Distribution Tropical Africa, Madagascar, tropical Asia, southern China, Japan, Taiwan, islands of the western Pacific, Malesia, New Guinea, northern Queensland, Melanesia, tropical America.

Fossils Flattened grooved unilocular endocarps, assigned to Icacinaceae, with deeply foveolate-reticulate surface and a papillate inner lining have been found in the Late Turonian to the Santonian of Central Europe and were described as Icacinicarya budvarensis. Similar endocarps from the Maastrichtian of Germany have been attributed to Icacinicarya papillaris and Iodes germanica. From the Palaeogene of eastern North America are many reports of fossil Icacinaceae and the extinct Stizocaryopsis was described from the Paleocene of Egypt. Fossilized wood, leaves and additional endocarps assigned to Icacinaceae are also known, e.g. from the Eocene of England (Icacinicarya and Palaeophytocrene described from the London Clay) and Germany. Fossil fruits of Palaeophytocrene (Phytocreneae) from Mid and Late Paleocene have been found in western North America and Colombia (Stull & al. 2012).

Habit Usually bisexual (sometimes andromonoecious?, gynomonoecious?, polygamomonoecious?, dioecious, androdioecious?, or gynodioecious?), evergreen trees, shrubs or lianas with non-axillary branch tendrils.

Vegetative anatomy hellogen ab initio superficial. Medullary vascular bundles present in Iodes. Secondary lateral growth in lianas often anomalous (via concentric/successive cambia or from simple cylindrical cambium; interxylary phloem rarely present; rarely with successive cambia). Vessel elements with usually simple (sometimes scalariform) perforation plates; lateral pits alternate to opposite, bordered or simple pits. Imperforate tracheary xylem elements tracheids, fibre tracheids or libriform fibres with simple or bordered pits, usually non-septate. Wood rays uniseriate and/or multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty, aliform, winged-aliform, confluent, scalariform, vasicentric, or banded. Tyloses sometimes present. Secondary phloem stratified, at least in young stems. Intraxylary phloem often present in lianas. Sieve tube plastids S type. Nodes usually 1:1, unilacunar with one leaf trace (sometimes 3:3, trilacunar with three traces, or 5:5, pentalacunar with five leaf traces). Secretory cavities (with latex) present or absent? Cortex with or without cristarque cells? Sclereids present. Asterosclereids sometimes present. Prismatic calciumoxalate crystals or groups of crystals usually in parenchyma cells (rhombic crystals, crystal sand, druses or styloid-like crystals sometimes present). Tanniniferous cells occasionally abundant.

Trichomes Hairs unicellular, often adpressed, with short stalk and T-shaped with body at right angles to stalk (icacinaceous hairs), sometimes unbranched, sometimes globular, or absent.

Leaves Usually alternate (spiral; occasionally opposite), simple, usually entire (rarely palmately lobed), often coriaceous, usually with conduplicate (rarely conduplicate-plicate) ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate and petiole with wing bundles, or bundle transection annular. Venation usually pinnate (rarely palmate), brochidodromous. Stomata usually cyclocytic (sometimes anomocytic, anisocytic or modified paracytic). Cuticular wax crystalloids? Domatia usually absent (sometimes as pits or pockets). Secretory cavities (with latex) present or absent? Mesophyll sometimes with sclerenchymatous idioblasts. Calciumoxalate druses occasionally present. Leaf margin usually entire (sometimes serrate or spinose-serrate).

Inflorescence Usually axillary (rarely terminal), paniculate, corymbose cymose, spicate or racemose, sometimes a thyrse. Floral prophylls (bracteoles) in Oncotheca triangular, two or three per flower.

Flowers Actinomorphic, small. Pedicel articulated. Hypogyny. Sepals (three to) five (or six)?, usually with imbricate to imbricate quincuncial (rarely valvate?) aestivation, persistent in fruit, usually more or less connate (rarely absent). Petals (three to) five (or six)?, usually with valvate (occasionally imbricate) aestivation, caducous, usually free (sometimes connate at base; rarely absent), with incurved apex and sometimes adaxial keel. Nectariferous disc usually absent (sometimes discoid, columellate or lobed).

Androecium Stamens (three to) five (or six)?, haplostemonous, antesepalous, alternipetalous. Filaments free from each other and usually from tepals (rarely adnate to corolla tube, epipetalous). Anthers usually dorsifixed (rarely basifixed), versatile or non-versatile, usually tetrasporangiate (in Oncotheca disporangiate, dithecal), usually introrse (sometimes extrorse or latrorse?), usually longicidal (dehiscing by longitudinal slits; rarely poricidal, dehiscing by apical pores?). Tapetum secretory, with multinucleate cells. Female flowers often with staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually triaperturate, colpate, colporoidate, colporate, pororate, porate och forate (rarely up to 7-aperturate or inaperturate), shed as monads, bicellular at dispersal. Exine tectate or tectate-perforate, sometimes with columellate infratectum, sometimes foveolate, foveate foveate-reticulate, echinate, or smooth.

Gynoecium Pistil composed of usually two (rarely one, three? or five) connate carpels. Ovary superior, unilocular (pseudomonomerous? to quinquelocular), without fleshy appendage. Style single, simple, short, or absent. Stigma wide (when style absent) or punctate, type? Stylodia in Oncotheca five, free, recurved, conduplicate. Stigmas ventral, punctate, type? Male flowers usually with pistillodium.

Ovules Placentation usually apical (axile in plurilocular ovaries). Ovules usually two (rarely one) per ovary, anatropous, pendulous, apotropous (sometimes epitropous), unitegmic (in Phytocrene apically bitegmic, with integuments free in micropyle), usually (thinly) crassinucellar (sometimes tenuinucellar). Integument four to more than ten cell layers thick. Funicular obturator present. Megagametophyte monosporous, Polygonum type. Synergids often with a filiform apparatus. Antipodal cells ephemeral. Endosperm development usually nuclear. Endosperm haustoria present or absent (rarely chalazal, often elongate). Endosperm sometimes ruminate. Embryogenesis?

Fruit A usually one-seeded (rarely two- to five-seeded) drupe, often flattened and/or ridged, with persistent calyx. Endocarp cells sometimes papillate.

Seeds Aril absent. Exotesta present. Endotesta present. Perisperm not developed. Endosperm copious or sparse, oily (sometimes starchy), sparse or absent (sometimes ruminate). Embryo straight or curved?, usually long, well differentiated, with chlorophyll. Cotyledons usually two (rarely three), foliaceous. Germination phanerocotylar or cryptocotylar.

Cytology n = 10, 11 (Pyrenacantha), 12, 20 (Pyrenacantha, Stachyanthus), 24 (Icacina), 25 (Oncotheca)

Phytochemistry Flavonols (quercetin), cyanidin, monoterpene secoiridoids, diterpenoids, ipecacalkaloids, camptothecine (monoterpene indole [quinolone] alkaloid), sesquiterpenoids, triterpenoid saponins, and cyanogenic compounds present. Route I iridoids? Ellagic acid not found. In Cassinopsis cassinopin (a glucoside), kaempferol trirhamnoside and verbascosides. Tannins present. Aluminium accumulated in some species.

Phytocrenaceae Arn. ex R. Br. in J. J. Bennett et R. Brown, Plant. Jav. Rar.: 244. 13 Mar 1852 [’Phytocreneae’]; Iodaceae Tiegh. in Bot. Jahresber. (Just) 25(2): 406. 19 Jan 1900 [’Jodaceae’]; Pleurisanthaceae Tiegh. in Bot. Jahresber. (Just) 25(2): 406. 19 Jan 1900; Sarcostigmataceae Tiegh. in Bot. Jahresber. (Just) 25(2): 406. 19 Jan 1900; Icacinales Tiegh. in Bot. Jahresber. (Just) 25(2): 406. 19 Jan 1900; Icacinanae Doweld, Tent. Syst. Plant. Vasc.: li. 23 Dec 2001

Distribution Tropical and southern Africa, Madagascar, tropical Asia, southern China, Japan, Taiwan, islands of the western Pacific, Malesia to New Guinea, northern Queensland, Melanesia, tropical America.

Vegetative anatomy Phellogen ab initio superficial. Medullary vascular bundles present in Iodes. Secondary lateral growth in lianas often anomalous (via concentric/successive cambia or from simple cylindrical cambium; Chlamydocarya and Sarcostigma with interxylary phloem; Icacina sometimes with successive cambia). Vessel elements with usually simple (sometimes scalariform) perforation plates; lateral pits alternate to opposite, bordered pits (in e.g. Cassinopsis). Imperforate tracheary xylem elements tracheids, fibre tracheids or libriform fibres with simple or bordered pits, usually non-septate. Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty, aliform, winged-aliform, confluent, scalariform, vasicentric, or banded. Tyloses sometimes present in Icacina and Mappia. Secondary phloem stratified. Intraxylary phloem often present in lianas (e.g. Sarcostigma). Sieve tube plastids S type. Nodes usually 1:1, unilacunar with one leaf trace (sometimes 3:3, trilacunar with three traces). Secretory cavities (with latex) present or absent? Cortex with or without cristarque cells? Sclereids present. Prismatic calciumoxalate crystals or groups of crystals usually in parenchyma cells (rhombic crystals in Alsodeiopsis, Desmostachys, Icacina and Lavigeria); crystal sand present in wood rays in Lavigeria; druses present in Cassinopsis; styloid-like crystals present in some species; chambered crystalliferous strands present in Cassinopsis.

Trichomes Hairs unicellular, often adpressed, with short stalk and T-shaped with body at right angles to stalk (icacinaceous hairs), sometimes unbranched, sometimes globular, or absent.

Leaves Usually alternate (spiral; in Cassinopsis and Iodes opposite), simple, usually entire (rarely palmately lobed), often coriaceous, usually with conduplicate (rarely conduplicate-plicate) ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate and petiole with wing bundles, or bundle transection annular (petiole in Iodes also with medullary bundles). Venation usually pinnate (in Hosiea palmate), brochidodromous. Stomata usually cyclocytic (sometimes anomocytic or anisocytic). Cuticular wax crystalloids? Domatia usually absent (sometimes as pits or pockets). Secretory cavities (with latex) present or absent? Mesophyll with or without sclerenchymatous idioblasts? Leaf margin usually entire (sometimes serrate or spinose-serrate, in Hosiea with long teeth).

Inflorescence Usually axillary (rarely terminal), paniculate, corymbose cymose, spicate or racemose.

Flowers Actinomorphic, small. Pedicel articulated. Hypogyny. Sepals (three to) five (or six)?, usually with imbricate (rarely valvate?) aestivation, persistent in fruit, usually more or less connate (in Phytocrene free or almost free; absent in Pyrenacantha). Petals (three to) five (or six)?, usually with valvate aestivation, usually free (sometimes connate at base; rarely absent), with incurved apex and sometimes adaxial keel. Nectariferous disc usually absent (sometimes discoid, columellate or lobed).

Androecium Stamens (three to) five (or six)?, haplostemonous, antesepalous, alternipetalous. Filaments free from each other and usually from tepals (rarely adnate to corolla tube, epipetalous). Anthers dorsifixed, versatile?, tetrasporangiate, usually introrse (sometimes latrorse?), usually longicidal (dehiscing by longitudinal slits; rarely poricidal, dehiscing by apical pores?). Tapetum secretory, with multinucleate cells. Female flowers often with staminodia.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually triaperturate, colpate, colporoidate, colporate, pororate, porate och forate (rarely up to 7-aperturate; in Stachyanthus inaperturate), shed as monads, bicellular at dispersal. Exine tectate, with ? infratectum, in Cassinopsis foveolate, in Lavigeria foveate, in Icacina foveate-reticulate, in Iodes and Polycephalium echinate.

Gynoecium Pistil composed of usually two (rarely one or three?) connate carpels. Ovary superior, unilocular (pseudomonomerous?), without fleshy appendage. Style single, simple, short, or absent. Stigma wide (when style absent) or punctate, type? Male flowers with pistillodium.

Ovules Placentation usually apical (axile in plurilocular ovaries). Ovules usually two (rarely one) per ovary, anatropous, pendulous, apotropous, unitegmic (in Phytocrene apically bitegmic, with integuments free in micropyle), usually (thinly) crassinucellar (sometimes tenuinucellar). Integument 7–10 (Pyrenacantha) or more than ten (Cassinopsis) cell layers thick. Funicular obturator present. Megagametophyte monosporous, Polygonum type. Synergids often with a filiform apparatus. Antipodal cells ephemeral. Endosperm development usually nuclear (in Nothapodytes cellular). Endosperm haustoria present or absent (in Nothapodytes chalazal, often elongate). Endosperm at least in Nothapodytes ruminate. Embryogenesis?

Fruit A one-seeded drupe, often flattened and/or ridged, with persistent calyx. Endocarp cells sometimes papillate.

Seeds Aril absent. Exotesta present. Endotesta present. Perisperm not developed. Endosperm copious or sparse, oily (in Merrilliodendron starchy), sparse or absent (sometimes ruminate). Embryo straight or curved?, usually long, well differentiated, with chlorophyll. Cotyledons two, foliaceous. Germination phanerocotylar or cryptocotylar.

Cytology n = 10, 11 (Pyrenacantha), 12, 20 (Pyrenacantha, Stachyanthus), 24 (Icacina)

Phytochemistry Flavonols (quercetin), cyanidin, monoterpene secoiridoids, diterpenoids, ipecacalkaloids, camptothecine (monoterpene indole [quinolone] alkaloid; in Merrilliodendron and Nothapodytes), sesquiterpenoids, triterpenoid saponins, and cyanogenic compounds present. Route I iridoids? Ellagic acid not found. In Cassinopsis cassinopin (a glucoside), kaempferol trirhamnoside and verbascosides. Aluminium accumulated in some species.

Use Timber, medicinal plants (seed oil from Sarcostigma), starch sources (tubers of Casimirella).

Systematics Cassinopsis (6; C. chapelieri, C. ciliata, C. ilicifolia, C. madagascariensis, C. tinifolia, C. tomentosa; southern and eastern Africa, Madagascar); Nothapodytes (11; East Asia, tropical Asia), Mappia (5; M. angustifolia, M. longipes, M. mexicana, M. multiflora, M. racemosa; southern Mexico, Central America, the Greater Antilles); Alsodeiopsis (11; tropical Africa), Pleurisanthes (7; P. artocarpi, P. brasiliensis, P. emarginata, P. flava, P. howardii, P. parviflora, P. simpliciflora; tropical South America), Merrilliodendron (1; M. megacarpum; the Philippines, islands in western Pacific), Lavigeria (1; L. macrocarpa; Cameroon, Gabon, Congo), Icacina (5; I. claessensii, I. guessfeldtii, I. mannii, I. oliviformis, I. trichantha; tropical Africa), Casimirella (7; C. ampla, C. beckii, C. crispula, C. diversifolia, C. guaranitica, C. lanata, C. rupestris; Central America, tropical South America), Leretia (2; L. cordata, L. racemosa; tropical South America); Mappianthus (1; M. iodoides; southern China), ‘Iodes’ (28; tropical regions in the Old World; paraphyletic; incl. Polyporandra?), Polyporandra (1; P. scandens; East Malesia to New Guinea, Melanesia; in Iodes?); Desmostachys (6; D. brevipes, D. longipes, D. oblongifolius, D. planchonianus, D. tenuifolius, D. vogelii; tropical Africa, Madagascar), Natsiatum (1; N. herpeticum; eastern Himalayas to Southeast Asia), Hosiea (2; H. japonica, H. sinensis; western and central China, Japan), Rhyticaryum (12; East Malesia to New Guinea and western Pacific islands), Sarcostigma (2; S. kleinii, S. paniculata; India to Southeast Asia and Malesia), Miquelia (7; M. assamica, M. caudata, M. celebica, M. kleinii, M. philippinensis, M. reticulata, M. thorelii; tropical Asia), Phytocrene (11; Southeast Asia, Malesia), Stachyanthus (4; S. cuneatus, S. donisii, S. occidentalis, S. zenkeri; tropical Africa), Pyrenacantha (c 35; tropical regions in the Old World)

Unplaced Icacinaceae Natsiatopsis (1; N. thunbergiifolia; Burma, southern Yunnan), Sleumeria (1; S. auriculata; northern Borneo).

Cassinopsis is sister to the remaining Icacinaceae (Byng & al. 2014, Stull & al. 2015)

The Icacina clade is morphologically homogenous and characterized by, e.g., vessel elements with simple perforation plates, lateral pits usually alternate, relatively short vessel elements and fibres, axial parenchyma banded and vasicentric, wood rays multiseriate, high and wide, and variation in the shape of the cambium. Almost all species are lianas.

Sarcostigma has interxylary phloem, few septated libriform fibres occurring together with normal fibres with bordered pits, and sparse biseriate and low multiseriate wood rays. Even the foliar anatomy and the pollen morphology are different from the other lianous Icacinaceae. Natsiatopsis and Sleumeria may be close to Natsiatum (Kårehed 2001, Byng & al. 2014).

One out of numerous most-parsimonious cladograms of Icacinaceae based on DNA sequence data and morphology (Kårehed 2001). Raphiostylis is usually placed in the Apodytes clade as sister to Apodytes. According to Angulo & al. (2013), Leretia, Icacina and Casimirella form a clade which is more related to other genera than to [Mappia+Nothapodytes].

Phylogeny of Icacinaceae based on Byng & al. (2014).

Cladogram of Icacinaceae based on DNA sequence data (Stull & al. 2015).

ONCOTHECACEAE Kobuski ex Airy Shaw

( Back to Icacinales )

Airy Shaw in Kew Bull. 18: 264. 8 Dec 1965

Oncothecales Doweld, Tent. Syst. Plant. Vasc.: li. 23 Dec 2001

Genera/species 1/2

Distribution New Caledonia.

Fossils Unknown.

Habit Bisexual, evergreen trees or shrubs.

Vegetative anatomy Phellogen ab initio in outer cortex. Vessel elements with scalariform perforation plates; lateral pits scalariform?, simple pits. Imperforate tracheary xylem elements tracheids or fibre tracheids with bordered pits?, non-septate. Wood rays uniseriate and multiseriate, heterocellular. Axial parenchyma apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty. Secondary phloem in young stems stratified. Sieve tube plastids S type. Nodes usually 5:5, pentalacunar with five leaf traces distally united into invaginated arc (sometimes 3:3, trilacunar with three traces). Asterosclereids present. Parenchyma cells with aggregations of calciumoxalate crystals. Druses abundant. Tanniniferous cells abundant.

Trichomes Hairs absent.

Leaves Alternate (spiral), simple, entire, coriaceous, with convolute? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection arcuate. Venation pinnate. Stomata modified paracytic or anisocytic. Cuticular wax crystalloids? Mesophyll with sclerenchymatous idioblasts. Calciumoxalate druses present. Leaf margin serrate or entire, with small caducous glands on teeth.

Inflorescence Axillary, usually panicle or thyrse. Floral prophylls (bracteoles) triangular, two or three per flower.

Flowers Actinomorphic, small. Hypogyny. Sepals five, with imbricate quincuncial aestivation, persistent, more or less free. Petals five, with imbricate aestivation, caducous, connate. Nectary? Disc absent.

Androecium Stamens five, haplostemonous, antesepalous, alternipetalous. Filaments free, adnate to corolla tube (epipetalous). Anthers basifixed, non-versatile, disporangiate (dithecal), extrorse, longicidal (dehiscing by longitudinal slits); connective in Oncotheca balansae (not in O. humboldtiana) acute and incurved above gynoecium. Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains (2–)3-colporate, shed as monads, bicellular? at dispersal. Exine tectate, with columellate infratectum, perforate, smooth.

Gynoecium Pistil composed of five syncarpous and connate antepetalous carpels. Ovary superior, quinquelocular, with five ridges, apically somewhat quinquelobate (with lobes incompletely closed). Stylodia five, free, recurved, conduplicate. Stigmas ventral, punctate, type? Pistillodium absent.

Ovules Placentation apical to axile. Ovules usually two (sometimes one) per carpel, anatropous, pendulous, epitropous, unitegmic, crassinucellar. Funicle long. Integument four to seven cell layers thick. Megagametophyte monosporous, Polygonum type? Endosperm development nuclear? Endosperm haustoria? Embryogenesis?

Fruit A two- to five-seeded drupe with thick quinquelocular pyrene and persistent calyx.

Seeds Aril absent. Testa thin. Exotesta? Endotesta? Perisperm not developed. Endosperm copious. Embryo straight, cylindrical, well differentiated, with chlorophyll? Hypocotyl long. Cotyledons usually two (rarely three), short. Germination?

Cytology n = 25

DNA

Phytochemistry Virtually unknown. Tannins present.

Use Unknown.

Systematics Oncotheca (2; O. balansae, O. humboldtiana; New Caledonia).

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LAMIIDAE Takht. ex Reveal

ICACINALES Tiegh.

ICACINACEAE (Benth.) Miers

ONCOTHECACEAE Kobuski ex Airy Shaw