Byttneria aculeata, copyright Heike Vibrans.

Belongs within: Malvales.
Contains: Rulingia, Malacothamnus, Sidalcea, Sida, Sphaeralcea, Bombacoideae, Malvoideae, Sterculioideae, Tilioideae, Dombeyoideae, Helicteroideae, Corchorus, Triumfetta, Grewia, Hermannia, Lasiopetalum.

The Malvaceae as used here in the broad sense is an assemblage of plants previously divided between the families Malvaceae, Bombacaceae, Sterculiaceae and Tiliaceae; polyphyly of the latter three families has lead to their inclusion in the first. Molecular data support a basal division of the Malvaceae sensu lato between Byttneriina, including taxa previously assigned to Sterculiaceae (Byttnerioideae) and Tiliaceae (Grewioideae), and Malvadendrina, including taxa previously assigned to all four original families (Baum et al. 1998). The Byttnerioideae, including the largest portion of the historical Sterculiaceae, are characterised by perfect, actinomorphic, pentamerous flowers with stamens in multiples of five and opposite the petals.

Characters (from the Angiosperm Phylogeny Website): Shrubs to trees (occasionally herbs); cyclopropenoid fatty acids, terpenoid-based quinones present, gums common; wood commonly fluoresces; pits not vestured; tile cells common; sieve tubes with non-dispersive protein bodies; hairs stellate/lepidote; leaves spiral or two-ranked, lamina vernation usually conduplicate(to plicate), margins entire or toothed, single vein running to the non-glandular apex, secondary venation palmate; inflorescence made up of modified cymose units ["bicolor units"]; calyx valvate; stamens (5-)many, in five groups opposite the petals, but fundamentally obdiplostemonous, basally connate, extrorse; tapetal cells 2(-4)-nucleate; gynoecium variable in orientation, style usually 5-branched apically, stigma usually dry; ovules 1-many carpel; micropyle zig-zag (endo- or exostomal), outer integument develops first, 2-4(-7) cells across, inner integument 2-7(-10) cells across, parietal tissue 3-8 cells, across, nucellar cap 2-5 cells across, hypostase present, obturator present or absent; inverse postament present; fruit a capsule (berry, schizocarp, etc.; muricate); testa multiplicative, vascularized, endotesta crystalliferous, tegmen multiplicative, endotegmic cells more or less thickened; endosperm often starchy, embryo often green; sporophytic self-incompatibility system often present.

<==Malvaceae [Bombacaceae, Eumalvales, Malvineae, Sterculiaceae, Sterculiineae]
    |--Malvadendrina BAN98
    |    |--Malvatheca BS04
    |    |    |--+--Bombacoideae BS04
    |    |    |  `--Malvoideae BS04
    |    |    `--+--Patinoa almirajo BS04, BAN98
    |    |       `--Ochroma BS04
    |    |            |--O. lagopus K06
    |    |            `--O. pyramidale BAN98
    |    `--+--Sterculioideae BS04
    |       `--+--+--Tilioideae BS04
    |          |  `--Dombeyoideae BS04
    |          `--+--Helicteroideae BS04
    |             `--Brownlowioideae BAN98
    |                  |--Mortoniodendron BAN98
    |                  `--+--Berrya [Berryeae] BAN98
    |                     |--Brownlowia [Brownlowieae] BS04
    |                     |    `--B. elata BAN98
    |                     |--Christiana BAN98
    |                     `--Jarandersonia BAN98
    `--Byttneriina [Byttneriaceae] BAN98
         |  i. s.: Oudemansia Miq. 1854 KC01
         |--Grewioideae BAN98
         |    |--Corchorus BAN98
         |    |--Sparmannia africana BAN98
         |    |--Triumfetta BAN98
         |    |--Grewia BAN98
         |    `--Heliocarpus BAN98
         |         |--H. americanus BAN98
         |         `--H. tomentosus BT87
         `--Byttnerioideae BAN98
              |--Hermannia BAN98
              |--Kleinhovia hospita BAN98, Cl78
              |--Leptonychia BAN98
              |--Lasiopetalum BAN98
              |--Theobroma [Theobromeae] BAN98
              |    |--T. bicolor BW64
              |    `--T. cacao BAN98
              `--Byttneria BAN98
                   |--*B. scabra BAN98
                   |--B. aculeata F09
                   |--B. carthaginensis RS99
                   `--B. herbacea P03

Malvaceae incertae sedis:
  Melochia D01
    |--M. corchorifolia LK14
    |--M. pyramidata LK14
    `--M. umbellata LK14
  Argyrodendron B00a
    |--A. peralatum W01
    `--A. trifoliolatum B00a [=Heritiera trifoliolata H90]
  Pterygota P02
  Melhania LK14
    |--M. futteyporensis PP07
    |--M. incana C16
    `--M. oblongifolia LK14
  Acropogon YY22
  Basiloxylon YY22
  Dicarpidium monoicum LK14
  Firmiana S02
    |--F. colorata S02
    |--F. malayana P88
    `--F. simplex [incl. Sterculia platanifolia] LO98
  Octolobus YY22
  Pterocymbium YY22
  Tarrietia argyrodendron YY22, H42
  Erythropsis Lindley ex Schott & Endlicher 1832 FT93
  Thomasia RL05
    |--T. cognata RL05
    |--T. foliosa GK00
    |--T. macrocalyx OS04
    |--T. paniculata GK00
    `--T. pauciflora GK00
  Keraudrenia H90
    |--K. corollata H90
    |    |--K. c. var. corollata H90
    |    `--K. c. var. denticulata H90
    |--K. hillii H90
    |--K. integrifolia H90
    `--K. velutina G04
         |--K. v. ssp. velutina G04
         `--K. v. ssp. elliptica G04
  Rulingia GK00
  Commersonia B00b
    |--C. bartramia Cr78
    `--C. fraseri B00b
  Hannafordia H90
    |--H. bissillii H90
    `--H. quadrivalvis KM08
  Malvastrum M99
    |--M. americanum [incl. M. spicatum] H90
    |--M. coromandelianum [incl. M. tricuspidatum] H90
    `--M. dissectum C01
  Ambroma augusta BB07
  Parabombacaceoxylon WM09
  Javelinoxylon WM09
  Abelmoschus M99
    |--A. esculentus M99
    |--A. ficulneus M99
    |--A. manihot H90
    `--A. moschatus LK14
  Fioria vitifolia LK14
  Waltheria LK14
    |--W. americana BTA75
    `--W. indica LK14
  Modiola H93
    |--M. caroliniana H93
    `--M. multifida C06
  Fugosia australis C16
  Eremalche H93
    |--E. exilis [=Malvastrum exile] H93
    |--E. parryi [=Malvastrum parryi] H93
    |    |--E. p. ssp. parryi H93
    |    `--E. p. ssp. kernensis [=Malvastrum kernense] H93
    `--E. rotundifolia [=Malvastrum rotundifolium] H93
  Herissantia crispa [=Abutilon crispum] H93
  Horsfordia H93
    |--H. alata H93
    `--H. newberryi H93
  Iiamna H93
    |--I. bakeri H93
    `--I. latibracteata H93
  Malacothamnus H93
  Malvella leprosa H93 [=Sida leprosa H90; incl. S. leprosa var. hederacea H93]
  Ayenia compacta [incl. A. californica] H93
  Bombacites formosus Berry 1916 CBH93
  Bombacoxylon CBH93
  Hibiscoxylon niloticum Kräusel 1939 CBH93
  Kosteletzkya CBH93
  Echiperiporites estelae CBH93
  Chattawaya paliformis Manchester 1980 CBH93
  Triplochitioxylon oregonensis Manchester 1979 CBH93
  Reevesiapollenites CBH93
  Gilesia biniflora H90
  Seringia arborescens H90
  Gynatrix pulchella [=Plagianthus pulchellus; incl. P. pulchellus var. tomentosus] H90
  Stegia DC. 1805 KC01
  Mansonia altissima S77
  Azanza lampas M72
  Sidalcea PT01
  Cephalohibiscus H03
  Kydia calycina P03
  Napaea YY22
  Anoda RN-F01
    |--A. cristata [incl. A. cristata var. digitata] H93
    |--A. hastata J87
    `--A. parviflora Cavanilles 1799 C99
  Lawrencia G04
    |--L. densiflora KM08
    |--L. glomerata [=Plagianthus glomeratus] H90
    |--L. spicata GK00
    |--L. squamata [=Plagianthus squamatus, Selenothamnus squamatus; incl. P. incanus] H90
    `--L. viridigrisea KM08
  Althea PT01
    |--A. cannabina H91
    |--A. hirsuta PT98
    |--A. officinalis PT01
    `--A. rosea KSM06
  Urena LK14
    |--U. americana [incl. U. reticulata] C55
    |--U. armitiana LK14
    |    |--U. a. var. armitiana LK14
    |    `--U. a. var. spenceri LK14
    `--U. lobata LK14
  Alcea Y98
    |--A. biennis PT98
    `--A. rosea [=Althaea rosea] H93
  Sida G04
  Sphaeralcea J87

*Type species of generic name indicated


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

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[H91] Hubálek, Z. 1991. Biogeographic indication of natural foci of tick-borne infections. In: Dusbábek, F., & V. Bukva (eds) Modern Acarology: Proceedings of the VIII International Congress of Acarology, held in České Budĕjovice, Czechoslovakia, 6–11 August 1990 vol. 1 pp. 255–260. SPB Academic Publishing: The Hague.

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Last updated: 26 July 2021.


Dorsum and venter of Monogalumnella neotricha, from Balogh & Balogh (1992 vol. 2).

Belongs within: Circumdehiscentiae.

The Galumnellidae are a group of galumnoid oribatid mites with slender chelicerae and an acuminate rostrum. The notogaster of galumnellids is commonly sculptured, though it is is smooth in Galumnopsis sensu stricto (Balogh & Balogh 1992).

Characters (from Balogh & Balogh 1992): Chelicerae peloptoid; rostrum acuminate. Prodorsum without true projecting lamellae, only with sclerotised lines on surface. Pteromorphae large, auriculate, movable (hinged).

    |--Monogalumnella Mahunka 1986 S04
    |    `--*M. neotricha Mahunka 1986 S04
    |--Iberogalumnella Arillo & Subías 1993 S04
    |    `--*I. alandalusica Arillo & Subías 1993 S04
    |--Trypogalumnella Mahunka 1995 S04
    |    |--*T. poronota Mahunka 1995 S04
    |    `--T. densoporosa Mahunka 1995 S04
    |--Galumnopsis Grandjean 1931 S04
    |    |--G. (Galumnopsis) S04
    |    |    |--*G. (G.) holoscripta Grandjean 1931 S04
    |    |    |--G. (G.) clavata Mahunka 1983 S04
    |    |    |--G. (G.) longisetus Mahunka 1983 S04
    |    |    |--G. (G.) rostrata Balogh 1962 S04
    |    |    |--G. (G.) ruginervis Balogh 1962 S04
    |    |    |--G. (G.) sagitta Balogh 1970 S04
    |    |    |--G. (G.) secunda (Sellnick 1923) [=Galumnella secunda] S04
    |    |    `--G. (G.) sellnicki Balogh 1960 S04
    |    `--G. (Porogalumnella Balogh 1968) S04
    |         |--G. (*P.) quadriporosa (Balogh 1968) [=*Porogalumnella quadriporosa] S04
    |         |--G. (P.) africana (Mahunka 1978) [=Porogalumnella africana] S04
    |         |--G. (P.) pulchella (Aoki & Hu 1993) [=Porogalumnella pulchella] S04
    |         |--G. (P.) reducta (Mahunka 1995) [=Porogalumnella reducta] S04
    |         `--G. (P.) setosa (Balakrishnan & Haq 1982) [=Porogalumnella setosa] S04
    `--Galumnella Berlese 1916 S04
         |  i. s.: G. indica Balakrishnan 1989 H01
         |--G. (Galumnella) S04
         |    |--*G. (G.) paradoxa Berlese 1916 S04
         |    |--G. (G.) apiculata Mahunka 1992 S04
         |    |--G. (G.) areolata Balogh 1960 S04
         |    |--G. (G.) cellularis Balogh & Mahunka 1967 S04
         |    |--G. (G.) geographica Mahunka 1995 S04
         |    |--G. (G.) nipponica Suzuki & Aoki 1970 [incl. G. angustifrons Aoki 1970] S04
         |    |--G. (G.) pauliani Balogh 1961 S04
         |    |--G. (G.) pilosa Engelbrecht 1972 S04
         |    |--G. (G.) punctipennis Balogh 1960 S04
         |    |--G. (G.) rugosa Balogh 1960 S04
         |    |--G. (G.) rugosula Balogh 1960 S04
         |    |--G. (G.) subareolata Mahunka 1969 S04
         |    |--G. (G.) tanzanica Mahunka 1984 S04
         |    `--G. (G.) woschitzi Balogh 1970 S04
         `--G. (Bigalumnella Mahunka 1994) S04
              `--G. (*B.) csavasorum (Mahunka 1994) [=*Bigalumnella csavasorum] S04

*Type species of generic name indicated


Balogh, J., & P. Balogh. 1992. The Oribatid Mites Genera of the World vol. 1. Hungarian Natural History Museum: Budapest.

[H01] Haq, M. A. 2001. Potential oribatid mite vectors of cestode parasites in Kerala, India. 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. 569–575. CSIRO Publishing: Melbourne.

[S04] Subías, L. S. 2004. Listado sistemático, sinonímico y biogeográfico de los ácaros oribátidos (Acariformes, Oribatida) del mundo (1758–2002). Graellsia 60 (número extraordinario): 3–305.


Cannartus tubarius, from here.

Belongs within: Corticata.

The Lithocyclia lineage of radiolarians originated in the middle Eocene with discoidal forms (Lithocyclia) that later evoleved into ellipsoidal forms (Cannartus and Ommatartus).

Characters (from Sanfilippo & Riedel 1980, as Coccodiscidae): Discoidal forms consisting of a lenticular cortical shell enclosing a small single or double medullary shell, and surrounded by an equatorial zone of spongy or concentrically-chambered structure, OR forms with ellipsoidal cortical shell, usually equatorially constricted and enclosing a single or double medullary shell, the opposite poles of the shell generally bearing spongy columns and/or single or multiple latticed caps.

    |--L. ocellus P98
    `--+--L. aristotelis P98
       `--+--L. angusta P98
          `--Cannartus P98
               |--C. tubarius P98
               `--+--C. violina P98
                  `--+--C. mammiferus P98
                     `--+--C. laticonus P98
                        |--+--‘Ommatartus’ hughesi P98
                        |  `--C. petterssoni P98
                        `--Ommatartus P98
                             |--O. antepenultimus P98
                             `--+--O. penultimus P98
                                `--+--O. avitus P98
                                   `--O. tetrathalamus P98

*Type species of generic name indicated


[P98] Prothero, D. R. 1998. Bringing Fossils to Life: An introduction to paleobiology. WCB McGraw-Hill: Boston.

Sanfilippo, A., & W. Riedel. 1980. A revised generic and suprageneric classification of the artiscins (Radiolaria). Journal of Paleontology 54 (5): 1008-1011.


Campascus minutus, from here.

Belongs within: Thecofilosea.

The Psammonobiotidae is a family of shelled amoeboids that live in the interstitial habitat between sand grains. Most genera in the family are marine, though the majority of Campascus species inhabit fresh water (Nicholls 2003).

Characters (from Nicholls 2003): Test covered with irregularly-shaped, flat particles, apparently of exogenous origin. Flared collar present surrounding oral aperture.

    |--Edaphonobiotus N03
    |--Alepiella Golemansky 1970 N03, G86
    |    `--A. tricornuta Golemansky 1970 G86
    |--Chardezia Golemansky 1970 G86
    |    `--C. caudata Golemansky 1970 G86
    |--Corythionelloides Sudzuki 1979 G86
    |    `--C. fumiana Sudzuki 1979 G86
    |--Micropsammella Golemansky 1970 N03, G86
    |    |--M. minima Chardez & Thomas 1980 G86
    |    `--M. retorta Golemansky 1970 [incl. M. retorta var. japonica Sudzuki 1979] G86
    |--Psammonobiotus Golemansky 1967 G86
    |    |--P. balticus Golemansky 1973 G86
    |    |--P. communis Golemansky 1967 G86
    |    |--P. golemanskyi Chardez 1971 G86
    |    |--P. linearis Golemansky 1970 G86
    |    |--P. minutus Golemansky 1970 G86
    |    |--P. plana Chardez 1971 G86
    |    `--P. septentrionalis Chardez 1977 G86
    `--Campascus Leidy 1877 N03
         |--*C. cornutus Leidy 1877 N03
         |--C. bidens Krascheninnikov 1923 N03
         |--C. cyphoderiformis Chardez & Gaspar 1984 N03
         |--C. dentatus Edmondson & Kingman 1913 N03
         |--C. interstitialis Golemansky 1981 N03
         |--C. minutus Penard 1899 N03
         |--C. simcoei Nicholls 2003 N03
         |--C. triqueter Penard 1891 N03
         `--C. vulgaris Valkanov 1936 N03

*Type species of generic name indicated


[G86] Golemansky, V. G. 1986. Rhizopoda: Testacea. In Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) (L. Botosaneanu, ed.) pp. 5-16. E. J. Brill / Dr. W. Backhuys: Leiden.

[N03] Nicholls, K. H. 2003. Form variation in Campascus minutus and a description of Campascus simcoei sp. n. (Testaceafilosea, Psammonobiotidae). European Journal of Protistology 39 (1): 103-112.


Palaeoephippium bifurcum, from here.

Belongs within: Radiozoa.

The Palaeoscenidiidae are a family of radiolarians known from the Ordovician to the Jurassic.

Characters (from MacDonald 2004): Five or more rays from ends of median bar or point center, separated into apical and basal hemispheres, may be rays between hemispheres. With or without shell, formed by branching of basal rays, less commonly by all rays.

Palaeoscenidiidae [Entactinaria]
    |--Palaeoscenidium Deflandre 1953 M04
    |--Palaeotrifidus ballator Renz 1990 M04
    |--Fukujius yamakoshii Furutani 1990 M04
    |--Palaeotripus Goodbody 1986 M04
    |    `--P. sexabrachiatus Renz 1990 M04
    |--Palaeopyramidium Goodbody 1986 M04
    |    `--*P. spinosum Goodbody 1986 M04
    |--Holdsworthum Furutani 1990 M04
    |    `--*H. japonicus Furutani 1990 M04
    |--Protoentactinia Kozur, Mostler & Repetski 1996 (see below for synonymy) M04
    |    |--*P. gracilispinosa Kozur, Mostler & Repetski 1996 M04
    |    `--P. tricorne (Goodbody 1986) [=Palaeoephippium tricorne] M04
    |--Goodbodium Furutani 1990 M04
    |    |--*G. flammatum (Goodbody 1986) [=Palaeoscenidium flammatum; incl. G. elegans Furutani 1990] M04
    |    |--G. nishiyamai Furutani 1990 M04
    |    `--G. rarispinosum (Goodbody 1986) [=Palaeoscenidium rarispinosum] M04
    |--Insolitignum MacDonald 1999 M04
    |    |--*I. dissimile (Goodbody 1986) [=Palaeoephippium dissimile; incl. I. peraniuma MacDonald 1999] M04
    |    |--I. cancellatum (Goodbody 1986) (see below for synonymy) M04
    |    `--I. vivanima MacDonald 1999 M04
    |--Palaeodecaradium Goodbody 1986 M04
    |    |--*P. umbelliforme Goodbody 1986 M04
    |    |--P. apertum (Goodbody 1986) [=Palaeoscenidium apertum; incl. Palaeotripus nudus Goodbody 1986] M04
    |    `--P. gordoni MacDonald 2004 M04
    `--Palaeoephippium Goodbody 1986 M04
         |--*P. bifurcum Goodbody 1986 [incl. P. aranea Goodbody 1986] M04
         |--P. echinatum Goodbody 1986 M04
         |--‘Palaeotripus’ impariramosus Goodbody 1986 M04
         |--P. multiramosum Goodbody 1986 M04
         |--P. octoramosum Renz 1990 M04
         |--P. radices Goodbody 1986 [incl. P. ramipendentes Goodbody 1986, P. reteforme Goodbody 1986] M04
         `--P. spinosum Goodbody 1986 [incl. Palaeotripus monospinosum Goodbody 1986] M04

Insolitignum cancellatum (Goodbody 1986) [=Palaeoscenidium cancellatum; incl. Palaeoephippium adraini MacDonald 1999, Palaeotripus cancellatus Goodbody 1986] M04

Protoentactinia Kozur, Mostler & Repetski 1996 [incl. Noblella Kozur, Mostler & Repetski 1996; Protoentactiniidae] M04

*Type species of generic name indicated


[M04] MacDonald, E. W. 2004. Palaeoscenidiidae (Radiolaria) from the Lower Silurian of the Cape Phillips Formation, Cornwallis Island, Nunavut, Canada. Journal of Paleontology 78: 257-274.


Lateral view, dorsal view of aspis, and ventral view of hysterosoma of Prototritia (Siciliophora) sicula, from Balogh & Balogh (1992, vol. 2).

Belongs within: Enarthronota.

The Protoplophoridae are a cosmopolitan family of oribatid mites with a ptychoid body form, able to withdraw the legs and close the aspis against the notogaster and ventral region, and with the notogaster divided by three transverse sutures (Balogh & Balogh 1992). A telescoping type L scissure divides the notogaster between the c and d setal rows.

<==Protoplophoridae [Protoplophorinae]
    |--Aedoplophora Grandjean 1932 NB-P09, S04
    |    `--A. glomerata Grandjean 1932 NB-P09, S04 [=Prototritia (Prototritia) glomerata S04]
    |--Cryptoplophora Grandjean 1932 S04
    |    |--*C. abscondita Grandjean 1932 S04
    |    `--C. asiatica Gordeeva, Niemi & Petrova-Nikitina 1998 S04
    |--Protoplophora Berlese 1910 S04
    |    |--*P. palpalis Berlese 1910 [incl. P. grandjeani Bernini 1983] S04
    |    `--P. oglasicola Bernini 1983 S04
    |--Arthrhoplophora Berlese 1910 [incl. Neoprototritia Shereef 1978, Triplophora Mahunka 1977] S04
    |    |--*A. paradoxa Berlese 1910 S04
    |    |--A. vulpes Berlese 1916 [incl. A. (Triplophora) berlesei Mahunka 1977] S04
    |    `--A. zachvatkini (Shereef 1978) [=Neoprototritia zachvatkini] S04
    |--Grandjeanoplophora Balogh & Mahunka 1979 S04
    |    |--*G. (Grandjeanoplophora) mauritanica (Grandjean 1932) [=Cryptoplophora mauritanica] S04
    |    `--G. (Tauroplophora Gordeeva, Niemi & Petrova-Nikitina 1998) S04
    |         `--G. (*T.) aureonotata (Gordeeva, Niemi & Petrova-Nikitina 1998) [=Tauroplophora aureonotata] S04
    |--Hauseroplophora Mahunka 1977 S04
    |    |--*H. soniae Mahunka 1977 S04
    |    |--H. flagellata Mahunka 1987 S04
    |    `--H. phitosi Mahunka 1977 S04
    |--Bursoplophora Subías & Pérez-Íñigo 1978 [incl. Csibiplophora Mahunka 1984] S04
    |    |--*B. iberica Subías & Pérez-Íñigo 1978 S04
    |    |--B. ethiopica Mahunka 1982 S04
    |    |--B. genavensium (Mahunka 1984) [=Csibiplophora genavensium] S04
    |    |--B. insularis Kahwash, Subías & Ruiz 1989 S04
    |    |--B. madagassica Mahunka 1994 S04
    |    |--B. meridionalis Bernini 1983 S04
    |    |--B. muraiae Mahunka & Mejía-Recamier 1998 S04
    |    `--B. tyrrhenica Bernini 1983 S04
    `--Prototritia Berlese 1916 S04
         |--P. (Prototritia) S04
         |    |--*P. (P.) armadillo (Berlese 1916) S04 (see below for synonymy)
         |    |--P. (P.) africana (Mahunka 1977) [=Aedoplophora africana] S04
         |    |--P. (P.) grandjeani (Mahunka 1977) [=Aedoplophora grandjeani] S04
         |    |--P. (P.) major (Jacot 1933) [=Aedoplophora major] S04
         |    `--P. (P.) palaciosi Mahunka & Mejía-Recamier 1998 S04
         `--P. (Siciliophora Bernini 1983) S04
              `--P. (*S.) sicula Bernini 1983 S04

*Prototritia (Prototritia) armadillo (Berlese 1916) S04 [=Arthrhoplophora (*P.) armadillo S04, Prothoplophora (*P.) armadillo BB92]

*Type species of generic name indicated


[BB92] Balogh, J. & P. Balogh. 1992. The Oribatid Mites Genera of the World vol. 1. Hungarian Natural History Museum: Budapest.

[NB-P09] Norton, R. A., & V. M. Behan-Pelletier. 2009. Suborder Oribatida. In: Krantz, G. W., & D. E. Walter (eds) A Manual of Acarology 3rd ed. pp. 430–564. Texas Tech University Press.

[S04] Subías, L. S. 2004. Listado sistemático, sinonímico y biogeográfico de los ácaros oribátidos (Acariformes, Oribatida) del mundo (1758–2002). Graellsia 60 (número extraordinario): 3–305.


Latero-ventral (A), dorsal (C) and basal (B, D) views of two specimens of Saitoum ambiguum, from Dumitrica & Zügel (2003).

Belongs within: Nassellaria.

Saitoum is a genus of radiolarian found from the Jurassic to the Cretaceous. Members of this genus have a globular cephalis with a distinct apical horn, and without verticils of spines on the feet (Dumitrica & Zügel 2003).

Saitoum Pessagno 1977 DZ03
    |--*S. pagei Pessagno 1977 DZ03
    |--S. ambiguum Dumitrica & Zügel 2003 DZ03
    |--S. bicorniculatum Dumitrica & Zügel 2003 DZ03
    |--S. curtipes Dumitrica & Zügel 2003 DZ03
    |--S. delicatum Dumitrica & Zügel 2003 DZ03
    |--S. dercourti Widz & de Wever 1993 DZ03
    |--S. differtum Dumitrica & Zügel 2003 DZ03
    |--S. elegans de Wever 1981 DZ03
    |--S. kapewinteri Dumitrica & Zügel 2003 DZ03
    |--S. labeosum Dumitrica & Zügel 2003 DZ03
    |--S. levium de Wever 1981 DZ03
    |--S. macilentum Dumitrica & Zügel 2003 DZ03
    |--S. reticulatum Dumitrica & Zügel 2003 DZ03
    `--S. virgulispina Dumitrica & Zügel 2003 DZ03

*Type species of generic name indicated


[DZ03] Dumitrica, P., & P. Zügel. 2003. Lower Tithonian mono- and dicyrtid Nassellaria (Radiolaria) from the Solnhofen area (southern Germany). Geodiversitas 25 (1): 5-72.


Anaphothrips sudanensis, from here.

Belongs within: Thripini.

Anaphothrips is a cosmopolitan genus of thrips associated with plant leaves, with recorded hosts including Poales, Asparagales, Lamiales, Asteraceae and Chenopodiaceae (Mound & Masumoto 2009).

Characters (from Mound & Masumoto 2009): Apterous, micropterous or macropterous; antennae 8- or 9-segmented, sensorium on III forked (simple in some species), on IV forked; III–VI usually with microtrichia present, often annulate; segment I without dorso-apical setae. Head often slightly projecting in front of eyes; 3 pairs of ocellar setae present; 3 or 4 pairs of postocular setae; eyes large, with or without 6 pigmented facets; maxillary palps 3-segmented, mouth cone relatively long, often slender and pointed at apex. Pronotum with no long setae (one species with posteromarginal S2 longer than remaining setae); ferna usually entire; basantra without setae. Mesonotal campaniform sensilla usually present but often small and inconspicuous. Metascutum with reticulate sculpture. Metapre-episternum usually weakly developed. Mesothoracic sternopleural sutures complete, furca with spinula; metathoracic furca without spinula. Tarsi 2-segmented. Fore wing with setae short, first vein with setal row irregularly spaced, second vein with many setae irregularly spaced; marginal cilia wavy; clavus with 4–9 marginal but no discal setae. Abdominal tergites and sternites without posteromarginal craspeda; tergites without lateral ctenidia, with median setae rarely close together, campaniform sensilla far from posterior margin; VI–VII usually with setae S3 reduced and much smaller than S4; setae on posterior tergites not elongate; tergite IX with two pairs of campaniform sensilla, mid-dorsal setae short and arising laterally; tergite X with dorsal split in distal half; sternites without discal setae; sternites III–VII with 3 pairs of marginal setae, median pair on VII ahead of posterior margin. Males (where known) with pore plates on sternite III or III–VII behind antecostal line, often large and C-shaped, but sometimes small, not C-shaped, or absent; tergite IX with or without dorsal paired thorn-like setae.

Anaphothrips Uzel 1895 [incl. Nakaharathrips Retana-Salazar 2007, Neophysopus Schmutz 1913] MM09
    |--*A. obscurus (Müller 1776) MM09 (see below for synonymy)
    |--A. ambiguus (Girault 1927) [=Neophysopus ambiguus] MM09
    |--A. aptilotus Mound & Masumoto 2009 MM09
    |--A. astrolomi Pitkin 1978 MM09
    |--A. atriplicis Mound & Masumoto 2009 MM09
    |--A. augustae Mound & Masumoto 2009 MM09
    |--‘Neophysopus’ aureolus Girault 1928 G28
    |--A. barringtoni Mound & Masumoto 2009 MM09
    |--A. barrowi Mound & Masumoto 2009 MM09
    |--A. callani Mound & Masumoto 2009 MM09
    |--A. carlylei Girault 1928 MM09
    |--A. cecili Girault 1928 [incl. Hemianaphothrips (Anaphothrips) concinnus Morison 1931] MM09
    |--A. chortinus Mound & Masumoto 2009 MM09
    |--A. cobari Mound & Masumoto 2009 MM09
    |--A. cucurbiti Pitkin 1978 MM09
    |--A. dalbyi Mound & Masumoto 2009 MM09
    |--A. desleyae Mound & Masumoto 2009 MM09
    |--A. dubius (Girault 1926) MM09 (see below for synonymy)
    |--A. epacrida Mound & Masumoto 2009 MM09
    |--A. eremophilae Mound & Masumoto 2009 MM09
    |--A. exocarpi Pitkin 1978 MM09
    |--A. exocarpoides Mound & Masumoto 2009 MM09
    |--‘Neophysopus’ fragariae Girault 1927 G27
    |--A. geijerae Mound & Masumoto 2009 MM09
    |--A. geleznowiae Mound & Masumoto 2009 MM09
    |--A. gillespiei Mound & Masumoto 2009 MM09
    |--A. glenysae Mound & Masumoto 2009 MM09
    |--A. incertus (Girault 1929) [=Limothrips incertus] MM09
    |--‘Neophysopus’ io Girault 1927 G27
    |--A. keatsi (Girault 1926) [=Heterothrips keatsi] MM09
    |--A. monga Mound & Masumoto 2009 MM09
    |--A. moundi Pitkin 1978 MM09
    |--A. newmani Moulton in Moulton & Newman 1935 MM09
    |--A. nimbus Mound & Masumoto 2009 MM09
    |--A. occidentalis Pitkin 1978 MM09
    |--A. orchis Mound & Masumoto 2009 MM09
    |--A. parsonsiae Mound & Masumoto 2009 MM09
    |--A. pulteneae Mound & Masumoto 2009 MM09
    |--A. secticornis PL12
    |--A. sudanensis Trybom 1911 MM96 (see below for synonymy)
    |--A. swezeyi Moulton 1928 MM09
    |--A. varii Moulton in Moulton & Newman 1935 MM09
    |--A. walchae Mound & Masumoto 2009 MM09
    |--A. westringiae Mound & Masumoto 2009 MM09
    |--A. whyalla Mound & Masumoto 2009 MM09
    |--A. woodi Pitkin 1978 MM09
    `--A. yalgooi Mound & Masumoto 2009 MM09

Anaphothrips dubius (Girault 1926) MM09 [=Heterothrips dubius MM09; incl. A. cinctiguttatus Girault 1927 MM09, A. regalis Girault 1928 MM09, Physothrips regalis GM79, Hemianaphothrips (Anaphothrips) tersus Morison 1930 MM09]

*Anaphothrips obscurus (Müller 1776) MM09 [=Thrips obscura MM09; incl. A. sexguttus Girault 1928 MM09, A. striatus (Osborn 1883) MM09, GM79, A. virgo MM96]

Anaphothrips sudanensis Trybom 1911 MM96 [=*Nakaharathrips sudanensis MM09; incl. A. bicinctus MM96, A. bicolor MM96, *Neophysopus medioflavus MM09]

*Type species of generic name indicated


[G27] Girault, A. A. 1927. Some new wild animals. Privately published (reprinted Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 208-210).

[G28] Girault, A. A. 1928. Some Insecta and a new All Highness. Privately published (reprinted Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 225-228).

[GM79] Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 1-400.

[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.

[MM09] Mound, L. A. & M. Masumoto. 2009. Australian Thripinae of the Anaphothrips genus-group (Thysanoptera), with three new genera and thirty-three new species. Zootaxa 2042: 1-76.

[PL12] Peñalver, E., C. C. Labandeira, E. Barrón, X. Delclòs, P. Nel, A. Nel, P. Tafforeau & C. Soriano. 2012. Thrips pollination of Mesozoic gymnosperms. Proceedings of the National Academy of Sciences of the USA 109 (22): 8623-8628.


Spiraling whitefly Aleurodicus dispersus, copyright Sam Fraser-Smith.

Belongs within: Sternorrhyncha.
Contains: Aleyrodinae.

The Aleyrodidae, whiteflies, are small plant-sucking bugs with immobile nymphs, some species of which are notorious crop pests. Adults are soft-bodied with reduced wing venation and are usually covered with a white, waxy powder (Carver et al. 1991).

Characters (from Grimaldi & Engel 2005): Adults usually covered with a mealy, flocculent wax, secreted by pairs of large glands on several abdominal sternites, spread over body using combs of hairs on hind legs. Antennae reduced to six or fewer segments; rostrum with four segments, segment two telescoping into segment one. Wing venation extremely reduced. First instar mobile; subsequent instars sedentary, with atrophied appendages; vasiform orifice present at tip of abdomen for expelling honeydew: anus opening into deep median fold in tergite eight, with lingula flicking away droplets forming therein; quiescent prepupal and pupal instars present prior to adulthood.

<==Aleyrodidae [Aleurodidae, Aleyrodina, Aleyrodoidea, Aleyrodomorpha, Scytinelytra]
    |--Heidea Schlee 1970 GE05, P92
    |    `--H. cretacica GE05
    |--Bernaea Schlee 1970 P92 [Bernaeidae, Bernaeinae GE05]
    |    `--B. neocomica GE05
    `--+--Siphonaleyrodinae S35
       |--Aleyrodinae S35
       |--Udamoselinae WEE70
       |    |--Udamoselis Enderlein 1909 S35
       |    `--Synaleurodicus Solomon 1935 WEE70, S35
       |         `--*S. hakeae Solomon 1935 S35
       `--Aleurodicinae S35
            |--Radialeurodicus Bondar 1923 S35
            `--Aleurodicus S35
                 |--A. antillensis SEH96
                 |--A. destructor CGW91 [incl. Aleyrodes albofloccosa (Froggatt 1918) WEE70, S35]
                 |--A. dispersus ZS10
                 |--A. dugesii Cockerell 1896 ZR96
                 `--A. holmesii (Maskell 1896) [=Aleurodes holmesii] K08

Aleyrodidae incertae sedis:
  Dialeurodes S35
    |--D. chittendeni SEH96
    |--D. citri S35
    |--D. citrifolii SEH96
    |--D. kirkaldyi CGW91
    `--D. pallidus G01
  Trialeurodes S35
    |--T. abutiloneus SEH96
    |--T. ferrnaldi SEH96
    |--T. floridensis SEH96
    |--T. packardi SEH96
    |--T. ricini SEH96
    |--T. vaporariorum S35
    `--T. variabilis SEH96
  Parabemisia myricae MHG04
  Aleuronympha K-P91
  Aleuroclava eucalypti CGW91
  Orchamoplatus citri CGW91
  Pealius LS96
    |--P. azaleae CGW91
    `--P. kelloggi LS96
  Aleurothrixus SEH96
    |--A. aepim M99
    |--A. floccosus [incl. A. howardi] SEH96
    |--A. porteri SEH96
    `--A. socialis M99
  Aleuroplatus SEH96
    |--A. coronata SEH96
    |--A. elemerae SEH96
    |--A. gelatinosus SEH96
    `--A. liquidambaris SEH96
  Aleuroglandulus malangae SEH96
  Crenidorsum SEH96
  Acaudaleyrodes citri SEH96
  Asterobemisia SEH96
    |--A. carpini SEH96
    `--A. paveli SEH96
  Bulgarialeurodes cotesii SEH96
  Siphoninus SEH96
    |--S. immaculatus SEH96
    `--S. phillyreae SEH96
  Aleurocybotus SEH96
    |--A. indicus SEH96
    `--A. occiduus SEH96
  Paraleyrodes SEH96
    |--P. naranjae SEH96
    `--P. perseae SEH96

*Type species of generic name indicated


[CGW91] Carver, M., G. F. Gross & T. E. Woodward. 1991. Hemiptera (bugs, leafhoppers, cicadas, aphids, scale insects etc.) In: CSIRO. The Insects of Australia: A textbook for students and research workers vol. 1 pp. 429–509. Melbourne University Press: Carlton (Victoria).

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[G01] Gupta, S. K. 2001. A conspectus of natural enemies of phytophagous mites and mites as potential biocontrol agents of agricultural pests in India. 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. 484–497. CSIRO Publishing: Melbourne.

[K08] Kirkaldy, G. W. 1908. A catalogue of the Hemiptera of Fiji. Proceedings of the Linnean Society of New South Wales 33: 345–391, pl. 4.

[K-P91] Kukalová-Peck, J. 1991. Fossil history and the evolution of hexapod structures. In: CSIRO. The Insects of Australia: A textbook for students and research workers vol. 1 pp. 141–179. Melbourne University Press: Carlton (Victoria).

[LS96] Liu, T.-X., & P. A. Stansly. 1996. Morphology of Nephaspis oculatus and Delphastus pusillus (Coleoptera: Coccinellidae), predators of Bemisia argentifolii (Homoptera: Aleyrodidae). Proceedings of the Entomological Society of Washington 98 (2): 292–300.

[MHG04] Maynard, G. V., J. G. Hamilton & J. F. Grimshaw. 2004. Quarantine—phytosanitary, sanitary and incursion management: an Australian entomological perspective. Australian Journal of Entomology 43 (3): 318–328.

[M99] Moraes, G. J. de. 1999. Pest status of the cassava green mite in Brazil and strategies for its control. In: Needham, G. R., R. Mitchell, D. J. Horn & W. C. Welbourn (eds) Acarology IX vol. 2. Symposia pp. 287–291. Ohio Biological Survey: Columbus (Ohio).

[P92] Poinar, G. O., Jr. 1992. Life in Amber. Stanford University Press: Stanford.

[SEH96] Schauff, M. E., G. A. Evans & J. M. Heraty. 1996. A pictorial guide to the species of Encarsia (Hymenoptera: Aphelinidae) parasitic on whiteflies (Homoptera: Aleyrodidae) in North America. Proceedings of the Entomological Society of Washington 98 (1): 1–35.

[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.

[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.

[ZS10] Zborowski, P., & R. Storey. 2010. A Field Guide to Insects in Australia 3rd ed. Reed New Holland: Sydney.

[ZR96] Zolnerowich, G., & M. Rose. 1996. A new species of Entedononecremnus (Hymenoptera: Chalcidoidea: Eulophidae) parasitic on the giant whitefly, Aleurodicus dugesii Cockerell (Homoptera: Aleyrodidae). Proceedings of the Entomological Society of Washington 98 (2): 369–373.

Last updated: 2 April 2022.


Great cuckoo dove Reinwardtoena reinwardtsi, photographed by Rob Hutchinson.

Belongs within: Columbaves.
Contains: Columbigallina, Aplopelia, Leptotila, Zenaida, Geotrygon, Macropygia, Patagioenas, Columba, Streptopelia, Stigmatopelia, Treroninae, Columbina.

The Columbidae include the pigeons and doves, a distinctive family of mostly herbivorous, strong-flying birds found worldwide. Earlier authors mostly divided this family between the Treroninae for the palaeotropical fruit pigeons and Columbinae for the remaining 'true pigeons', with a few more small subfamilies for certain distinctive taxa, but this scheme has not been supported by molecular phylogenetic analyses (Shapiro et al. 2002; Jetz et al. 2012). Instead, a clade of New World taxa including the genera Geotrygon (quail-doves), Leptotila and Zenaida (mourning doves) may form one of the basal divisions within the columbids. Its sister-clade, containing the remaining species, is most diverse in the Old World, though it does include the New World ground doves (Claravis, Columbina and related genera). The now-extinct passenger pigeon Ectopistes migratorius of North America may be a relatively basal member of this clade; the passenger pigeon was a slender, long-winged and -tailed pigeon adapted for strong, rapid flight.

Characters (from Austin 1961): Stout-bodied, neck rather short, head small. Bill short, slender, rounded, usually thickening toward tip and thinner in the middle; nostrils emerging through fleshy cere at base of bill. Plumage a thick, heavy coat of strong-shafted feathers, loosely attached to thin skin so as to drop out easily.

Columbidae (see below for synonymy)
    |--+--+--+--‘Geotrygon’ veraguensis BKB15
    |  |  |  `--Leptotila SS02
    |  |  `--+--Zenaida SS02
    |  |     `--Geotrygon SS02
    |  `--+--+--‘Geotrygon’ purpurata BKB15
    |     |  `--‘Geotrygon’ saphirina BKB15
    |     `--+--‘Geotrygon’ versicolor BKB15
    |        `--+--‘Geotrygon’ montana BKB15
    |           |    |--G. m. montana MS55
    |           |    `--G. m. martinica MS55
    |           `--‘Geotrygon’ violacea BKB15
    `--+--+--Ectopistes Swainson 1827 BKB15, B94 [Ectopistinae]
       |  |    `--E. migratorius SS02
       |  `--+--+--Macropygia SS02
       |     |  `--Reinwardtoena JT12
       |     |       |--R. browni JT12
       |     |       |--R. crassirostris JT12
       |     |       `--R. reinwardtsi JT12
       |     |            |--R. r. reinwardtsi T62
       |     |            |--R. r. albida T62
       |     |            |--R. r. brevis T62
       |     |            `--R. r. griseotincta T62
       |     `--+--Patagioenas BKB15
       |        `--+--Columba SS02
       |           `--+--Streptopelia SS02
       |              `--+--Stigmatopelia JT12
       |                 `--Nesoenas SS02
       |                      |--N. mayeri SS02
       |                      `--N. picturata JT12 [=Streptopelia picturata BKB15, Turtur picturatus S66]
       |                           |--N. p. picturata L81
       |                           |--‘Streptopelia’ p. aldabrana L81
       |                           |--‘Streptopelia’ p. chuni L81
       |                           |--‘Streptopelia’ p. comorensis L81
       |                           |--‘Streptopelia’ p. rostrata L81
       |                           `--‘Streptopelia’ p. saturata L81
       `--+--Treroninae SS02
          `--+--Claravis Oberholser 1899 JT12, B94 [incl. Peristera Swainson 1827 B94]
             |    |--C. geoffroyi HSS13
             |    |--C. godefrida JT12
             |    |--C. mondetoura JT12
             |    |--C. pretiosa JT12
             |    |--‘Peristera’ principalis Hartlaub in Dohrn 1866 D66
             |    |--‘Peristera’ simplex D66
             |    `--‘Peristera’ tympanistria S66
             `--+--Uropelia campestris JT12
                `--+--Columbina SS02
                   `--Metriopelia JT12
                        |--M. aymara JT12
                        `--+--M. melanoptera JT12
                           `--+--M. ceciliae JT12 [=Gymnopelia ceciliae T62]
                              |    |--M. c. ceciliae T62
                              |    |--M. c. obsoleta T62
                              |    `--M. c. zimmeri [=Gymnopelia ceciliae zimmeri] T62
                              `--M. morenoi JT12

Columbidae incertae sedis:
  Zenaidura T62
    |--Z. auriculata T62
    |    |--Z. a. auriculata T62
    |    `--Z. a. stenura T62
    |--Z. graysoni RN72
    `--Z. macroura T62
         |--Z. m. macroura T62
         |--Z. m. carolinensis T62
         `--Z. m. marginella T62
  Columbigallina T62
  Tympanistria tympanistria T62
    |--T. t. tympanistria T62
    `--T. t. fraseri T62
  Oreopeleia T62
    |--O. chiriquensis RN72
    |--O. chrysia RN72
    |--O. martinica RN72
    |--O. montana S18
    `--O. mystacea T62
         |--O. m. mystacea T62
         `--O. m. sabae T62
  Aplopelia JT12
  Coryphoenas T62
  Nesopelia galapagoensis RN72
  Oxypelia T62
  Lophophaps T62
    |--L. ferruginea Gould 1865 WS48 (see below for synonymy)
    `--L. leucogaster R87
  Osculatia T62
  Melopelia asiatica FS55
  Starnoenas Bonaparte 1838 [Starnoenadinae] B94
    `--S. cyanocephala JT12
  Erythroenas pulcherrima B66
  Scardafella SS02
    |--S. inca L81
    `--S. squammata (Lesson 1831) ME04
  Dysmoropelia dekarchiskos HSS13
  Rupephaps Worthy, Hand et al. 2009 CC10
    `--*R. taketake Worthy, Hand et al. 2009 CC10
  Calopelia puella PB27
    |--C. p. puella PB27
    `--C. p. brehmeri PB27

Columbidae [Claravinae, Columbae, Columbiformes, Columbinae, Columbini, Palumbinae, Peristeridae, Peristerinae, Turturidae]

Lophophaps ferruginea Gould 1865 WS48 [=Geophaps plumifera ferruginea M03; incl. L. ferruginea mungi Mathews 1912 WS48]

*Type species of generic name indicated


Austin, O. L., Jr. 1961. Birds of the World: A survey of the twenty-seven orders and one hundred and fifty-five families. Paul Hamlyn: London.

[B66] Bartlett, A. D. 1866. Notes on the breeding of several species of birds in the Society's gardens during the year 1865. Proceedings of the Zoological Society of London 1866: 76–79.

[B94] Bock, W. J. 1994. History and nomenclature of avian family-group names. Bulletin of the American Museum of Natural History 222: 1–281.

[BKB15] Burleigh, J. G., R. T. Kimball & E. L. Braun. 2015. Building the avian tree of life using a large-scale, sparse supermatrix. Molecular Phylogenetics and Evolution 84: 53–63.

[CC10] Checklist Committee (OSNZ). 2010. Checklist of the Birds of New Zealand, Norfolk and Macquarie Islands, and the Ross Dependency, Antarctica 4th ed. Ornithological Society of New Zealand and Te Papa Press: Wellington.

[D66] Dohrn, H. 1866. Synopsis of the birds of Ilha do Principe, with some remarks on their habits and descriptions of new species. Proceedings of the Zoological Society of London 1866: 324–332.

[FS55] Felten, H., & J. Steinbacher. 1955. Zur Vogelfauna von El Salvador. Senckenbergiana Biologica 36 (1–2): 9–19.

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

[JT12] Jetz, W., G. H. Thomas, J. B. Joy, K. Hartmann & A. Ø. Mooers. 2012. The global diversity of birds in space and time. Nature 491: 444–448.

[L81] Long, J. L. 1981. Introduced Birds of the World: The worldwide history, distribution and influence of birds introduced to new environments. Reed: Sydney.

[ME04] Mayr, G., & P. G. P. Ericson. 2004. Evidence for a sister group relationship between the Madagascan mesites (Mesitornithidae) and the cuckoos (Cuculidae). Senckenbergiana Biologica 84 (1–2): 1–17.

[MS55] Mertens, R., & J. Steinbacher. 1955. Die im Senckenberg-Museum vorhandenen Arten ausgestorbener, aussterbender oder seltener Vögel. Senckenbergiana Biologica 36 (3–4): 241–265.

[M03] Morcombe, M. 2003. Field Guide to Australian Birds 2nd ed. Steve Parish Publishing.

[PB27] Pilsbry, H. A., & J. Bequaert. 1927. The aquatic mollusks of the Belgian Congo, with a geographical and ecological account of Congo malacology. Bulletin of the American Museum of Natural History 53 (2): 69–602, pls 10–77.

[R87] Ramsay, E. P. 1887. List of Western Australian birds collected by Mr. Cairn, and Mr. W. H. Boyer-Bower, at Derby and its vicinity, with remarks on the species. Proceedings of the Linnean Society of New South Wales, series 2, 1 (4): 1085–1100.

[RN72] Rutgers, A., & K. A. Norris (eds) 1972. Encyclopaedia of Aviculture vol. 1. London, Blandford Press.

[S66] Schlegel, H. 1866. Communication from, on mammals and birds collected in Madagascar. Proceedings of the Zoological Society of London 1866: 419–426.

[SS02] Shapiro, B., D. Sibthorpe, A. Rambaut, J. Austin, G. M. Wragg, O. R. P. Bininda-Emonds, P. L. M. Lee & A. Cooper. 2002. Flight of the dodo. Nature 295: 1683.

[S18] Stone, W. 1918. Birds of the Panama Canal Zone, with special reference to a collection made by Mr. Lindsey L. Jewel. Proceedings of the Academy of Natural Sciences of Philadelphia 70: 239–280.

[T62] Tendeiro, J. 1962. Estudos sobre malófagos: revisão monográfica do género Columbicola Ewing (Ischnocera, Philopteridae). Memórias da Junta de Investigações do Ultramar, ser. 2, 32: 7–460.

[WS48] Whittell, H. M., & D. L. Serventy. 1948. A systematic list of the birds of Western Australia. Public Library, Museum and Art Gallery of Western Australia, Special Publication 1: 1–126.

Last updated: 29 August 2019.


Ruddy cuckoo dove Macropygia emiliana, photographed by Johannes Pfleiderer.

Belongs within: Columbidae.

Macropygia, the cuckoo doves, is a genus of pigeons found in southern Asia and Australasia. Members of this genus have brown plumage, usually more boldly barred in the females, and long, graduated tails.

<==Macropygia Swainson 1837 [Macropygiinae] B94
    |--M. amboinensis J06
    |    |--M. a. amboinensis J06
    |    |--M. a. albiceps J06
    |    |--M. a. quinkan M03
    |    |--M. a. robinsoni M03
    |    `--M. a. sanghirensis R02
    |--M. emiliana JT12
    |--M. mackinlayi JT12
    |--M. magna JT12
    |--M. nigrirostris JT12
    |--M. phasianella T62 [=M. amboinensis phasianella M03]
    |--M. ruficeps T62
    |    |--M. r. ruficeps T62
    |    `--M. r. assimilis T62
    |--M. rufipennis JT12
    |--M. tenuirostris JT12 [=M. phasianella tenuirostris T62]
    `--M. unchall T62
         |--M. u. unchall T62
         `--M. u. tusalia T62

*Type species of generic name indicated


[B94] Bock, W. J. 1994. History and nomenclature of avian family-group names. Bulletin of the American Museum of Natural History 222: 1-281.

[JT12] Jetz, W., G. H. Thomas, J. B. Joy, K. Hartmann & A. Ø. Mooers. 2012. The global diversity of birds in space and time. Nature 491: 444-448.

[J06] Johnstone, R. E. 2006. The birds of Gag Island, Western Papuan islands, Indonesia. Records of the Western Australian Museum 23 (2): 115-132.

[M03] Morcombe, M. 2003. Field Guide to Australian Birds, 2nd ed. Steve Parish Publishing.

[R02] Riley, J. 2002. Population sizes and the status of endemic and restricted-range bird species on Sangihe Island, Indonesia. Bird Conservation International 12: 53-78.

[T62] Tendeiro, J. 1962. Estudos sobre malófagos: Revisão monográfica do género Columbicola Ewing (Ischnocera, Philopteridae). Memórias da Junta de Investigações do Ultramar, ser. 2, 32: 7-460.