Kimulidae

Minuella dimorpha, from Pérez González & Kury (2007).
Belongs within: Laniatores.

The Kimulidae is a small family of short-legged harvestmen found in northern South America.

Characters (from Pérez González & Kury 2007): Dorsal scutum bell-shaped with laterals of carapace convex; opisthosomal scutum widest at groove II and slightly constricted at area III or IV (Tegipiolus without any constriction); opisthosomal scutum outline in lateral view high, but somewhat flattened, not rounded convex. Ocularium prominent, granular, armed with a medial spiniform apophysis erect or curved or sinuous or inclined anteriorly. Mesotergum with four areas, area I longer than the others; mesotergal areas typically densely granular but unarmed; in a few species area I possesses a pair or transverse row of pointed tubercles. Lateral margin of scutum may have eularged spiniform tubercles, larger at area II. Free tergites with pointed corners or unpaired median apophyses. Free Sternites may bear varied armature, rows of spines or unpaired spines. Chelicerae with well-marked bulla, without remarkable armature. Pedipalps with setiferous tubercles never strongly developed; femur convex dorsally wlth ventral and dorsal scattered setiferous tubercles, group of two basal setiferous tubercles with basalmost tubercle always much reduced and always with mesal subdistal setiferous tubercle; patella with one mesal subdistal setiferous tubercle; tibia a little elongate, usually longer than tarsus, marginated with lateroectal and lateromesal rows of two to seven setiferous tubercles; tarsus with lateroectal and lateromesal rows of three to four setiferous tubercles. Legs III-IV without tarsal process and scopula; male coxa IV well developed, visible under scutum in dorsal view, roughly armed with heavy granules or acute tubercles; trochanter IV with characteristic ventral spine; femur IV very incrassate, with ventral row of strong spiniform apophyses; tibia and metatarsus IV roughly tuberculate with very enlarged and globose ventral tubercles at apices; tarsal formula 4(2):6-13(2):5:5-6 (except in Fudeci, 3:5:5:5). Penis with truncus cylindrical without well-defined ventral plate; pars distalis well-differentiated from pars basalis by one sulcus or the border between both easily recognisable by the upper limit of the pars basalis striate area; pars distalis laterally armed with three or four strong spatulate spines, in some species lanceolate, in others rounded, and ventrally with four small acute setae; Pars distalis with lamina ventralis surrounding capsula interna (conductors + stylus), apical extremes of the lamina ventralis enlarged, fingerlike, touching one another dorsally, the apical region of pars distalis commonly with two folds that can be ventrally entire or divided; two rigid conductors that can be lamelar or greatly developed and enlarged.

Kimulidae [Minuidae, Minuinae]
    |--Acanthominua Sørensen 1932 K03
    |    `--*A. tricarinata Sørensen 1932 K03
    |--Euminua Sørensen 1932 K03
    |    `--*E. brevitarsa Sørensen 1932 K03
    |--Euminuoides Mello-Leitão 1935 K03
    |    `--*E. longitarsa (Sørensen 1932) [=Euminua longitarsa, E. longitarsis (l. c.)] K03
    |--Fudeci González-Sponga 1997 PGK07, K03
    |    `--*F. curvifemur González-Sponga 1997 K03
    |--Metakimula Avram 1973 PGK07, K03
    |    `--*M. botosaneanui (Avram 1973) [=Kimula (Metakimula) botosaneanui] K03
    |--Tegipiolus Roewer 1949 PGK07, K03
    |    `--*T. pachypus Roewer 1949 K03
    |--Microminua Sørensen 1932 K03
    |    |--*M. parvula Sørensen 1932 K03
    |    `--M. soerenseni Soares & Soares 1954 K03
    |--Pseudominua Mello-Leitão 1933 K03
    |    |--*P. convolvulus (Sørensen 1932) [=Euminua convolvulus] K03
    |    `--P. peruviana Roewer 1963 K03
    |--Kimula Goodnight & Goodnight 1942 PGK07
    |    |--*K. elongata Goodnight & Goodnight 1942 K03
    |    |--K. banksi Šilhavý 1969 K03
    |    |--K. cokendolpheri Pérez González & Armas 2000 K03
    |    |--K. goodnightiorum Šilhavý 1969 K03
    |    |--K. levii Šilhavý 1969 K03
    |    |--K. tuberculata Goodnight & Goodnight 1943 K03
    |    `--K. turquinensis Šilhavý 1969 K03
    `--Minuella Roewer 1949 PGK07 [incl. Minua Sørensen 1932 (nom. inv.) PGK07, K03]
         |--*M. dimorpha (Sørensen 1932) [=Minua dimorpha; incl. Minua insolens Sørensen 1932] K03
         |--M. scabra (Sørensen 1932) [=*Minua scabra] K03
         |--M. barloventensis González-Sponga 1987 [=Minua barloventensis] K03
         |--M. choroniensis González-Sponga 1987 [=Minua choroniensis] K03
         |--M. crassa González-Sponga 1987 [=Minua crassa] K03
         |--M. denticulata González-Sponga 1987 [=Minua denticulata] K03
         |--M. elias (Sørensen 1932) [=Minua elias] K03
         |--M. guatopensis González-Sponga 1987 [=Minua guatopensis] K03
         |--M. momoyana González-Sponga 1998 [=Minua momoyana] K03
         |--M. montis González-Sponga 1987 [=Minua montis] K03
         |--M. nebulae González-Sponga 1987 [=Minua nebulae] K03
         |--M. parva González-Sponga 1987 [=Minua parva] K03
         |--M. pinturelensis González-Sponga 1987 [=Minua pinturelensis] K03
         |--M. punctiacuta González-Sponga 1987 [=Minua punctiacuta] K03
         `--M. venefica González-Sponga 1987 [=Minua venefica] K03

*Type species of generic name indicated

REFERENCES

[K03] Kury, A. B. 2003. Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Ibérica de Aracnología, volumen especial monográfico 1: 1-337.

[PGK07] Pérez González, A., & A. B. Kury. 2007. Kimulidae Pérez González, Kury, and Alonso-Zarazaga, new name. In Harvestmen: The Biology of Opiliones (R. Pinto-da-Rocha, G. Machado & G. Giribet, eds) pp. 207-209. Harvard University Press: Cambridge (Massachusetts).

Escadabiidae

Undescribed species of Baculigerus, from Kury & Pérez González (2007).
Belongs within: Laniatores.

The Escadabiidae is a small family of short-legged harvestmen that has so far only been recorded from coastal regions and inland caves of Brazil.

Characters (from Kury & Pérez González 2007): Dorsal scutum 2.5–3.5 mm long. Dorsal scutum campaniform, with sides straight, hourglass-shaped in a few species. Ocularium small, with a few granules and sometimes a median spine. Mesotergum divided into four areas by straight transverse grooves, without relevant armature, area I longer than others. Sternites smooth and unarmed or with huge lateral projections. Chelicerae weak and not sexually dimorphic; basichelicerite short with well-marked bulla. Pedipalps more or less as long as dorsal scutum, without special modifications; femur with two or three ventral setiferous tubercles, patella with one mesal setiferous tubercle; tibia and tarsus each with a row of a few ventromesal and ventroectal setiferous tubercles. All legs short and granulous; coxa IV of male coarsely granulate with dorsoapical apophysis made up of two subequal branches, lateral border parallel to main axis of body; femora I–II straight, IV curved subbasally, strongly incrassate in males of Escadabius; tibiae I–II with differentiated porose distal area that may develop into a deep notch and/or huge apophysis. Penis with pair of rigid conductors and reversible capsula interna; ventral plate not defined; pars distalis well separated from pars basalis by constriction (absent in Escadabius), with ventral lamina apicalis that may be armed with small lateral spines; pars distalis with ventral keel-shaped protuberance; pair of well-developed rigid conductors that do not sink deeply into glans socket (vestigial in Escadabius); stylus very large and apically surrounded by well-developed hornlike parastylar collar.

Escadabiidae
    |--Baculigerus Soares 1979 KPG07, K03
    |    `--*B. littoris Soares 1979 K03
    |--Jim Soares 1979 KPG07, K03
    |    `--*J. benignus Soares 1979 K03
    |--Recifesius Soares 1978 KPG07, K03
    |    `--*R. pernambucanus Soares 1978 K03
    |--Spaeleoleptes Soares 1966 KPG07, K03
    |    `--*S. spaeleus Soares 1966 K03
    `--Escadabius Roewer 1949 KPG07, K03
         |--*E. ventricalcaratus Roewer 1949 K03
         |--E. schubarti Roewer 1949 K03
         `--E. spinicoxa Roewer 1949 K03

*Type species of generic name indicated

REFERENCES

[K03] Kury, A. B. 2003. Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Ibérica de Aracnología, volumen especial monográfico 1: 1-337.

[KPG07] Kury, A. B., & A. Pérez González. 2007. Escadabiidae Kury and Pérez in Kury, 2003. In Harvestmen: The Biology of Opiliones (R. Pinto-da-Rocha, G. Machado & G. Giribet, eds) pp. 191-194. Harvard University Press: Cambridge (Massachusetts).

Pinaceae

Norway spruce Picea abies, photographed by MPF.

Belongs within: Coniferophyta.
Contains: Abies, Pinus subg. Pinus, Pinus subg. Strobus.

The Pinaceae include the pines (Pinus), spruces (Picea), cedars (Cedrus) and related taxa. This is a primarily Northern Hemisphere family, with some species being the dominant tree species within their range. Pinus species differ from other members of the family in having cones that are biennial or triennial, while other genera grow new cones each year. Members of the genus Picea, the spruces, have whorled branches with needles attached singly in a spiral pattern. Larix, larches, are deciduous, dropping their needles over winter.

Characters (from J. W. Thieret): Trees (occasionally shrubs), evergreen (annually deciduous in Larix), resinous and aromatic, monoecious. Bark smooth to scaly or furrowed. Lateral branches well developed and similar to leading (long) shoots or reduced to well-defined short (spur) shoots (Pinus, Larix); twigs terete, sometimes clothed by persistent primary leaves or leaf bases; longest internodes less than 1cm; buds conspicuous. Roots fibrous to woody, unspecialized. Leaves (needles) simple, shed singly (except whole fascicles shed in Pinus), alternate and spirally arranged but sometimes proximally twisted so as to appear 1- or 2-ranked, or fascicled, linear to needlelike, sessile to short-petiolate; foliage leaves either borne singly (spirally) on long shoots or in tufts (fascicles) on short shoots; juvenile leaves (when present) borne on long shoots, scalelike; resin canals present. Pollen cones maturing and shed annually, solitary or clustered, axillary, ovoid to ellipsoid or cylindric; sporophylls overlapping, bearing 2 abaxial microsporangia (pollen sacs); pollen spheric, 2-winged, less commonly with wings reduced to frill (in Tsuga sect. Tsuga), or not winged (in Larix and Pseudotsuga). Seed cones maturing and shed in 1-3 seasons or long-persistent, sometimes serotinous (not opening upon maturity but much later: Pinus), compound, axillary, solitary or grouped; scales overlapping, free from subtending included or exserted bracts for most of length, spirally arranged, strongly flattened, at maturity relatively thin to strongly thickened and woody (in Pinus), with 2 inverted, adaxial ovules. Seeds 2 per scale, elongate terminal wing partially decurrent on seed body (wing short or absent in some species of Pinus); aril lacking; cotyledons 2-12[-18].

Pinaceae [Abietaceae, Abietinae, Pinales]
    |  i. s.: Pinites succinifer Göppert 1836 E01 [=Pinus succinifera P02]
    |         Nothotsuga CD07
    |         Compsostrobus neotericus MM12
    |         Schizolepidopsis HS17
    |--Cedrus EH06
    |    |--C. atlantica C55
    |    |--C. brevifolia D30
    |    |--C. deodara EH06
    |    `--C. libani C55
    `--+--+--+--Abies EH06
       |  |  `--Keteleeria EH06
       |  |       |--K. davidiana EH06
       |  |       `--K. evelyniana MM12
       |  `--+--Pseudolarix amabilis EH06
       |     `--Tsuga EH06
       |          |--T. canadensis SS96
       |          |--T. chinensis CP02
       |          |--T. dumosa SN88
       |          |--T. heterophylla C90
       |          |--T. mertensiana EH06
       |          `--T. sieboldii [incl. Abies tsuga] LO98
       `--+--+--Pseudotsuga EH06
          |  |    |--P. douglasii B88
          |  |    |--P. macrocarpa B14
          |  |    |--P. menziesii EH06
          |  |    `--P. taxifolia SW03
          |  `--Larix EH06
          |       |--L. decidua EH06
          |       |--L. gmelinii L02
          |       |    |--L. g. var. gmelinii F92
          |       |    `--L. g. var. japanica F92
          |       |--L. groenlandii TH03
          |       |--L. himalaica SN88
          |       |--L. kaemferi F92
          |       |--L. koreana F92
          |       |--L. laricina TH03
          |       |--L. leptolepis F92
          |       |--L. occidentalis MM12
          |       |--L. olbensis F92
          |       |--L. sibirica F92
          |       `--L. subczewii F92
          `--+--+--Cathaya argyrophylla EH06
             |  `--Picea EH06
             |       |--P. abies EH06
             |       |--P. alcoquiana [incl. Abies menziessi] LO98
             |       |--P. breweriana H93
             |       |--P. canadensis R38
             |       |--P. contorta F56
             |       |    |--P. c. var. contorta F56
             |       |    `--P. c. var. latifolia F56
             |       |--P. engelmannii H93
             |       |--P. excelsa M47
             |       |--P. glauca BFD01
             |       |--P. glehnii M03
             |       |--P. jezoensis JKW03
             |       |--P. mariana BFD01
             |       |--P. morrisonicola MM12
             |       |--P. orientalis U01
             |       |--P. rubens SR03
             |       |--P. sitchensis MM12
             |       `--P. smithiana MM12
             `--+--Pityostrobus bernissartensis MM12
                `--Pinus EH06
                     |  i. s.: P. abies S00
                     |         P. ayacahuite CG05
                     |         P. cooperi CG05
                     |         P. fenzeliana L02
                     |         P. kochiana CK05
                     |         P. larix S00
                     |         P. luchuensis SM03
                     |         P. maritima C55
                     |         P. montana SR03
                     |         P. occidentalis J87
                     |         P. palaeostrobus S89
                     |         P. pallasiana CK05
                     |         P. pringlei CG05
                     |         P. quadrifolia H93
                     |         P. rudis CG05
                     |         P. torreyana [incl. P. torreyana ssp. insularis] H93
                     |         P. uncinata C55
                     |--P. subg. Pinus EH06
                     `--P. subg. Strobus EH06

*Type species of generic name indicated

REFERENCES

[B88] Bouček, Z. 1988. Australasian Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International: Wallingford (UK).

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

[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.

[C55] Candolle, A. de. 1855. Géographie Botanique Raisonée: Ou exposition des faits principaux et des lois concernant la distribution géographique des plantes de l’époque actuelle vol. 2. Librairie de Victor Masson: Paris.

[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.

[CK05] Chemeris, A. N., & N. M. Kovblyuk. 2005. A contribution to the knowledge of the harvestman fauna of the Crimea (Arachnida: Opiliones). Arthropoda Selecta 14 (4): 305–328.

[CP02] Chung, S.-W., & C.-I. Peng. 2002. Senecio kuanshanensis (Asteraceae), a new species from southern Taiwan. Botanical Bulletin of Academia Sinica 43: 155–159.

[CG05] Cognato, A. I., N. E. Gillette, R. C. Bolaños & F. A. H. Sperling. 2005. Mitochondrial phylogeny of pine cone beetles (Scolytinae, Conophthorus) and their affiliation with geographic area and host. Molecular Phylogenetics and Evolution 36: 494–508.

[C90] Crawford, R. L. 1990. Discovery of Caviphantes saxetorum in North America; status of Scironis tarsalis (Araneida, Linyphiidae). Journal of Arachnology 18: 235–236.

[D30] Druce, G. C. 1930. Account of a botanical tour in Cyprus. Proceedings of the Linnean Society of London 141: 50–52.

[EH06] Eckert, A. J., & B. D. Hall. 2006. Phylogeny, historical biogeography, and patterns of diversification for Pinus (Pinaceae): phylogenetic tests of fossil-based hypotheses. Molecular Phylogenetics and Evolution 40 (1): 166–182.

[E01] Engel, M. S. 2001. A monograph of the Baltic amber bees and evolution of the Apoidea (Hymenoptera). Bulletin of the American Museum of Natural History 259: 1–192.

[F92] Fan Z. 1992. Key to the Common Flies of China 2^nd ed. Science Press: Beijing.

[F56] Foote, R. H. 1956. Gall midges associated with cones of western forest trees (Diptera: Itonididae). Journal of the Washington Academy of Sciences 46 (2): 48–57.

[HS17] Herrera, F., G. Shi, N. Ichinnorov, M. Takahashi, E. V. Bugdaeva, P. S. Herendeen & P. R. Crane. 2017. The presumed ginkgophyte Umaltolepis has seed-bearing structures resembling those of Peltaspermales and Umkomasiales. Proceedings of the National Academy of Sciences of the USA: E2385–E2391.

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

[JKW03] Jacobs, K., T. Kirisits & M. J. Wingfield. 2003. Taxonomic re-evaluation of three related species of Graphium, based on morphology, ecology and phylogeny. Mycologia 95 (4): 714–727.

[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.

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

[L02] Li F. 2002. Psocoptera of China vol. 2. Science Press: Beijing.

[MM12] Mao, K., R. I. Milne, L. Zhang, Y. Peng, J. Liu, P. Thomas, R. R. Mill & S. S. Renner. 2012. Distribution of living Cupressaceae reflects the breakup of Pangea. Proceedings of the National Academy of Sciences of the USA 109 (20): 7793–7798.

[M47] Mayr, E. 1947. Ecological factors in speciation. Evolution 1: 263–288.

[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.

[P02] Ponomarenko, A. G. 2002. Superorder Scarabaeidea Laicharting, 1781. Order Coleoptera Linné, 1758. The beetles. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 164–176. Kluwer Academic Publishers: Dordrecht.

[R38] Ross, H. H. 1938. The Nearctic species of Pikonema, a genus of spruce sawflies (Hymenoptera, Tenthredinidae). Proceedings of the Entomological Society of Washington 40 (1): 17–20.

[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.

[S00] Saussure, C. de. 1800. De l'influence du sol sur quelques parties constituantes des végétaux. Bulletin des Sciences, par la Societé Philomathique de Paris 2 (40): 124–125.

[SS96] Simberloff, D., & P. Stilling. 1996. How risky is biological control? Ecology 77 (7): 1965–1974.

[SR03] Skovgaard, K., S. Rosendahl, K. O’Donnell & H. I. Nirenberg. 2003. Fusarium commune is a new species identified by morphological and molecular phylogenetic data. Mycologia 95 (4): 630–636.

[SW03] Song, D., & Q. Wang. 2003. Systematics of the longicorn beetle genus Coptomma Newman (Coleoptera: Cerambycidae: Cerambycinae). Invertebrate Systematics 17: 429–447.

[S89] Squinabol, S. 1889. Res Ligusticae. VII.—Cenno preliminare sulla flora fossile di Santa Giustina. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 73–76.

[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.

[TH03] Tedford, R. H., & C. R. Harington. 2003. An Arctic mammal fauna from the Early Pliocene of North America. Nature 425: 388–390.

[U01] Urhan, R. 2001. Two new species of the genus Zercon Koch from Turkey (Acari: Gamasida: Zerconidae). Genus 12 (4): 589–597.

Last updated: 2 December 2018.

Erdtmanithecales

Reconstruction of microsporangiate organs of Eucommiitheca hirsuta, from Friis & Pedersen (1996).
Belongs within: Spermatophyta.

The Erdtmanithecales are a group of Cretaceous plants known from reproductive structures and seeds associated with Eucommiidites pollen; vegetative structures have not yet been identified for the group.

Characters (from Friis & Pedersen 1996): Male reproductive organs of closely spaced microsporangiate units; microsporangiate units stalked and peltate, bearing numerous sporangia in a radial arrangement around stalk; dehiscence of sporangia by longitudinal slits. Pollen grains of the Eucommiidites type dispersed singly; single grains elliptical in equatorial outline, bisymmetrical with two planes of symmetry through the polar areas and with a distal main colpus; main colpus typically expanded at the ends and flanked by two colpi or a single ring colpus in a proximal to almost equatorial position; pollen wall ultrastructure with granular infratectal structure and thick laminated endexine. Seeds small, orthotropous; megaspore membrane well developed; micropylar tube long and narrow, formed from inner envelope (inner integument).

<==Erdtmanithecales
    |--Erdtmanispermum balticum FC07
    |--Erdtmanitheca texensis FC07
    |--Eucommiitheca hirsuta FC07
    |--Bayeritheca hughesii FC07
    |--Eucommiidites FC07
    |    |--E. minor Groot & Penny 1960 I02
    |    `--E. troedsonii Erdtmann 1948 I02
    `--Spermatites FC07
         |--S. pattensis FC07
         `--S. patuxensis FC07

*Type species of generic name indicated

REFERENCES

Friis, E. M., & K. R. Pedersen. 1996. Eucommiitheca hirsuta, a new pollen organ with Eucommiidites pollen from the Early Cretaceous of Portugal. Grana 35 (2): 104-112.

[FC07] Friis, E. M., P. R. Crane, K. R. Pedersen, S. Bengtson, P. C. J. Donoghue, G. W. Grimm & M. Stampanoni. 2007. Phase-contrast X-ray microtomography links Cretaceous seed with Gnetales and Bennettitales. Nature 450: 549-552.

[I02] Ibrahim, M. I. A. 2002. Late Albian-Middle Cenomanian palynofacies and palynostratigraphy, Abu Gharadig-5 well, Western Desert, Egypt. Cretaceous Research 23: 775-788.

Dioscoreaceae

Dioscorea madecassa, copyright RBG Kew.

Belongs within: Dioscoreales.

The Dioscoreaceae is a pantropical family mostly composed of tuberous vines, some species of which (in the genus Dioscorea) are grown as food plants. Black bryony Tamus communis is a widespread plant of the forest understory in the Mediterranean and adjoining regions.

Characters (from L. Raz):Vines (sometimes herbs), geophytes perennating from fleshy rhizomes or tubers. Stems renewed annually or occasionally persisting for more than a single growing season, twining-climbing, procumbent, or seldom erect; vascular bundles commonly arranged in 2 concentric circles. Leaves alternate, opposite, or whorled at basal nodes; blades simple or occasionally digitately compound, typically cordate, reticulate-veined; principal veins arcuate, ascending from base to apex; margins entire or sometimes palmately lobed; stipules absent, rarely stipulate, never with tendrils; petioles typically with distinct pulvinus at each end. Inflorescences axillary, solitary or in fascicles of 2–7, branched and paniculate, or unbranched and spicate or racemose, ultimate inflorescence unit cymose, sessile or pedunculate, often reduced to 1 flower. Flowers unisexual, staminate and pistillate flowers on different plants, rarely staminate and pistillate flowers on same plant or bisexual; perianth epigynous, rotate, campanulate, or funnelform; tepals 6, petaloid, in 2 similar whorls of 3, connate at least basally; stamens 6, in 2 whorls of 3, inner whorl sometimes sterile or absent; filaments distinct (sometimes connate basally or completely into tube), inserted at base of perianth; anthers 2-locular, tetrasporangiate, dehiscing by longitudinal slits, thecae distinct or sometimes connate, rarely divided by a long-appendiculate connective; pistils 3; ovary (1–)3-locular, ovules anatropous, 2(–many) per locule, placentation axile or parietal; styles 3, ± distinct. Fruits capsular, rarely baccate or samarate, dehiscence loculicidal. Seeds flattened or globose, winged or unwinged, embryo small, endosperm copious, starchless.

Dioscoreaceae
    |--Epipetrum YY22
    |--Rajania ovata J87
    |--Testudinaria YY22
    |--Borderea YY22
    |--Higinbothamia YY22
    |--Tamus communis DS04
    |    |--T. c. ssp. communis PT98
    |    `--T. c. ssp. cretica PT98
    `--Dioscorea WA08
         |  i. s.: D. aculeata C55
         |         D. alata C55
         |         D. belophylla HU08
         |         D. bulbifera C55
         |         D. decaryana WA08
         |         D. deltoides C55
         |         D. elephantipes MS10
         |         D. glabra P03
         |         D. hamiltonii SR07
         |         D. hispida P03
         |         D. hombuka WA08
         |         D. kimiae Wilkin in Wilkin, Andrianantenaina et al. 2008 WA08
         |         D. mayottensis WA08
         |         D. nummularia H03
         |         D. pentaphylla C55
         |         D. polygonoides DS04
         |         D. retusa DS04
         |         D. sambiranensis WA08
         |         D. sterilis WA08
         |         D. transversa VT13
         |         D. villosa EBS98
         |--D. sect. Campanuliflorae WA08
         |    |--D. acuminata Baker 1885 [incl. D. bararum Perrier 1928] WA08
         |    |--D. maciba Jum. & Perrier 1910 WA08
         |    `--D. madecassa Perrier 1928 (see below for synonymy) WA08
         `--D. sect. Seriflorae WA08
              |--D. seriflora Jum. & Perrier 1910 (see below for synonymy) WA08
              `--D. tsaratananensis Perrier 1928 WA08

Dioscorea madecassa Perrier 1928 [incl. D. karatana Wilkin in Wilkin et al. 2000, D. knuthii Perrier 1928] WA08

Dioscorea seriflora Jum. & Perrier 1910 [incl. D. ovifotsy Perrier 1928, D. porosa Boivin ms, D. tanalarum Perrier 1928] WA08

*Type species of generic name indicated

REFERENCES

[C55] Candolle, A. de. 1855. Géographie Botanique Raisonée: Ou exposition des faits principaux et des lois concernant la distribution géographique des plantes de l’époque actuelle vol. 2. Librairie de Victor Masson: Paris.

[DS04] Davis, J. I., D. W. Stevenson, G. Petersen, O. Seberg, L. M. Campbell, J. V. Freudenstein, D. H. Goldman, C. R. Hardy, F. A. Michelangeli, M. P. Simmons, C. D. Specht, F. Vergara-Silva & M. Gandolfo. 2004. A phylogeny of the monocots, as inferred from rbcL and atpA sequence variation, and a comparison of methods for calculating jackknife and bootstrap values. Systematic Botany 29 (3): 467–510.

[EBS98] Elliott, K. J., L. R. Boring & W. T. Swank. 1998. Changes in vegetation structure and diversity after grass-to-forest succession in a southern Appalachian watershed. American Midland Naturalist 140: 219–232.

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

[HU08] Hore, U., & V. P. Uniyal. 2008. Diversity and composition of spider assemblages in five vegetation types of the Terai Conservation Area, India. Journal of Arachnology 36 (2): 251–258.

[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.

[MS10] Moore, M. J., P. S. Soltis, C. D. Bell, J. G. Burleigh & D. E. Soltis. 2010. Phylogenetic analysis of 83 plastid genes further resolves the early diversification of eudicots. Proceedings of the National Academy of Sciences of the USA 107 (10): 4623–4628.

[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.

[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 (1–4): 121–142.

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

[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.

[WA08] Wilkin, P., W. P. Andrianantenaina, V. Jeannoda & A. Hladik. 2008. The species of Dioscorea L. (Dioscoreaceae) from Madagascar with campanulate tori, including a new species from eastern Madagascar. Kew Bulletin 63 (4): 583–600.

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

Last updated: 24 January 2018.

Liliales

Rough bindweed Smilax aspera, photographed by Carsten Niehaus.

Belongs within: Monocotyledoneae.
Contains: Colchicaceae, Liliaceae.

The Liliales is a major clade within the monocotyledons, many species of which are significant garden plants. Many species are bulbous or rhizomatous herbs; others are climbers, such as the greenbriers Smilax and supplejacks Ripogonum. The Corsiaceae are small mycoheterotrophic herbs that lack chlorophyll, found in Australasia and South America. Campynema lineare, green mountain lily, is a small, somewhat grass-like herb with green flowers found in subalpine and alpine regions of Tasmania.

Synapomorphies (from the Angiosperm Phylogeny Website): Storage fructans, chelidonic acid, steroidal saponins present; leaves elliptical, fine venation reticulate; inflorescence terminal; tepals large (sometimes small), free, tepal nectaries present; anthers extrorse; style often long, stigma capitate; ovules many/carpel, parietal tissue absent, nucellar cap present; perianth deciduous; tegmen with cellular structure; endosperm with thick pitted walls, hemicellulosic; mitochondrial sdh3 gene lost.

Liliales [Alstroemeriales]
    |  i. s.: Lachnanthes tinctoria MM96
    |--Arachnitis [Corsiaceae] DS04
    |    `--A. uniflora DS04
    `--+--Campynema [Campynemataceae] DS04
       |    `--C. lineare DS04
       `--+--+--Petermannia [Petermanniaceae] DS04
          |  |    `--P. cirrosa DS04
          |  `--+--Luzuriaga Ruiz & Pav. 1802 (nom. cons.) DS04, ME70 [Luzuriagaceae]
          |     |    |--*L. radicans ME70
          |     |    |--L. marginata ME70
          |     |    `--L. parviflora (Hook. f.) Kunth 1850 (see below for synonymy) ME70
          |     `--+--Colchicaceae DS04
          |        `--Alstroemeriaceae DS04
          |             |--Bomarea HE80
          |             |    |--B. edulis J87
          |             |    `--B. multiflora HE80
          |             `--Alstroemeria SaaRai07
          |                  |--A. aurantiaca Don in Sweet 1833 HE80
          |                  |--A. caryophyllaea DS04
          |                  `--A. pulchella Linnaeus 1781 HE80
          `--+--+--Trillium [Trilliaceae] SaaRai07
             |  |    |--T. grandiflorum DS04
             |  |    |--T. kamtschaticum T03
             |  |    `--T. pusillum R56
             |  |         |--T. p. var. pusillum R56
             |  |         `--T. p. var. virginianum R56
             |  `--Melanthiaceae DS04
             |       |  i. s.: Melanthium YY22
             |       |--Chamaelirium luteum DS04
             |       `--+--Amianthium muscaetoxicum DS04
             |          `--Veratrum DS04
             |               |--V. album DS04
             |               |--V. nigrum C55
             |               `--V. viride DS04
             `--+--+--Philesia [Philesiaceae] DS04
                |  |    `--P. buxifolia DS04
                |  `--Ripogonum Forster & Forster 1776 DS04, ME70 [Ripogonaceae]
                |       |--*R. scandens Forster & Forster 1776 ME70 (see below for synonymy)
                |       |--R. discolor DS04
                |       `--R. elseyanum DS04
                `--+--+--Liliaceae SaaRai07
                   |  `--Calochortus [Calochortaceae] DS04
                   |       |--C. longebarbatus RS98
                   |       `--C. minimus DS04
                   `--Smilax [Smilacaceae] DS04
                        |--S. aspera Linnaeus 1753 PL04
                        |--S. australis LK14
                        |--S. bona-nox WM14
                        |--S. havanensis J87
                        |--S. lanceifolia D07
                        |--S. mexicana GJ09
                        |--S. perfoliata BB07
                        |--S. rotundifolia DS04
                        `--S. zeylanica SanRav07

Luzuriaga parviflora (Hook. f.) Kunth 1850 [=Callixene parviflora Hook. f. 1844, Enargea parviflora (Hook. f.) Skottsb. 1915; incl. C. melantha Col. 1885] ME70

*Ripogonum scandens Forster & Forster 1776 ME70 [incl. Rhipogonum parviflorum C06, Smilax ripogonum Forst. f. 1786 ME70]

*Type species of generic name indicated

REFERENCES

[BB07] Baishya, A. K., & P. J. Bora. 2007. Cross community ethno-medico botany of Dibru-Saikhowa Biosphere Reserve, Assam. Bulletin of the Botanical Survey of India 49 (1–4): 121–154.

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

[C06] Cheeseman, T. F. 1906. Manual of the New Zealand Flora. John Mackay, Government Printer: Wellington.

[D07] Dash, S. S. 2007. Useful plants of Kabi Sacred Grove, Sikkim. Bulletin of the Botanical Survey of India 49 (1–4): 79–88.

[DS04] Davis, J. I., D. W. Stevenson, G. Petersen, O. Seberg, L. M. Campbell, J. V. Freudenstein, D. H. Goldman, C. R. Hardy, F. A. Michelangeli, M. P. Simmons, C. D. Specht, F. Vergara-Silva & M. Gandolfo. 2004. A phylogeny of the monocots, as inferred from rbcL and atpA sequence variation, and a comparison of methods for calculating jackknife and bootstrap values. Systematic Botany 29 (3): 467–510.

[GJ09] Gagné, R. J., & M. Jaschhof. 2009. Cecidomyiidae (gall midges). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 1 pp. 293–314. NRC Research Press: Ottawa.

[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).

[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.

[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.

[ME70] Moore, L. B., & E. Edgar. 1970. Flora of New Zealand vol. 2. Indigenous Tracheophyta: Monocotyledones except Gramineae. A. R. Shearer, Government Printer: Wellington (New Zealand).

[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.

[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.

[RS98] Ramsey, J., & D. W. Schemske. 1998. Pathways, mechanisms, and rates of polyploid formation in flowering plants. Annual Review of Ecology and Systematics 29: 467–501.

[R56] Reed, C. F. 1956. Hyla cinerea in Maryland, Delaware, and Virginia, with notes on the taxonomic status of Hyla cinerea evittata. Journal of the Washington Academy of Sciences 46 (10): 328–332.

[SaaRai07] Saarela, J. M., H. S. Rai, J. A. Doyle, P. K. Endress, S. Mathews, A. D. Marchant, B. G. Briggs & S. W. Graham. 2007. Hydatellaceae identified as a new branch near the base of the angiosperm phylogenetic tree. Nature 446: 312–315.

[SanRav07] Sankar, R. V., K. Ravikumar, N. M. Ganesh Babu & D. K. Ved. 2007. Botany of Anapady MPCA, Palghat district, Kerala with special emphasis on species of conservation concern. Bulletin of the Botanical Survey of India 49 (1–4): 165–172.

[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.

[WM14] Wickett, N. H., S. Mirarab, N. Nguyen, T. Warnow, E. Carpenter, N. Matasci, S. Ayyampalayam, M. S. Barker, J. G. Burleigh, M. A. Gitzendanner, B. R. Ruhfel, E. Wafula, J. P. Der, S. W. Graham, S. Mathews, M. Melkonian, D. E. Soltis, P. S. Soltis, N. W. Miles, C. J. Rothfels, L. Pokorny, A. J. Shaw, L. DeGeronimo, D. W. Stevenson, B. Surek, J. C. Villarreal, B. Roure, H. Philippe, C. W. dePamphilis, T. Chen, M. K. Deyholos, R. S. Baucom, T. M. Kutchan, M. M. Augustin, J. Wang, Y. Zhang, Z. Tian, Z. Yan, X. Wu, X. Sun, G. K.-S. Wong & J. Leebens-Mack. 2014. Phylotranscriptomic analysis of the origin and early diversification of land plants. Proceedings of the National Academy of Sciences of the USA 111 (45): E4859–E4868.

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

Last updated: 27 June 2018.

Araceae

Anthurium jenmanii, photographed by Phil Nelson.

Belongs within: Alismatanae.
Contains: Orontioideae, Lemnaceae, Aroideae, Monsteroideae.

The Araceae are a family of flowering plants in which most species have the flowers reduced and clustered onto a compact spike that is in turn subtended by a large, often colourful bract. Basal aroids include the settler's twine or boorgay Gymnostachys anceps of eastern Australia, distinguished from other aroids by its linear leaves, parallel venation, and a flowering shoot of a complex synflorescence consisting of three to seven short, perennating floral sympodia seperated from each other by a distinct peduncular axis with each sympodium subtended by a leaf-like bract (Nie et al. 2006).

Characters (from the Angiosperm Phylogeny Website): Cyanogenic glucoside triglochinin, flavone C-glycosides present; dimorphic root hypodermis present; sieve tube plastids also with starch; pseudopetiole bundles scattered; stomata unorientated, also anomo- and tetracytic; leaves usually with pseudopetiole, lamina and (pseudo)midrib, tertiary venation cross to reticulate, base with lateral (occasionally auriculate) flanges; inflorescence densely spicate [spadix], unbranched, inflorescence bract well developed, ± coloured, ± surrounding spadix [spathe]; flowers sessile, floral bracts absent; flowers 2-3-merous, in latter case median member of outer whorl of tepals adaxial, tepals ± hooded, with single trace, free (sometimes connate); anthers extrorse; pollen with ektexine present; septal nectaries absent; carpels (basally) ascidiate, fusion usually congenital, loculus usually with secretion, style at most short, stigma also wet; ovules ± unvascularized; fruit a berry; testa multiplicative, ≥5 cells across, often parenchymatous, or with exotesta and/or endotesta and mesotesta lignified, tegmen collapsed; x = 16; cotyledon not photosynthetic.

<==Araceae [Arales, Aranae, Ariflorae, Calleae]
    |--+--Orontioideae NS06
    |  `--Gymnostachys [Gymnostachydoideae] NS06
    |       `--G. anceps Brown 1810 CD07
    `--+--Lemnaceae TB04
       `--+--+--Aroideae NS06
          |  `--Lasioideae TB04
          |       |--Anaphyllopsis americana TB04
          |       `--Dracontium polyphyllum NS06
          `--+--Monsteroideae NS06
             `--Pothoideae TB04
                  |--Pothos [Potheae] NS06
                  |    |--P. ovalifolius TB04
                  |    `--+--P. junghuhnii TB04
                  |       `--P. scandens TB04
                  `--Anthurium [Anthurieae] TB04
                       |--A. guildingii TB04
                       |--A. jenmannii TB04
                       `--A. scandens J87

Araceae incertae sedis:
  Zantedeschia HE80
    |--Z. aethiopica (Linnaeus) Sprengel 1826 HE80
    `--Z. albomaculata HE80
  Adelonema YY22
  Aglaodorum YY22
  Alocasiophyllum YY22
  Amauriella YY22
  Aphyllarum YY22
  Asterostigma YY22
  Biarum YY22
  Bucephalandra YY22
  Caladiopsis YY22
  Callopsis YY22
  Chamaecladon YY22
  Chlorospatha YY22
  Diandriella YY22
  Gamogyne YY22
  Gearum YY22
  Gonatanthus YY22
  Gorgonidium YY22
  Hapaline YY22
  Helicodiceros YY22
  Hydrosme YY22
  Lagenandra YY22
  Mangonia YY22
  Microcasia YY22
  Philonotion YY22
  Pinellia YY22
  Piptospatha YY22
  Plesmonium YY22
  Porophyrospatha YY22
  Pseudohydrosme YY22
  Remusatia YY22
  Rhynchopyle YY22
  Scaphispatha YY22
  Schismatoglottis YY22
  Schizocasia YY22
  Spathantheum YY22
  Stylochiton YY22
  Synantherias YY22
  Syngonium YY22
  Taccarum YY22
  Thaumatophyllum YY22
  Theriophonum YY22
  Typhonium SVD02
    |--T. diversifolium O88
    |--T. horsfieldii (Miquel) Steenis 1948 (see below for synonymy) SVD02
    |--T. liliifolium LK14
    |--T. nudibaccatum LK14
    |--T. pedatum Schott 1857 SVD02
    `--T. peltandroides LK14
  Typhonodorum YY22
  Ulearum YY22
  Xenophya YY22
  Zomicarpa YY22
  Zomicarpella YY22
  Synandrospadix YY22
  Pothoidium YY22
  Dracunculus vulgaris Schott in Schott & Endlicher 1832 HE80
  Cryptocoryne P88
    |--C. ciliata P88
    |--C. cordata P88
    `--C. crispatula YG03
  Cyrtosperma S68
  Lasia spinosa BB07
  Anaphyllum wightii UB06
  Urospatha MM96
    |--U. caudata MM96
    `--U. friedrichsthallii MM96
  Landoltia punctata LK14
  Richardia africana Kunth 1818 C06

Typhonium horsfieldii (Miquel) Steenis 1948 [=Sauromatum horsfieldii Miquel 1856; incl. T. fallax Brown 1880, Heterostalis pedata Schott 1864 non T. pedatum Schott 1857] SVD02

*Type species of generic name indicated

REFERENCES

[BB07] Baishya, A. K., & P. J. Bora. 2007. Cross community ethno-medico botany of Dibru-Saikhowa Biosphere Reserve, Assam. Bulletin of the Botanical Survey of India 49 (1–4): 121–154.

[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.

[C06] Cheeseman, T. F. 1906. Manual of the New Zealand Flora. John Mackay, Government Printer: Wellington.

[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).

[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.

[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.

[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.

[NS06] Nie, Z.-L., H. Sun, H. Li & J. Wen. 2006. Intercontinental biogeography of subfamily Orontioideae (Symplocarpus, Lysichiton, and Orontium) of Araceae in eastern Asia and North America. Molecular Phylogenetics and Evolution 40 (1): 155–165.

[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.

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

[SVD02] Sasikala, K., E. Vajravelu & P. Daniel. 2002. Typhonium horsfieldii (Miq.) Steenis, family Araceae, a new report for India. Journal of the Bombay Natural History Society 99 (3): 581–583.

[S68] Simons, E. L. 1968. African Oligocene mammals: introduction, history of study, and faunal succession. Peabody Museum of Natural History, Yale University, Bulletin 28: 1–21.

[TB04] Tam, S.-M., P. C. Boyce, T. M. Upson, D. Barabé, A. Bruneau, F. Forest & J. S. Parker. 2004. Intergeneric and infrafamilial phylogeny of subfamily Monsteroideae (Araceae) revealed by chloroplast trnL-F sequences. American Journal of Botany 91 (3): 490–498.

[UB06] Udayan, P. S. & I. Balachandran. 2006. Cleistanthus sankunnianus Sivar. & Indu Balach.—a rare and little known endemic plant rediscovered from wild populations in Kollam district of Kerala state, India. Bulletin of the Botanical Survey of India 48 (1–4): 217–218.

[YG03] Yadav, S. R., & S. P. Gaikwad. 2003. A revision of the Indian Aponogetonaceae. Bulletin of the Botanical Survey of India 45 (1–4): 39–76.

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

Last updated: 24 April 2018.

Mesangiospermae

Hornwort Ceratophyllum demersum, copyright Eyeweed.

Belongs within: Angiospermae.
Contains: Chloranthaceae, Magnoliidae, Monocotyledoneae, Ranunculales, Proteales, Buxanae, Gunneridae.

The Mesangiospermae were defined by Cantino et al. (2007) as the most inclusive crown clade including Platanus occidentalis but excluding Nymphaea odorata and Austrobaileya scandens; as such, it represents the clade of flowering plants excluding the basal ANITA grade. The clade is supported by molecular analyses, but reliable morphological synapomorphies have not yet been identified. The Mesangiospermae contains five well-supported clades—the monocotyledons, eudicotyledons, magnoliids, Chloranthaceae and Ceratophyllum—but relationships between these clades remain unsettled. Ceratophyllum, hornworts, is a cosmopolitan genus of aquatic plants bearing rings of forked leaves and lacking true roots.

The Eudicotyledoneae is the clade containing the majority of species previously grouped in flowering plant classifications as 'dicotyledons' (the characters uniting dicotyledons in the broad sense are now regarded as plesiomorphies of flowering plants as a whole). Eudicots are distinguished by the possession of tricolpate pollen grains; it is possible that the loss of oil cells in the mesophyll and the dry fruit wall also represents a synapomorphy of the clade (Cantino et al. 2007). Cantino et al. (2007) defined Eudicotyledoneae as the least inclusive clade containing Ranunculus, Platanus, Sabia, Trochodendron, Buxus and Helianthus.

<==Mesangiospermae [Ranales] CD07
    |--+--Chloranthaceae WM14
    |  `--Magnoliidae WM14
    `--+--Monocotyledoneae APG16
       `--+--Ceratophyllum Linnaeus 1753 KC01 [Ceratophyllaceae, Ceratophyllales APG16]
          |    |--C. demersum BL04
          |    |--C. furcatispinum CBH93
          |    `--C. submersum BL04
          `--Eudicotyledoneae (see below for synonymy) APG16
               |--Ranunculales SR07
               `--+--Proteales APG16
                  `--+--+--Buxanae APG16
                     |  `--Gunneridae SR07
                     `--Trochodendraceae [Trochodendrales] APG16
                          |--Tetracentron [Tetracentraceae, Tetracentroideae] CD07
                          |--Trochodendron [Trochodendroideae] CD07
                          |    `--T. aralioides Siebold & Zucc. 1838 CD07
                          `--Trochodendroides CBH93
                               |--T. arctica Z02
                               `--T. rhomboideus (Lesquereux) Berry 1922 CBH93

Eudicotyledoneae [Dilleniidae, Dillenianae, Ranunculanae, Ranunculidae, Sapindineae, Tricolpatae, Trochodendrineae] APG16

*Type species of generic name indicated

REFERENCES

[APG16] Angiosperm Phylogeny Group. 2016. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Botanical Journal of the Linnean Society 181: 1–20.

[BL04] Barkman, T. J., S.-H. Lim, K. M. Salleh & J. Nais. 2004. Mitochondrial DNA sequences reveal the photosynthetic relatives of Rafflesia, the world’s largest flower. Proceedings of the National Academy of Sciences of the USA 101 (3): 787–792.

[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.

[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.

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

[SR07] Saarela, J. M., H. S. Rai, J. A. Doyle, P. K. Endress, S. Mathews, A. D. Marchant, B. G. Briggs & S. W. Graham. 2007. Hydatellaceae identified as a new branch near the base of the angiosperm phylogenetic tree. Nature 446: 312–315.

[WM14] Wickett, N. H., S. Mirarab, N. Nguyen, T. Warnow, E. Carpenter, N. Matasci, S. Ayyampalayam, M. S. Barker, J. G. Burleigh, M. A. Gitzendanner, B. R. Ruhfel, E. Wafula, J. P. Der, S. W. Graham, S. Mathews, M. Melkonian, D. E. Soltis, P. S. Soltis, N. W. Miles, C. J. Rothfels, L. Pokorny, A. J. Shaw, L. DeGeronimo, D. W. Stevenson, B. Surek, J. C. Villarreal, B. Roure, H. Philippe, C. W. dePamphilis, T. Chen, M. K. Deyholos, R. S. Baucom, T. M. Kutchan, M. M. Augustin, J. Wang, Y. Zhang, Z. Tian, Z. Yan, X. Wu, X. Sun, G. K.-S. Wong & J. Leebens-Mack. 2014. Phylotranscriptomic analysis of the origin and early diversification of land plants. Proceedings of the National Academy of Sciences of the USA 111 (45): E4859–E4868.

[Z02] Zherikhin, V. V. 2002. Insect trace fossils. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 303–324. Kluwer Academic Publishers: Dordrecht.

Last updated: 18 July 2021.

Austrobaileyales

Japanese star anise Illicium anisatum, photographed by Kenpei.

Belongs within: Angiospermae.

The Austrobaileyales is a small clade (about 100 species) of flowering plants found in the Indo-Australian region and southern North America. The clade has been shown to occupy a basal position in angiosperm phylogeny.

Synapomorphies (from the Angiosperm Phylogeny Website): Tiglic acid present; vessels solitary; nodes 1:2; petiole bundle(s) arcuate; mucilaginous extragynoecial compitum present, outer integument 5-7 cells thick; fruit a berrylet; perianth deciduous; mesotestal cells ± sclerotic; endosperm starchy.

<==Austrobaileyales [Illiciales]
    |--Austrobaileya [Austrobaileyaceae] SR07
    |    `--A. scandens White 1933 CD07
    `--+--Trimeniaceae FF01
       |    |--Piptocalyx C01
       |    `--Trimenia C01
       `--Illiciineae T00
            |--Schisandraceae SR07
            |    |--Kadsura T00
            |    `--Schisandra SR07
            |         |--S. chinensis WZ99
            |         `--S. sphenanthera SR07
            `--Illicium [Illiciaceae] SR07
                 |--I. anisatum DS04
                 |--I. ekmanii J87
                 |--I. geiseltalense JD05
                 |--I. parviflorum DS04
                 `--I. religiosum LO98

*Type species of generic name indicated

REFERENCES

[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.

[C01] Carlquist, S. 2001. Observations on the vegetative anatomy of Austrobaileya: Habital, organographic and phylogenetic conclusions. Botanical Journal of the Linnean Society 135 (1): 1-11.

[DS04] Davis, J. I., D. W. Stevenson, G. Petersen, O. Seberg, L. M. Campbell, J. V. Freudenstein, D. H. Goldman, C. R. Hardy, F. A. Michelangeli, M. P. Simmons, C. D. Specht, F. Vergara-Silva & M. Gandolfo. 2004. A phylogeny of the monocots, as inferred from rbcL and atpA sequence variation, and a comparison of methods for calculating jackknife and bootstrap values. Systematic Botany 29 (3): 467-510.

[FF04] Friedman, W. E., & S. K. Floyd. 2001. Perspective: The origin of flowering plants and their reproductive biology – a tale of two phylogenies. Evolution 55 (2): 217-231.

[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.

[JD05] Junge, F. W., M. Dolezych, H. Walther, T. Böttger, A. Kühl, L. Kunzmann, P. Morgenstern, T. Steinberg & R. Stange. 2005. Ein Fenster in Landschaft und Vegetation vor 37 Millionen Jahren: Lithologische, sedimentgeochemische und paläobotanische Befunde aus einem Paläoflusssystem des Weißelsterbeckens. Mauritiana 19 (2): 185-273.

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

[SR07] Saarela, J. M., H. S. Rai, J. A. Doyle, P. K. Endress, S. Mathews, A. D. Marchant, B. G. Briggs & S. W. Graham. 2007. Hydatellaceae identified as a new branch near the base of the angiosperm phylogenetic tree. Nature 446: 312-315.

[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.

[WZ09] Wang Y., Zhou J., Tan N., Ding Z. & Jiang X. 1999. Cyclic dipeptides from Schizandra chinensis and their syntheses. Yaoxue Xuebao [Acta Pharmaceutica Sinica] 34: 19-22.

Aroideae

Philodendron insigne, copyright Bernard Dupont.

Belongs within: Araceae.
Contains: Arisaema.

The Aroideae is a clade within the family Araceae with the spadix (flower-spike) divided into a male and female region. Members of the Aroideae except the Zamioculcadeae are united by the presence of lactifers and of pollen with a thin thin ectexine, thick and spongy endexine and massive intine (Angiosperm Phylogeny Website). Notable members of the clade include taro Colocasia esculenta, widely grown for its edible corms. Many, such as species of Philodendron, are popular ornamental plants grown primarily for their foliage. The titan arum Amorphophallus titanum produces the world's largest unbranched inflorescence, reaching a height of up to 2.5 metres.

Synapomorphies (from the Angiosperm Phylogeny Website, as 'Zamioculcadoideae + Aroideae'): Spathe differentiated into tube plus blade, spadix differentiated into zones with staminate and carpellate flowers; flowers imperfect; pollen inaperturate.

<==Aroideae
    |--Zamioculcas [Zamioculcadeae] TB04
    |    `--Z. zamiifolia TB04
    `--+--+--Nephthytideae TB04
       |  |    |--Anchomanes difformis TB04
       |  |    `--Nephthytis afzelii TB04
       |  `--Aglaonema [Aglaonemateae] TB04
       |       |--A. griffithii P88
       |       |--A. modestum TB04
       |       `--A. oblongifolium K03
       `--+--Anubias [Anubiadeae] TB04
          |    `--A. barteri TB04
          |         |--A. b. var. barteri TB04
          |         `--A. b. var. caladiifolia TB04
          |--+--Culcasieae TB04
          |  |    |--Culcasia saxatilis TB04
          |  |    `--+--Cercestis stigmaticus TB04
          |  |       `--Rhektophyllum mirabile TB04
          |  `--+--Dieffenbachia [Dieffenbachieae] TB04
          |     |    `--D. pittieri TB04
          |     `--Spathicarpa [Spathicarpeae] TB04
          |          `--S. sagittifolia TB04
          |--+--Montrichardia [Montrichardieae] TB04
          |  |    `--M. arborescens [incl. M. aculeata] BTA75
          |  `--+--Homalomena [Homalomeneae] TB04
          |     |    |--H. propinga K03
          |     |    `--H. rubescens TB04
          |     `--Philodendron [Philodendreae, Philodendroideae] TB04
          |          |  i. s.: P. callosum NS06
          |          |         P. consanguineum J87
          |          |         P. sinuatum MM96
          |          |--P. insigne TB04
          |          |--P. rudgeanum TB04
          |          |--P. solimoesense TB04
          |          `--+--P. fragrantissimum TB04
          |             `--+--P. billietiae TB04
          |                `---P. melinonii TB04
          `--+--+--Caladieae TB04
             |  |    |--Caladium TB04
             |  |    |    |--‘Arum’ arborescens V01
             |  |    |    |--‘Arum’ auritum V01
             |  |    |    |--C. bicolor TB04 [=Arum bicolor V01]
             |  |    |    |--‘Arum’ helleborifolium V01
             |  |    |    |--C. nymphaeaefolium V01
             |  |    |    |--‘Arum’ sagittaefolium V01
             |  |    |    |--‘Arum’ seguinum V01
             |  |    |    `--C. solocasia AP80
             |  |    `--Xanthosoma TB04
             |  |         |--X. sagittifolium TB04
             |  |         `--X. undipes C09
             |  `--Thomsonieae TB04
             |       |--Pseudodracontium FPC04
             |       `--Amorphophallus TB04
             |            |--A. galbra LK14
             |            |--A. gigas B14
             |            |--A. paeoniifolius TB04
             |            |    |--A. p. var. paeoniifolius TB04
             |            |    `--A. p. var. campanulatus TB04
             |            `--A. titanum CG12
             `--+--Peltandra [Peltandreae] TB04
                |    `--P. virginica TB04
                `--+--+--Arisaema TB04
                   |  `--Pistia [Pistieae] TB04
                   |       `--P. stratiotes TB04
                   `--Colocasieae [Colocasioideae] TB04
                        |--Protarum FPC04
                        |--Steudnera FPC04
                        |--Colocasia HE80
                        |    |--C. esculenta (Linnaeus) Schott in Schott & Endlicher 1832 HE80 (see below for synonymy)
                        |    |    |--C. e. var. esculenta LK14
                        |    |    `--C. e. var. aquatilis LK14
                        |    `--C. macrorrhiza M87
                        `--Alocasia TB04
                             |--A. cuprea H03
                             |--A. denudata P88
                             |--A. macrorrhiza (Linnaeus) Don in Sweet 1839 HE80
                             `--A. odora TB04

Aroideae incertae sedis:
  Arum HE80
    |--A. dioscoridis PT98
    |--A. dracunculus D97
    |--A. italicum Miller 1768 HE80
    `--A. maculatum TG88
  Arisarum [Arisareae] FPC04
    `--A. vulgare PT98
  Ambrosina [Ambrosineae] FPC04
  Calla [Calloideae] NS06
    `--C. palustris NS06

Colocasia esculenta (Linnaeus) Schott in Schott & Endlicher 1832 HE80 [=Arum esculentum V01, Caladium esculentum HE80; incl. C. esculenta var. antiquorum HE80, Arum colocasia C55]

*Type species of generic name indicated

REFERENCES

[AP80] Andrews, K. L., & S. L. Poe. 1980. Spider mites of El Salvador, Central America (Acari: Tetranychidae). Florida Entomologist 63 (4): 502–505.

[BTA75] Baker, E. W., D. M. Tuttle & M. J. Abbatiello. 1975. The false spider mites of northwestern and north central Mexico (Acarina: Tenuipalpidae). Smithsonian Contributions to Zoology 194: 1–23.

[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. 1855. Géographie Botanique Raisonée: Ou exposition des faits principaux et des lois concernant la distribution géographique des plantes de l’époque actuelle vol. 2. Librairie de Victor Masson: Paris.

[C09] Chaverri, L. G. 2009. Culicidae (mosquitos, zancudos). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 1 pp. 369–388. NRC Research Press: Ottawa.

[CG12] Clouse, R., & G. Giribet. 2012. On the Cyphophthalmi (Arachnida, Opiliones) types from the Museo Civico di Storia Naturale "Giacomo Doria". Bulletin of the Museum of Comparative Zoology 160 (5): 241–257.

[D97] Duméril, A. M. C. 1797. Extrait d'une dissertation sur l'organe de l'odorat dans les insectes. Bulletin des Sciences, par la Societé Philomathique de Paris 1 (5): 34.

[FPC04] Friis, E. M., K. R. Pedersen & P. R. Crane. 2004. Araceae from the Early Cretaceous of Portugal: evidence on the emergence of monocotyledons. Proceedings of the National Academy of Sciences of the USA 101 (47): 16565–16570.

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

[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).

[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.

[K03] Kulip, J. 2003. An ethnobotanical survey of medicinal and other useful plants of Muruts in Sabah, Malaysia. Telopea 10 (1): 81–98.

[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.

[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.

[M87] Mueller, F. von. 1887. Some hitherto undescribed plants of New South Wales. Proceedings of the Linnean Society of New South Wales, series 2, 1 (4): 1105–1110.

[NS06] Nie, Z.-L., H. Sun, H. Li & J. Wen. 2006. Intercontinental biogeography of subfamily Orontioideae (Symplocarpus, Lysichiton, and Orontium) of Araceae in eastern Asia and North America. Molecular Phylogenetics and Evolution 40 (1): 155–165.

[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.

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

[TB04] Tam, S.-M., P. C. Boyce, T. M. Upson, D. Barabé, A. Bruneau, F. Forest & J. S. Parker. 2004. Intergeneric and infrafamilial phylogeny of subfamily Monsteroideae (Araceae) revealed by chloroplast trnL-F sequences. American Journal of Botany 91 (3): 490–498.

[TG88] Tepfer, D., A. Goldmann, N. Pamboukdjian, M. Maille, A. Lepingle, D. Chevalier, J. Dénarié & C. Rosenberg. 1988. A plasmid of Rhizobium meliloti 41 encodes catabolism of two compounds from root exudate of Calystegium sepium. Journal of Bacteriology 170 (3): 1153–1161.

[V01] Ventenat, C. 1801. Dissertation sur le genre Arum. Bulletin des Sciences, par la Societé Philomathique de Paris 2 (46): 171–172.

Last updated: 13 November 2018.