lepista’ Zalessky 1926 non Wallengren 1863 (n. d.) LT64b
    `--*L. ornata Zalessky 1926 LT64b
  ‘Lobularia’ Costa 1839 non Lamarck 1816 (n. d.) LT64b
    `--*L. vesiculosa Costa 1839 LT64b
  Lyrina Zborzewski 1834 LT64b
    `--*L. fischeri Zborzewski 1834 LT64b
  ‘Mesopora’ Ehrenberg 1854 non Wesmael 1852 LT64b
    `--*M. chloris Ehrenberg 1854 LT64b
  Metarotaliella Grell 1962 (n. d.) LT64b
    `--*M. parva Grell 1962 LT64b
  Mirfa de Gregorio 1890 LT64b
    `--*M. subtetraedra de Gregorio 1890 LT64b
  Mirga de Gregorio 1930 (n. d.) LT64b
    `--*M. permiana (de Gregorio 1930) [=Orbulina (*Mirga) permiana] LT64b
  Molnaria Zalessky 1926 (n. d.) LT64b
    `--*M. spinulata Zalessky 1926 LT64b
  ‘Monocystis’ Ehrenberg 1854 non Stein 1848 (n. d.) LT64b
    `--*M. arcella (Ehrenberg 1854) [=Miliola (*Monocystis) arcella] Lt64b
  Nummulitella Dorren 1948 (n. d.) LT64b
    `--*N. polystylata Dorren 1948 LT64b
  Oncobotrys Ehrenberg 1856 (n. d.) LT64b
    `--*O. buccinum Ehrenberg 1856 LT64b
  Ozawaina Lee 1927 (n. d.) LT64b
    `--*O. antiquior (Rouillier & Vosinsky 1849) LT64b (see below for synonymy)
  Orthocerina d’Orbigny in de la Sagra 1839 (n. d.) LT64b
    |--*O. quadrilatera (d’Orbigny in de la Sagra 1839) [=Nodosaria (*Orthocerina) quadrilatera] LT64b
    |--O. clavulus [=Nodosaria (Orthocerina) clavulus] LT64b
    `--O. murchisoni LT64b
  Otostomum Ehrenberg 1872 (n. d.) LT64b
    `--*O. strophoconus Ehrenberg 1872 LT64b
  ‘Pectinaria’ Zalessky 1926 non Lamarck 1818 (n. d.) LT64b
    `--*P. costata Zalessky 1926 LT64b
  Physomphalus Ehrenberg 1856 (n. d.) LT64b
    `--*P. porosus Ehrenberg 1856 LT64b
  Pleurostomina Costa 1862 (n. d.) LT64b
    `--*P. bimucronata Costa 1862 LT64b
  Pleurotrema Ehrenberg 1840 (n. d.) LT64b
    `--*P. calcarina Ehrenberg 1840 LT64b
  Pseudastrorhizula Wetzel 1940 (n. d.) LT64b
    `--*P. eisenacki Wetzel 1940 LT64b
  Pteroptyx Ehrenberg 1873 (n. d.) LT64b
    `--*P. vespertilio Ehrenberg 1873 LT64b
  ‘Renulina’ Blake 1876 nec Lamarck 1804 nec de Blainville 1825 (n. d.) LT64b
    `--*R. sorbyana Blake 1876 LT64b
  Rhabdella d’Archiac & Haime 1853 (n. d.) LT64b
    `--*R. malcolmi d’Archiac & Haime 1853 LT64b
  Rhynchoplecta Ehrenberg 1854 (n. d.) LT64b
    `--*R. punctata Ehrenberg 1854 LT64b
  Rotalites Lamarck 1801 (n. d.) LT64b
    `--*R. tuberculosa Lamarck 1801 LT64b
  Semseya Franzenau 1893 (n. d.) LT64b
    `--*S. lamellata Franzenau 1863 LT64b
  Siderospira Ehrenberg 1845 (n. d.) LT64b
    `--*S. indica (Ehrenberg 1845) [=Siderolina indica] LT64b
  Spiroplectina Schubert 1902 (n. d.) LT64b
  Spiropleurites Ehrenberg 1854 (n. d.) LT64b
  Synspira Ehrenberg 1854 (n. d.) LT64b
    `--*S. triquetra Ehrenberg 1854 LT64b
  Trioxeia de Folin 1888 (n. d.) LT64b
    `--*T. edwardsi de Folin 1888 LT64b
  Upsonella Moore 1959 (n. d.) LT64b
    `--*U. typus Moore 1959 LT64b
  Volutaria Zalessky 1926 (n. d.) LT64b
    `--*V. potoniei Zalessky 1926 LT64b
  Volvotextularia Termier & Termier 1950 (n. d.) LT64b
    `--*V. polymorpha Termier & Termier 1950 LT64b
  Parredicta porifera LJ98
  0--Hemisphaerammininae PH03
  `--Notodendrodes antarctikos PH03
  Hyperamminidae C40

Inorganic: Astrorhiza cretacea Franke 1928 H75

‘Ammodiscus’ lindahli Carpenter & Jeffreys 1870 [incl. *Arenistella agglutinans Fischer in de Folin & Périer 1875] LT64a

Astrorhiza Sandahl 1858 [=Arastrorhizum Rhumbler 1913; incl. Arenistella Fischer in de Folin & Périer 1875, Astrodiscus Schulze 1875 non Ludwig 1866, Haeckelina Bessels 1875] LT64a

Dicarinella asymmetrica [incl. Globotruncana (Globotruncana) ventricosa carinata, G. fundiconulosa, G. lobata] M08

Foraminifera [Allogromidiaceae, Allogromiida, Allogromiidea, Allogromiidia, Allogromiina, Allogromioidea, Ammoasconidae, Ammodinetta, Ammodiscacea, Ammodiscida, Arastrorhiznia, Archimonothalamia, Arenacea, Arenacidae, Arforaminifera, Arreticularia, Arrhabdamidia, Arrhabdamnia, Ascoforaminifera, Asiphoidea, Astrorhizicae, Astrorhizida, Astrorhizidaceae, Astrorhizidae, Astrorhizidea, Astrorhizina, Astrorhizinae, Ataxophragmiida, Botellinidae, Dendrophryida, Dendrophryidae, Foraminiferae, Foraminiferiae, Foraminiferida, Haplophragmiacea, Lagynacea, Lagynidea, Lituolida, Lituolidaceae, Lituolidea, Metammida, Microcometides, Monosomatia, Monostegua, Monothalamia, Perforata, Polysomatia, Polythalamea, Polythalamiis, Protammida, Psammatostichostegia, Reticulariida, Rhabdamminae, Rhabdamminidae, Rhabdamminina, Rhabdammininae, Saccorhizidae, Serpuleidae, Siphonoforaminifera, Textulariacea, Textulariaceae, Textularida, Textularidae, Textularidea, Textularidia, Textulariidea, Textulariina, Thalamia, Thalamophora, Tubulata]

*Ozawaina antiquior (Rouillier & Vosinsky 1849) LT64b [=Nummulina antiquior C40, Fusulinella (*Ozawaina) antiquior LT64b, Orobias antiquior C40]

Planocassidulina norcrossi (Cushman 1933) [=Cassidulina norcrossi; incl. C. kasiwazakiensis Husezima & Maruhasi 1944, C. nakamurai Uchio 1951] H03

Rhabdammina Sars in Carpenter 1869 PH03, LT64a [=Arrhabdammum Rhumbler 1913 LT64a; incl. Oculosiphon Avnimelech 1952 LT64a, Rhabdammina Eimer & Fickert 1899 non Sars in Carpenter 1869 LT64a]

Rhabdammina linearis Brady 1879 [=*Oculosiphon linearis, *Rhabdammina (sensu Eimer & Fickert) linearis] LT64a

Rhizammina Brady 1879 PHT13, LT64a [=Arrhizammum Rhumbler 1913 LT64as; incl. Testulorhiza Avnimelech 1952 LT64a, Testulosiphon Avnimelech 1952 LT64a]

*Schackoina cenomana (Schacko 1897) [=Siderolina cenomana, Hantkenina (*Schackoina) cenomana] LT64b

Schizammina Heron-Allen & Earland 1929 [incl. Arpsammosiphoum Rhumbler 1913, Psammosiphon Rhumbler 1911 non Vine 1882] LT64a

Schizammina wedmoriensis (Chapman 1895) [=Nodosinella wedmoriensis, *Arpsammosiphoum wedmoriense, *Psammosiphon wedmoriense] LT64a

*Type species of generic name indicated


[AS12] Adl, S. M., A. G. B. Simpson, C. E. Lane, J. Lukeš, D. Bass, S. S. Bowser, M. W. Brown, F. Burki, M. Dunthorn, V. Hampl, A. Heiss, M. Hoppenrath, E. Lara, E. Le Gall, D. H. Lynn, H. McManus, E. A. D. Mitchell, S. E. Mozley-Stanridge, L. W. Parfrey, J. Pawlowski, S. Rueckert, L. Shadwick, C. L. Schoch, A. Smirnov & F. W. Spiegel. 2012. The revised classification of eukaryotes. Journal of Eukaryotic Microbiology 59 (5): 429–493.

[BL79] Basov, V. A., B. G. Lopatin, I. S. Gramberg, A. I. Danjushevskaya, V. Ya. Kaban’kov, V. M. Lazurkin & D. K. Patrunov. 1979. Lower Cretaceous lithostratigraphy near Galicia Bank. Initial Reports of the Deep Sea Drilling Project 47: 683–717.

[B49] Battey, M. H. 1949. The geology of the Tuakau-Mercer area, Auckland. Transactions and Proceedings of the Royal Society of New Zealand 77 (3): 429–455.

[C-S03] Cavalier-Smith, T. 2003. Protist phylogeny and the high-level classification of Protozoa. European Journal of Protistology 39 (4): 338–348.

[CM01] Cózar, P., & B. L. Mamet. 2001. Planohowchinia (Lasiodiscidae, Foraminiferida), a new Late Viséan genus, southwestern Spain. Journal of Foraminiferal Research 31 (3): 228–232.

[C40] Cushman, J. A. 1940. Foraminifera: Their classification and economic use 3rd ed. Harvard University Press: Cambridge (Massachusetts).

[E03] Edgell, H. S. 2003. Upper Devonian Charophyta of Western Australia. Micropaleontology 49 (4): 359–374.

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[G96] Gooday, A. J. 1996. Xenophyophores (Protista), including two new species, from two abyssal sites in the northeast Atlantic Ocean. Journal of Foraminiferal Research 26 (3): 193–208.

[GAR03] Groves, J. R., D. Altiner & R. Rettori. 2003. Origin and early evolutionary radiation of the order Lagenida (Foraminifera). Journal of Paleontology 77 (5): 831–843.

[H03] Hanagata, S. 2003. Miocene–Pliocene Foraminifera from the Niigata oil-fields region, northeastern Japan. Micropaleontology 49 (4): 293–340.

[H75] Häntzschel, W. 1975. Treatise on Invertebrate Paleontology pt W. Miscellanea Suppl. 1. Trace Fossils and Problematica 2nd ed. The Geological Society of America: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[H90] Hayward, B. W. 1990. Use of foraminiferal data in analysis of Taranaki Basin, New Zealand. Journal of Foraminiferal Research 20: 71–83.

[HS02] Höfling, R., & R. W. Scott. 2002. Early and Mid-Cretaceous buildups. SEPM Special Publication 72: 521–548.

[JW99] Jian, Z.-M., L.-J. Wang, M. Kienast, M. Sarnthein, W. Kuhnt, H.-L. Lin & P.-X. Wang. 1999. Benthic foraminiferal paleoceanography of the South China Sea over the last 40,000 years. Marine Geology 156: 159–186.

[KS02] Kanjilal, S., & M. S. Srinivasan. 2002. New bivalves from the Miocene of Little Andaman Island, Bay of Bengal. Journal of the Geological Society of India 60: 527–536.

[K04] Kobayashi, F. 2004. Late Permian foraminifers from the Limestone Block in the Southern Chichibu Terrane of west Shikoku, SW Japan. Journal of Paleontology 78 (1): 62–70.

[LJ98] Li, Q., N. P. James, B. McGowran, Y. Bone & J. Cann. 1998. Synergetic influence of water masses and Kangaroo Island barrier on foraminiferal distribution, Lincoln and Lacepede shelves, South Australia: a synthesis. Alcheringa 22 (2): 153–176.

[LT64a] Loeblich, A. R., Jr & H. Tappan. 1964a. Sarcodina: chiefly “thecamoebians” and Foraminiferida. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt C. Protista 2 vol. 1. The Geological Society of America and The University of Kansas Press.

[LT64b] Loeblich, A. R., Jr & H. Tappan. 1964b. Sarcodina: chiefly “thecamoebians” and Foraminiferida. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt C. Protista 2 vol. 2. The Geological Society of America and The University of Kansas Press.

[LA03] Longet, D., J. M. Archibald, P. J. Keeling & J. Pawlowski. 2003. Foraminifera and Cercozoa share a common origin according to RNA polymerase II phylogenies. International Journal of Systematic and Evolutionary Microbiology 53: 1735–1739.

[MO81] Mancini, E. A., & G. E. Oliver. 1981. Planktic foraminifers from the Tuscahoma Sand (upper Paleocene) of southwest Alabama. Micropaleontology 27 (2): 204–225.

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Section of Pseudovermiporella sodalica, from Loeblich & Tappan (1964).

Belongs within: Miliolida.
Contains: Spiroloculina.

The Nubeculariidae are a group of often spreading or branched calcareous foraminifera known from the Middle Carboniferous to the present. Members of the subfamilies Nubeculariinae and some Nodobaculariinae have the test attached to their substrate; other members of the family have the test free (Loeblich & Tappan 1964).

Characters (from Loeblich & Tappan 1964): Test free or attached, planispiral or irregularly coiled, at least in early stages, later spreading or branched; aperture simple, rounded or slitlike or more rarely cribrate.

<==Nubeculariidae (see below for synonymy)
    |--Discospirina Munier-Chalmas 1902 (see below for synonymy) LT64
    |    `--D. italica (Costa 1856) (see below for synonymy) LT64
    |--Spiroloculininae LT64
    |    |--Spiroloculina LT64
    |    |--Cribrolinoides Cushman & LeRoy 1939 LT64
    |    |    `--*C. curta (Cushman 1917) [=Quinqueloculina disparilis var. curta] LT64
    |    `--Planispirinoides Parr 1950 LT64
    |         `--*P. bucculenta (Brady 1884) [=Miliolina bucculenta] LT64
    |--Nubeculariinae [Nubeculinellinae] LT64
    |    |  i. s.: Cornuspiramia Cushman 1928 [incl. Rhizonubecula Le Calvez 1935] LT64
    |    |           |--*C. antillarum (Cushman 1922) [=Nubecularia antillarum] LT64
    |    |           `--C. adherens (Le Calvez 1935) [=*Rhizonubecula adherens] LT64
    |    |         Nubeculopsis Collins 1958 LT64
    |    |           `--*N. queenslandica Collins 1958 LT64
    |    |         Webbina d’Orbigny in de la Sagra 1839 [=Arwebbum Rhumbler 1913, Webbum Rhumbler 1913] LT64
    |    |           `--*W. rugosa d’Orbigny in Barker-Webb & Berthelot 1839 [=*Arwebbum rugosum, *Webbum rugosum] LT64
    |    |--Nubeculinella Cushman 1930 C40, LT64
    |    |    `--*N. bigoti Cushman 1930 LT64
    |    |--Nubecularia Defrance 1825 C40
    |    |    |--*N. lucifuga Defrance 1825 LT64
    |    |    `--N. nitida Chapman 1909 F71
    |    `--+--Sinzowella Cushman 1933 C40
    |       |    `--*S. deformis (Karrer & Sinzow 1877) [=Nubecularia novorossica var. deformis] LT64
    |       `--+--Calcituba von Roboz 1884 C40, LT64
    |          |    `--*C. polymorpha von Roboz 1884 LT64
    |          `--Squamulina Schultze 1854 C40 (see below for synonymy)
    |               `--*S. laevis Schultze 1854 LT64
    |--Nodobaculariinae [Nodophthalmidiinae] LT64
    |    |--Nubeculina Cushman 1924 LT64
    |    |    `--*N. divaricata (Brady 1879) LT64 [=Sagrina divaricata LT64, Nubecularia divaricata C40]
    |    |--Orthella Bykova in Kiparisova et al. 1956 LT64
    |    |    `--*O. paalzowi Bykova in Kiparisova et al. 1956 LT64
    |    |--Meandroloculina Bogdanovich 1935 LT64
    |    |    |--*M. bogatschovi Bogdanovich 1935 LT64
    |    |    `--M. litoralis LT64
    |    |--Nodobaculariella Cushman & Hanzawa 1937 LT64
    |    |    |--*N. japonica Cushman & Hanzawa 1937 LT64
    |    |    `--N. atlantica C40
    |    |--Nodobacularia Rhumbler 1895 [incl. Gymnesina Colom 1959, Pseudonubeculina Bartenstein & Brand 1949] LT64
    |    |    |--*N. tibia (Jones & Parker 1860) [=Nubecularia tibia] LT64
    |    |    |--N. glomerosa (Colom 1959) [=*Gymnesina glomerosa] LT64
    |    |    `--‘Nubecularia’ nodulosa Chapman 1891 (see below for synonymy) LT64
    |    |--Vertebralina d’Orbigny 1826 LT64
    |    |    |--*V. striata d’Orbigny 1826 LT64
    |    |    |--V. catena H04
    |    |    |--V. furcata H04
    |    |    |--V. insignis H04
    |    |    `--V. mucronata H04
    |    `--Nodophthalmidium Macfadyen 1939 (see below for synonymy) LT64
    |         |--*N. compressum (Rhumbler 1906) [=Nodobacularia compressa] LT64
    |         |--N. costatum (Bogdanovich 1952) [=*Sarmatiella costata] LT64
    |         |--N. milletti C40
    |         |--N. obscurum (Bogdanovich 1960) [=*Foraminella obscura] LT64
    |         `--N. primum LT64
    `--Ophthalmidiinae [Holostreptae, Opthalmidiinae] LT64
         |--Ophthalmina Rhumbler 1936 LT64
         |    `--*O. kilianensis Rhumbler 1936 LT64
         |--Eosigmoilina Ganelina in Kiparisova et al. 1956 LT64
         |    `--*E. explicata Ganelina in Kiparisova et al. 1956 LT64
         |--Wiesnerella Cushman 1933 LT64
         |    `--*W. auriculata (Egger 1893) [=Planispirina auriculata] LT64
         |--Cornuloculina Burbach 1886 [=Hauerinella Schubert 1921] LT64
         |    `--*C. inconstans (Brady 1879) [=Hauerina inconstans, *Hauerinella inconstans] LT64
         |--Galeanella Kristan 1958 (see below for synonymy) LT64
         |    `--*G. tollmanni (Kristan 1957) [=*Galea tollmanni] LT64
         |--Glomulina Rhumbler 1936 LT64
         |    `--*G. fistulescens Rhumbler 1936 LT64
         |--Zoyaella Loeblich & Tappan 1962 [=Ceratina Goës 1894 nec Latreille 1802 nec Menge 1868] LT64
         |    `--*Z. trochamminoides (Goës 1894) [=*Ceratina trochamminoides, Fischerina trochamminoides] LT62
         |--Ophthalmidium Kübler & Zwingli 1870 (see below for synonymy) LT64
         |    |--*O. liasicum (Kübler & Zwingli 1866) [=*Oculina liasica] LT64
         |    |--O. acutimargo (Brady 1884) [=Spiroloculina acutimargo, *Spirophthalmidium acutimargo] LT64
         |    |--O. inconstans C40
         |    `--O. porosum Zwingli & Kübler 1870 C40
         |--Edentostomina Collins 1958 LT64
         |    |--*E. cultrata (Brady 1879) [=Miliolina cultrata] LT64
         |    |--E. durrandii [=Miliolina durrandii] LT64
         |    |--E. milletti [=Biloculina milletti] LT64
         |    `--E. rupertiana [=Miliolina rupertiana] LT64
         `--Pseudovermiporella Elliott 1958 LT64
              |--*P. sodalica Elliott 1958 VH01
              |--P. elliotti Erk & Bilgütay 1960 [=Vermiporella elliotti] VH01
              |--P. longipora (Praturlon 1963) VH01
              |--‘Anthracoporella’ mercurii Elliott 1968 VH01
              `--‘Vermiporella’ sumatrana Pia 1937 [incl. V. nipponica Endo in Endo & Kanuma 1954] VH01

Discospirina Munier-Chalmas 1902 [=Cyclophthalmidium Lister in Lankester 1903, Discospirinia (l. c.), Krumbachina Wiesner 1920; Discospiriniinae, Discospirininae, Krumbachininae] LT64

Discospirina italica (Costa 1856) [=Pavonina italica; incl. Orbitolites tenuissimus Carpenter in Carpenter & Jeffreys 1870, *Cyclophthalmidium tenuissimum, *Discospirina tenuissima, *Krumbachina tenuissima] LT64

Galeanella Kristan 1958 [=Galea Kristan 1957 nec Meuschen 1787 nec Smith 1817 nec Meyen 1833 nec Moerch 1852] LT64

Nodophthalmidium Macfadyen 1939 [incl. Foraminella Bogdanovich 1960 nec Sowerby 1835 nec Levinsen 1909; Sarmatiella Bogdanovich 1952] LT64

‘Nubecularia’ nodulosa Chapman 1891 [=*Pseudonubeculina nodulosa; incl. Lagenammina pyriformis, Nodophthalmidium pyriforme] LT64

Nubeculariidae [Calcitubida, Inaequilateralidae, Inequilateralidae, Nubecularida, Nubecularidae, Nubecularina, Nubecularinae, Ophtalmidiidae, Ophthalmididae, Ophthalmidiidae, Ovulinetta, Vertebralinidae]

Ophthalmidium Kübler & Zwingli 1870 [=Oculina Kübler & Zwingli 1866 non Lamarck 1816; incl. Spiropthalmidium Cushman 1927, Spirophthalmidium] LT64

Squamulina Schultze 1854 C40 [=Arsquamulum Rhumbler 1913 LT64; Squamulinida, Squamulinidae, Squamulinidea]

*Type species of generic name indicated


[C40] Cushman, J. A. 1940. Foraminifera: Their classification and economic use 3rd ed. Harvard University Press: Cambridge (Massachusetts).

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1–167.

[H04] Haeckel, E. 1899–1904. Kunstformen der Natur. Bibliographisches Institut: Leipzig und Wien.

[LT62] Loeblich, A. R., Jr & H. Tappan. 1962. Six new generic names in the Mycetozoida (Trichiidae) and Foraminiferida (Fischerinidae, Buliminidae, Caucasinidae, and Pleurostomellidae), and a redescription of Loxostomum (Loxostomidae, new family). Proceedings of the Biological Society of Washington 75: 107–114.

[LT64] Loeblich, A. R., Jr & H. Tappan. 1964. Sarcodina: chiefly “thecamoebians” and Foraminiferida. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt C. Protista 2 vol. 1. The Geological Society of America and The University of Kansas Press.

[VH01] Vachard, D., M. Hauser, R. Martini, L. Zaninetti, A. Matter & T. Peters. 2001. New algae and problematica of algal affinity from the Permian of the Aseelah Unit of the Batain Plain (East Oman). Geobios 34 (4): 375–404.