Ixia amethystina, from Lambley Nursery.

Belongs within: Iridaceae.

The Ixieae are a group of iridaceous plants in which the rootstock forms a corm; flowers are sessile, subtended basally by paired bracts, and usually arranged in distichous spikes.

<==Ixieae [Ixioideae]
    |--Gladiolus Linné 1753 M99, G77
    |    |--G. byzantinus HE80
    |    |--G. × cardinalis HE80
    |    |--G. caryophyllaceus RL05
    |    |--G. italicus [incl. G. seqetum] H93
    |    |--G. natalensis (Ecklon) Hooker 1831 [incl. G. psittacinus] HE80
    |    |--G. tristis H93
    |    `--G. undulatus Linnaeus 1767 [incl. G. cuspidatus] HE80
    `--Ixia MG06
         |  i. s.: I. maculata Linnaeus 1763 HE80
         |         I. paniculata HE80
         |         I. polystachya Linnaeus 1762 HE80
         `--I. sect. Dichone MG06
              |--I. amethystina Manning & Goldblatt 2006 MG06
              |--I. brevituba MG06
              |--I. curvata MG06
              `--I. trifolia MG06

*Type species of generic name indicated


[G77] Geerinck, D. J. L. 1977. Iridaceae. Flora Malesiana, Series I—Spermatophyta, Flowering Plants 8 (2): 77–84.

[HE80] Healy, A. J., & E. Edgar. 1980. Flora of New Zealand vol. 3. Adventive cyperaceous, petalous and spathaceous monocotyledons. P. D. Hasselberg, Government Printer: Wellington (New Zealand).

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

[MG06] Manning, J. C., & P. Goldblatt. 2006. New species of Iridaceae from the Hantam-Roggeveld Centre of Endemism, and the Bokkeveld, Northern Cape, South Africa. Bothalia 36 (2): 139–145.

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

[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.


Heliocheilus cramboides, copyright Donald Hobern.

Belongs within: Noctuidae.
Contains: Heliothis.

The Heliothinae, bollworms and related forms, are a group of moths whose larvae feed mostly on the reproductive organs (flowers, fruits and seeds) of plants, with some species being significant horticultural pests.

Characters (from Matthews 1999): Small to medium-sized, robust moths. Frons generally rounded, bulging with well-developed ventral lip. Labial palps moderately long, weakly upcurved. Antennal filiform or minimally biserrate in male, filiform in female. Compound eye smooth, unlashed. Head and thorax covered with mixed hairlike and spathulate scales. Hind wing venation trifine; vein M2, if present, reduced and lying in median fold of hind wing. Male genitalia with valve usually simple and strap-like, quite long and narrow, rather flat, often curved distally, usually lacking processes except sometimes with a small ampulla; apical corona usually consisting of one or more rows of spine-like setae, occasionally absent; sacculus generally not well distinguished. Aedeagus short or long, straight or slightly curved, often bearing a denticulate 'scobinate' patch at apex; scobinate patch may occur at base of vesica; vesica varying from simple and sac-like to long and coiled, usually a number of diverticula present from a 'basal pouch'; vesica may bear cornuti. Female genitalia with papillae anales (ovipositor) varying from soft and pad-like with may setae to sclerotised and knife-like with fewer setae; ductus bursae varying from short to long, opening posteriorly through ostium bursae, variably ribbed and sclerotised; appendix bursae usually present, with ductus seminalis lying at apex; fundus bursae usually bearing from one to four signa. Larvae with spiny skin; prothoracic lateral setae with transverse position in later instars.

Heliothinae [Heliothidae] M99
    |  i. s.: Eutricopis nexilis M99
    |         Heliothodes diminutiva M99
    |         Periphanes ZK11
    |--Pyrrhia umbra (Hufnagel 1766) M99, JP05
    |--Melaporphyria M99
    |--+--Protadisura M99
    |  `--+--+--Heliolonche M99
    |     |  `--Schinia M99
    |     |       |--S. dobla M99
    |     |       `--S. florida M99
    |     `--Adisura Moore 1881 [incl. Astonycha Turner 1920] M99
    |          |--*A. atkinsoni Moore 1881 M99
    |          |--A. cana M99
    |          |--A. litarga (Turner 1920) [=*Astonycha litarga] M99
    |          |--A. marginalis (Walker 1858) (see below for synonymy) M99
    |          |--A. parva M99
    |          `--A. stigmatica M99
    `--+--Heliothis M99
       |--+--Australothis Matthews 1991 [=Australoverpa Matthews 1991] M99
       |  |    |--*A. rubrescens (Walker 1858) (see below for synonymy) M99
       |  |    |--A. exopisso Matthews 1999 M99
       |  |    |--A. tertia (Roepke 1941) [=Heliothis tertia] M99
       |  |    `--A. volatilis M99
       |  `--Helicoverpa Hardwick 1965 M99
       |       |  i. s.: H. assulta (Guenée 1852) (see below for synonymy) M99
       |       |         H. fletcheri M99
       |       |         H. hardwicki Matthews 1999 M99
       |       |         H. prepodes (Common 1985) [=Heliothis prepodes] M99
       |       |--H. punctigera (Wallengren 1860) [=Heliothis punctigera; incl. Chloridea marmada Swinhoe 1918] M99
       |       `--+--H. gelotopoeon M99
       |          `--+--H. hawaiiensis M99
       |             `--+--*H. armigera (Hübner 1803–1808) M99 (see below for synonymy)
       |                `--+--H. confusa M99
       |                   `--H. zea M99
       `--Heliocheilus Grote 1865 [incl. Canthylidia Butler 1886, Raghuva Moore 1881, Rhodosea Grote 1883] M99
            |--*H. paradoxus Grote 1865 M99
            |--H. abaccheutus Matthews 1999 M99
            |--H. aberrans (Butler 1886) [=Heliothis aberrans] M99
            |--H. albipunctella M99
            |--H. albivenata (Montague 1914) (see below for synonymy) M99
            |--H. aleurota (Lower 1902) (see below for synonymy) M99
            |--H. atrilinea (Turner 1943) [=Canthylidia atrilinea] M99
            |--H. canusina (Swinhoe 1901) (see below for synonymy) M99
            |--H. cistella (Swinhoe 1901) [=Canthylidia cistella] M99
            |--H. cladotus Swinhoe 1901 (see below for synonymy) M99
            |--H. confertissima (Walker 1865) [=Leucania confertissima, *Raghuva confertissima] M99
            |--H. confundens (Warren 1913) [=Canthylidia confundens] M99
            |--H. cramboides (Guenée 1852) (see below for synonymy) M99
            |--H. eodora (Meyrick 1902) [=Heliothis eodora; incl. Canthylidia tenuistria Turner 1902] M99
            |--H. ferruginosa (Turner 1911) [=Canthylidia ferruginosa; incl. C. epigrapha Turner 1920] M99
            |--H. flavitincta (Lower 1908) [=Melicleptria flavitincta] M99
            |--H. halimolimnus Matthews 1999 M99
            |--H. ionola (Swinhoe 1901) [=Adisura ionola, Melicleptria ionola] M99
            |--*Rhodosea’ julia Grote 1883 M99
            |--H. melibaphes (Hampson 1903) [=Melicleptria melibaphes] M99
            |--H. mesoleuca (Lower 1902) [=Canthylidia mesoleuca, Melicleptria mesoleuca] M99
            |--H. moribunda (Guenée 1852) M99 (see below for synonymy)
            |--H. neurota (Lower 1903) (see below for synonymy) M99
            |--H. pallida (Butler 1886) (see below for synonymy) M99
            |--H. puncticulata (Warren 1913) [=Canthylidia puncticulata] M99
            |--H. ranalaetensis Matthews 1999 M99
            |--H. rhodopolia (Turner 1911) [=Canthylidia rhodopolia] M99
            |--H. thelycritus Matthews 1999 M99
            `--H. vulpinotatus Matthews 1999 M99

Adisura marginalis (Walker 1858) [=Anthophila marginalis; incl. Heliothis delicia Felder & Rogenhofer 1874, A. dulcis Moore 1881, A. purgata Warren 1913, A. similis Moore 1881] M99

*Australothis rubrescens (Walker 1858) [=Thalpophila rubrescens; incl. Chloridea aresca Turner 1911, C. caesia Warren 1913, Heliothis hyperchroa Turner 1920, C. rufa Warren 1913] M99

*Helicoverpa armigera (Hübner 1803–1808) M99 [=Noctua armigera M99, Heliothis (Helicoverpa) armigera M83; incl. N. barbara Fabricius 1794 (nom. rej.) M99, Helicoverpa armigera commoni Hardwick 1965, Heliothis conferta Walker 1857 M99, Helic. armigera conferta M99, Heliothis armigera ab. fusca Cockerell 1889 M99, Heliothis guidelii Costantino 1922 M99, Heliothis obsoleta (Fabricius 1793) M99, GM79, Heliothis pulverosa Walker 1857 M99, Heliothis rama Bhattacharjee & Gupta 1972 M99, Chloridea obsoleta ab. rufa Warren 1911 non C. rufa Warren 1913 M99, Heliothis uniformis Wallengren 1860 M99]

Helicoverpa assulta (Guenée 1852) [=Heliothis assulta; incl. Helic. assulta afra Hardwick 1965, Heliothis separata Walker 1857, Heliothis succinea Moore 1881, Heliothis temperata Walker 1857] M99

Heliocheilus albivenata (Montague 1914) [=Melicleptria albivenata; incl. M. bipartita Strand 1924, M. stramineipicta Strand 1924, Canthylidia zorophanes Turner 1929, C. zurophanes (l. c.)] M99

Heliocheilus aleurota (Lower 1902) [=Melicleptria aleurota, Canthylidia alleurota (l. c.); incl. C. cana Turner 1943, C. aleurota ab. spissata Warren 1913] M99

Heliocheilus canusina (Swinhoe 1901) [=Canthylidia canusina; incl. C. anemodes Lower 1902, C. canusina ab. intacta Warren 1913] M99

Heliocheilus cladotus Swinhoe 1901 [incl. Canthylidia cladotus ab. densata Warren 1913, C. cladotus ab. hyalina Warren 1913, C. cladotus ab. hyalosticta Warren 1913, C. cramboides ab. semigrisea Warren 1913] M99

Heliocheilus cramboides (Guenée 1852) [=Leucania cramboides; incl. Canthylidia capnoneura Turner 1932, C. discolor Warren 1913, Heliocheilus discolor, C. cramboides ab. intensa Warren 1913, C. nervosa Turner 1943, Heliothis neurias Meyrick 1902, Heliocheilus neurias, C. cramboides ab. semiochrea Warren 1913, C. sericea Warren 1913, C. sulphurea Warren 1913, C. sumbensis Warren 1913, C. venata Warren 1913, Heliocheilus venata] M99

Heliocheilus moribunda (Guenée 1852) M99 [=Leucania moribunda M99, Canthylidia moribunda P27; incl. C. arenosa Turner 1943 M99, Leucania invaria Walker 1856 M99, C. invaria pallescens Warren 1913 M99]

Heliocheilus neurota (Lower 1903) [=Tarache neurota; incl. Canthylidia clathrata Warren 1913, C. crocopepla Turner 1925, Oruza crocotoschema Turner 1936, C. eurhythma Turner 1915, C. clathrata ab. exesa Warren 1913, C. clathrata ab. notata Warren 1913, C. clathrata ab. pallida Warren 1913 non C. pallida Butler 1886] M99

Heliocheilus pallida (Butler 1886) [=*Canthylidia pallida; incl. C. albida Warren 1913, H. osmidus Swinhoe 1901, C. rosea Warren 1913] M99

*Type species of generic name indicated


[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.

[JP05] Jungmann, E., & U. Poller. 2005. Neunachweise und Wiederfunde für die Schmetterlingsfauna (Insecta; Lepidoptera) des Altenburger Landes. Mauritiana 19 (2): 317–326.

[M83] Martin, N. A. 1983. Miscellaneous observations on a pasture fauna: an annotated species list. DSIR Entomology Division Report 3: 1–98.

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

[P27] Philpott, A. 1927. The maxillae in the Lepidoptera. Transactions and Proceedings of the New Zealand Institute 57: 721–746.

[ZK11] Zahiri, R., I. J. Kitching, J. D. Lafontaine, M. Mutanen, L. Kaila, J. D. Holloway & N. Wahlberg. 2011. A new molecular phylogeny offers hope for a stable family level classification of the Noctuoidea (Lepidoptera). Zoologica Scripta 40 (2): 158–173.


Luculia gratissima, copyright A. J. T. Johnsingh.

Belongs within: Gentianales.
Contains: Canthium, Rubioideae, Ixoroideae, Cinchonoideae.

The Rubiaceae are a cosmopolitan group of flowering plants usually bearing radial flowers with four-lobed corollas, often associated in tight cymes or clusters.

Characters (from Hickman 1993): Annual, perennial, shrub, vine, or tree. Leaves generally opposite, entire; stipules generally on stalk, sometimes leaf-like (then leaves apparently whorled and stipules considered leaves), adjacent pairs sometimes fused. Inflorescence a cyme, panicle, cluster, or flower solitary, generally terminal and more or less axillary. Flower generally bisexual; calyx generally more or less four-lobed, sometimes absent; corolla generally radial, four-lobed; stamens epipetalous, alternate with corolla lobes, generally included; ovary generally inferior, chambers generally 2 or 4, style 1, more or less fused if 2. Fruit two or four nutlets or a berry, drupe, or capsule.

<==Rubiaceae [Cinchoneae, Coffeoideae, Coptosapelteae]
    |--Luculia RB01
    |    |--L. grandifolia RB01
    |    |--L. gratissima SK02
    |    `--L. pinceana RB01
    `--+--Rubioideae RB01
       |--Coptosapelta RB01
       |    |--C. diffusa BM00
       |    `--C. flavescens RB01
       `--+--Ixoroideae RB01
          `--Cinchonoideae RB01

Rubiaceae incertae sedis:
  Atractogyne YY22
  Siphonandrium YY22
  Alibertia edulis BJ99
  Amajoua YY22
  Anthispermum YY22
  Basanacantha YY22
  Byrsophyllum YY22
  Crocyllis YY22
  Duroia YY22
  Garapatica YY22
  Kotchubaea YY22
  Melanopsidium YY22
  Morindopsis YY22
  Stachyarrhena YY22
  Thieleodoxa YY22
  Bataprine YY22
  Bobea YY22
  Cuviera YY22
  Hodgkinsonia YY22
  Myonima YY22
  Pachystylus YY22
  Timonius timon YY22, WP99
  Bathysa YY22
  Pouchetia Richard ex de Candolle 1830 FT93
    `--P. gilletii BJ99
  Psydrax LK14
    |--P. odorata LK14
    |    |--P. o. ssp. odorata LK14
    |    `--P. o. ssp. arnhemica LK14
    |--P. pendulina LK14
    `--P. suaveolens G04
  Rutidea orientalis BM00
  Schenckia blumenaviensis BM00
  Aidia BJ99
    |--A. micrantha BJ99
    `--A. racemosa VT13
  Aoranthe penduliflora BJ99
  Bertiera breviflora BJ99
  Burchellia bubalina BJ99
  Calochone redingii BJ99
  Canthium BJ99
  Casasia clusiifolia BJ99
  Chomelia BJ99
  Cremaspora triflora BJ99
  Cubanola domingensis BJ99
  Didymosalpinx norae BJ99
  Euclinia longiflora BJ99
  Feretia aeruginescens BJ99
  Genipa americana BJ99
  Glossostipula concinna BJ99
  Gonzalagunia affinis BJ99
  Heinsia crinita BJ99
  Hippotis BJ99
  Kailarsenia ochreata BJ99
  Keetia zanzibarica BJ99
  Kraussia floribunda BJ99
  Ladenbergia pavonii BJ99
  Leptactina platyphylla BJ99
  Massularia acuminata BJ99
  Meyna BJ99
    |--M. laxiflora P03
    `--M. tetraphylla BJ99
  Oxyanthus BJ99
    |--O. pyriformis BJ99
    `--O. zanguebaricus BJ99
  Paracoffea melanocarpa BJ99
  Porterandia crosbyi BJ99
  Posoqueria latifolia BJ99
  Pseudosabicea arborea BJ99
  Psilanthus mannii BJ99
  Rachicallis americana BJ99
  Ramosmania rodriguesii BJ99
  Randia BJ99
    |--R. aculeata BJ99
    |--R. benthamina WB-P93
    |--R. fitzalani [=Atractocarpus fitzalani] BJ99
    |--R. karstenii MM96
    `--R. moorei BJ99
  Rosenbergiodendron longiflorum BJ99
  Rothmannia longiflora BJ99
  Sabicea BJ99
    |--S. speiosa M09
    `--S. villosa BJ99
  Sukunia longipes BJ99
  Tamridaea capsulifera BJ99
  Tarenna BJ99
    |--T. cymosa BJ99
    |--T. dallachiana LK14
    |    |--T. d. ssp. dallachiana LK14
    |    `--T. d. ssp. expandens LK14
    |--T. neurophylla BJ99
    `--T. pentamera LK14
  Tricalysia BJ99
    |--T. cryptocalyx BJ99
    |--T. malaccensis J06
    `--T. ovalifolia BJ99
  Plectronia Linnaeus 1767 BR65
    `--P. latifolia [=Canthium latifolium] BR65
  Mastixiodendron pachyclados (Sch.) Melch. 1925 [=Fagraea pachyclados Sch. in Sch. & Laut. 1905] C95
  Chione venosa J87
  Stevensia hotteana J87
  Xeromphis spinosa KJ05
  Thecagonum ovalifolium P03
  Wendlandia P03
    |--W. exserta P03
    `--W. tinctoria P03
  Caelospermum reticulatum B00
  Kelloggia galioides H93
  Ixorophyllum anceps CBH93
  Paleorubiaceophyllum eocenicum (Berry) Roth & Dilcher 1979 CBH93
  Rubiaceocarpum markgrafii Kräusel 1939 [=R. markgrafi] CBH93
  ‘Triporotetradites’ nachterstedtensis CBH93
  Retiniphyllum AS02
  Ravnia triflora [incl. Lagenanthus parviflorus Ewan 1952] SK02
  Didymochlamys Hook. 1872 KC01
  Disperma Gmel. 1792 KC01
  Higginsia Pers. 1805 KC01
  Jackia Wall. 1823 KC01
  Piringa Juss. 1820 KC01
  Psathyra Spreng. 1818 KC01
  Schachtia Karst. 1859 KC01
  ‘Ludwigia’ erigata Linné 1767 [incl. L. triflora Desr. in Lamk 1792 (nom. illeg.)] R77

*Type species of generic name indicated


[AS02] Albert, V. A., & L. Struwe. 2002. Gentianaceae in context. In: Struwe, L., & V. A. Albert (eds) Gentianaceae: Systematics and Natural History pp. 1–20. Cambridge University Press: Cambridge.

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

[B00] Braby, M. F. 2000. Butterflies of Australia: their identification, biology and distribution vol. 1. CSIRO Publishing: Collingwood (Victoria).

[BJ99] Bremer, B., R. K. Jansen, B. Oxelman, M. Backlund, H. Lantz & K.-J. Kim. 1999. More characters or more taxa for a robust phylogeny—case study from the coffee family (Rubiaceae). Systematic Biology 48 (3): 413–435.

[BM00] Bremer, B., & J.-F. Manen. 2000. Phylogeny and classification of the subfamily Rubioideae (Rubiaceae). Plant Systematics and Evolution 225: 43–72.

[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.

[C95] Conn, B. J. 1995. Loganiaceae. In: Conn, B. J. (ed.) Handbooks of the Flora of Papua New Guinea vol. 3 pp. 132–188. Melbourne University Press: Carlton (Australia).

[FT93] Fensome, R. A., F. J. R. Taylor, G. Norris, W. A. S. Sarjeant, D. I. Wharton & G. L. Williams. 1993. A classification of living and fossil dinoflagellates. Micropaleontology Special Publication 7: i–viii, 1–351.

[G04] Gibson, N. 2004. 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.

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

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

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

[KJ05] Katewa, S. S., A. Jain, B. L. Chaudhary & P. Galav. 2005. Some unreported medicinal uses of plants from the tribal area of Southern Rajasthan. Bulletin of the Botanical Survey of India 47: 121–130.

[KC01] Kirk, P. M., P. F. Cannon, J. C. David & J. A. Stalpers. 2001. Ainsworth & Bisby's Dictionary of the Fungi 9th ed. CAB International: Wallingford (UK).

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

[M09] Mayr, G. 2009. Paleogene Fossil Birds. Springer.

[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.

[P03] Paul, T. K. 2003. Botanical observations on the Purulia pumped storage hydropower project area, Bagmundi Hills, Purulia district, West Bengal. Bulletin of the Botanical Survey of India 45: 121–142.

[R77] Raven, P. H. 1977. Onagraceae. Flora Malesiana, Series I—Spermatophyta, Flowering Plants 8 (2): 98–113.

[RB01] Razafimandimbison, S. G., & B. Bremer. 2001. Tribal delimitation of Naucleeae (Cinchonoideae, Rubiaceae): inference from molecular and morphological data. Systematics and Geography of Plants 71: 515–538.

[SK02] Struwe, L., J. W. Kadereit, J. Klackenberg, S. Nilsson, M. Thiv, K. B. von Hagen & V. A. Albert. 2002. Systematics, character evolution, and biogeography of Gentianaceae, including a new tribal and subtribal classification. In: Struwe, L., & V. A. Albert (eds) Gentianaceae: Systematics and Natural History pp. 21–309. Cambridge University Press: Cambridge.

[VT13] Vigilante, T., J. Toohey, A. Gorring, V. Blundell, T. Saunders, S. Mangolamara, K. George, J. Oobagooma, M. Waina, K. Morgan & K. Doohan. 2013. Island country: aboriginal connections, values and knowledge of the Western Australian Kimberley islands in the context of an island biological survey. Records of the Western Australian Museum Supplement 81: 145–182.

[WB-P93] Walter, D. E., & V. M. Behan-Pelletier. 1993. Systematics and ecology of Adhaesozetes polyphyllos sp.nov. (Acari: Oribatida: Licneremaeoidea), a leaf-inhabiting mite from Australian rainforests. Canadian Journal of Zoology 71: 1024–1040.

[WP99] Walter, D. E., & H. C. Proctor. 1999. Mites: Ecology, Evolution and Behaviour. CABI Publishing: Wallingford (UK).

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


Noni Morinda citrifolia, copyright Wilfredor.

Belongs within: Rubioideae.

The Morindeae are a group of rubiaceous plants bearing flowers in head inflorescences, and fleshy drupes with one or two ovules per carpel.

    |--Morindinae BM00
    |    |  i. s.: Gynochthodes coriacea BM00
    |    |         Pogonolobus BM00
    |    |--Appunia guatemalensis BM00
    |    `--+--Coelospermum balansanum BM00
    |       `--Morinda BM00
    |            |--M. canthoides LK14
    |            |--M. citrifolia RB01
    |            |--M. elliptica P88
    |            `--M. jasminoides LK14
    `--+--+--Mitchella repens BM00
       |  `--Damnacanthus indicus BM00
       `--Prismatomerinae BM00
            |--Renellia [incl. Didymoecium] BM00
            |--Motleyia BM00
            `--Prismatomeris BM00
                 |--P. beccarianum BM00
                 `--P. labordei BM00

*Type species of generic name indicated


[BM00] Bremer, B., & J.-F. Manen. 2000. Phylogeny and classification of the subfamily Rubioideae (Rubiaceae). Plant Systematics and Evolution 225: 43–72.

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

[P88] Polunin, I. 1988. Plants and Flowers of Malaysia. Times Editions: Singapore.

[RB01] Razafimandimbison, S. G., & B. Bremer. 2001. Tribal delimitation of Naucleeae (Cinchonoideae, Rubiaceae): inference from molecular and morphological data. Systematics and Geography of Plants 71: 515–538.


Faramea cf. multiflora, copyright Scott Zona.

Belongs within: Rubioideae.

The Coussareeae are a small but morphologically diverse group of flowering plants found in the Americas, many of which have tetramerous flowers, flattened berry-like or thin-walled capsular fruits, and flattened seeds (Bremer & Manen 2000).

Characters (from Bremer & Manen 2000): Herbs (creeping in Coccocypselum), subshrubs, shrubs, or small trees. Stipules entire, cleft, as a ridge with appendages, or deeply bilobed. Flowers homo- or heterostylous, often tetramerous, white, blue, or bright yellow (Cruckshanksia, Oreopolus). Ovary 1–2-1ocular, with 1–2 or many (Coccocypselum) ovules per locule. Fruits often flattened (not Coccocypselum), fleshy, white (Coussarea) or often blue (Faramea, Coccocypselum) berries, schizocarps (Declieuxia), or thin-walled capsules (Cruckshanksia, Oreopolus), with 1–2 or many seeds, often flattened. Pollen 3-colporate or 2–4-porate. Chromosome basic number x = 10, 11? with 2- or 4-ploidy level.

<==Coussareeae [Coccocypseleae, Cruckshanksieae]
    |  i. s.: Cruckshanksia hymenodon BM00
    |         Oreopolus glacialis BM00
    |         Declieuxia fruticosa BM00
    |         Heterophyllaea BM00
    |         Schizocolea BM00
    |--Coccocypselum hirsutum BM00
    `--+--Coussarea BM00
       |    |--C. contracta BM00
       |    `--C. macrophylla BM00
       `--Faramea BM00
            |--F. multiflora BM00
            `--F. porophylla BM00

*Type species of generic name indicated


[BM00] Bremer, B., & J.-F. Manen. 2000. Phylogeny and classification of the subfamily Rubioideae (Rubiaceae). Plant Systematics and Evolution 225: 43–72.


Mountain saucer-flower Crusea diversifolia, copyright Sue Carnahan.

Belongs within: Spermacoceae.

Crusea, saucer flowers, is a genus of small annual plants found in southern North America.

    |--C. coronata F79
    `--+--+--C. coccinea F79
       |  |    |--C. c. var. breviloba F79
       |  |    `--+--C. c. var. coccinea F79
       |  |       `--C. c. var. chiriquensis F79
       |  `--C. megalocarpa F79
       `--+--C. diversifolia F79
          `--+--C. lucida F79
             `--+--C. parvifolia F79
                |--C. psyllioides F79
                `--+--C. calcicola F79
                   |--C. setosa F79
                   `--+--C. wrightii var. wrightii F79
                      `--+--C. longiflora F79
                         `--+--C. wrightii var. angustifolia F79
                            `--+--C. calocephala F79
                               `--C. hispida F79
                                    |--C. h. var. hispida F79
                                    `--C. h. var. grandiflora F79

*Type species of generic name indicated


[F79] Farris, J. S. 1979. Review: Cladistics and Plant Systematics. Systematic Zoology 28 (3): 400–411.


Epicrus canestrinii, copyright Matthew Shepherd.

Belongs within: Gamasina.

The Epicriidae are a group of mites known from the Northern Hemisphere with species found in habitats such as moss, leaf litter, decaying wood or animal nests (Lindquist et al. 2009).

Characters (from Lindquist et al. 2009): Female and male dorsal shield entire, usually expanded laterally and consolidated with peritrematic shields, which in turn are expanded posteriorly onto the ventrolateral surface behind coxae IV, ornamented with 11–19 pairs of dorsal glands and a network of large bi-, tri-, or qundrifurcate tubercles. Stigmata prominent, typically on protuberances, peritremes vestigial or lacking. Tritosternum reduced to stubby base and abbreviated laciniae. Sternal setae I on paired jugular plates, poroids may be absent. Female sternal shield with 2–3 pairs of setae, apparently without poroids; 4th sternal setae usually free on soft integument or rarely inserted on metasternal plates or on posterior edge of sternal shield; epigynal shield with two pairs (occasionally one pair) of setae and well separated posteriorly from an anal or small ventrianal shield. Males with sternitigenital shield separated from ventral and anal shields or from ventrianal or anal shield (the latter sometimes integrated with posterior margin of dorsal shield). Posterior margins of coxae IV with single or multiple inguinal glands. Leg I elongated, without tarsal claws and with two to several setae that are more or less elongated and minutely clubbed apically. Peripodomeric suture of tarsi II–IV usually with a dorsal intercalary sclerite bearing two setae. Femur I with thirteen setae, including four ventral setae and five dorsal setae; tibia I with four ventral setae, six dorsal setae, and 2 anterolateral setae; tibia III with nine setae (2 1/1, 2/1 2); genu III with ten setae, including two ventral setae. Palptarsal claw three-tined (occasionally two-tined). Corniculi small, slender, much shorter than internal malae. Female tocospermous. Male with genital aperture located between coxae III and without spurs or modified spinelike setae on legs II–IV; male chelicerae not obviously elaborated for sperm transfer. Larva lacking pygidial shield; protonymphal pygidial shield with eight pairs of setae, setae S4S5 on shield; dorsal shield of deutonymph entire.

<==Epicriidae [Epicrioidea] LKW09
    |--Neoepicrius LKW09
    |--Adenoepicrius LKW09
    |--Berlesiana Turk 1943 ET79
    |    |--*B. cirrata (Berlese 1916) [=Epicrius cirratus] ET79
    |    `--B. denticulata ET79
    `--Epicrius Canestrini & Fanzago 1878 (see below for synonymy) FH93
         |--E. mollis (Kramer 1876) [=Gamasus mollis; incl. *E. geometricus Canestrini & Fanzago 1887] ET79
         |--E. canestrinii S61
         |--E. menzeli Schweizer 1922 S61
         |--E. spinituberculatus ET79
         `--E. washingtonianus Berlese 1916 FH93

Epicrius Canestrini & Fanzago 1878 [incl. Diepicrius Berlese 1916, Epicriella Willmann 1953, Eugamasus Absolon 1899 non Berlese 1892, Parasejus Trägårdh 1910] FH93

*Type species of generic name indicated


[ET79] Evans, G. O., & W. M. Till. 1979. Mesostigmatic mites of Britain and Ireland (Chelicerata: Acari-Parasitiformes). An introduction to their external morphology and classification. Transactions of the Zoological Society of London 35: 139–270.

[FH93] Farrier, M. H., & M. K. Hennessey. 1993. Soil-inhabiting and free-living Mesostigmata (Acari-Parasitiformes) from North America: an annotated checklist with bibliography and index. North Carolina Agricultural Research Service, North Carolina State University, Technical Bulletin 302: i–xvi, 1–408.

[LKW09] Lindquist, E. E., G. W. Krantz & D. E. Walter. 2009. Order Mesostigmata. In: Krantz, G. W., & D. E. Walter (eds) A Manual of Acarology 3rd ed. pp. 124–232. Texas Tech University Press.

[S61] Schweizer, J. 1961. Die Landmilben der Schweiz (Mittelland, Jura und Alpen): Parasitiformes Reuter, mit 246 Arten und Varietäten und 268 meist kombinierten Originalzeichnungen. Denkschriften der Schweizerischen Naturforschenden Gesellschaft [Mémoires de la Société Helvétique des Sciences Naturelles] 84: i–vii, 1–207.